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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

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

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Fossil remains for the family Gomphotheriidae The main objective of this study is to describe and
found in Mexico were reviewed by ALBERDI & identify the almost complete skeleton of Stegomasto-
CORONA (2005), with particular attention to Stego- don collected in the Basin of Chapala Lake (ALBERDI
mastodon, including the mention of the study spe- et al. 2002 a), and now deposited on the collections
cimen. In North America, there are few records for of the Guadalajara’s Paleontological Museum “Fe-
this genus, generally recording isolated molars (see derico A. Solórzano Barreto”. We also compare
OSBORN 1936). known gomphothere remains from North, Central, and
Stegomastodon is known in North America from South America; review the geographic distribution
the Blancan to the Irvingtonian, and during this last of southern forms throughout the Americas, their
period it migrated to the south of the continent, where possible origin, and the paleoenvironment in which
it is found in localities from middle and late Pleisto- they lived.
cene (BELL et al. 2004; WOODBURNE et al. 2006). It is
the least represented gomphothere in Mexican sedi-
2. The Chapala Locality
ments, and its presence could be related to either
warm or humid conditions, which might have been The Chapala specimen was found in April 2000 by
found in localities as far as Santa Anita, Baja Cali- Mr. JUAN SANTOS in sediments of the north rim of
fornia; Rancho El Ocote, Guanajuato; and La Goleta, Lake Chapala, in the town of Santa Cruz de la Soledad
Michoacán (ALBERDI & CORONA 2005). Other records (20º 17’ 59” N, 103º 09’ 17” W, 1522 mosl), Jalisco
of its presence include: Yepómera (Chihuahua), (Fig. 1). The remains were found at the bottom of the
Ameca (Jalisco), Zacualtipan (Hidalgo), and Valse- lake sediments in very fine sand; they represent a
quillo (Puebla). 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 pisiforms; right and left cuneiforms or pyramidals;
SOLÓRZANO BARRETO, curator of Guadalajara’s right lunate; right and left scaphoids; right and left
Regional Museum, to evaluate the findings and the unciforms; right and left magna; right and left tra-
feasibility to excavate them. Based on SOLÓRZANO’s pezoids; among the metacarpals, there are a possible
report, a team was organized to salvage the skeleton, right MCV, right and left MCIV, right and left MCIII,
including OTTO SCHONDUBE B. as archaeologist; right and left MCII, and right and left MCI; as for the
FEDERICO SOLÓRZANO BARRETO, paleontologist; anterior phalanges: there are the first phalanx for right
CARLOS SANTOS, conservator; ROBERTO PERALTA, fingers IV, III and I, and second phalanx for right
restorer, and JAVIER JUÁREZ W., biologist from the fingers III and II, as well as some other fragments and
Guadalajara’s Paleontological Museum. The remains two deteriorated third phalanges.
were deposited in the collections of Guadalajara’s From the hind limb, there is the almost complete
Paleontological Museum “Federico A. Solórzano pelvis, right femur and distal epiphysis of left femur;
Barreto” (MPG). both patellae; both complete tibiae; complete right
The finding of fossil remains in the region of the fibula and distal portion of left fibula; from the tarsal
Lake Chapala is common, and most are isolated and bones: right and left calcanea; right and left astragali;
disarticulated, whereas the gomphothere remains right and left naviculars; right and left cuboids; right
were surprisingly complete, clearly with anatomical external and internal cuneiforms; from the meta-
association and without traces of carnivore activity or tarsals: right and left MTI, MTII, MTIII, MTIV, and
any movement after deposition. The previous facts, MTV; for the hind phalanges: first phalanges of left
along with the crushed skull, suggest either a cata- fingers II, III and IV, and right fingers I and IV;
strophic death or a fast burial event for the animal. second phalanges for left fingers III and IV. Also,
A sample of molar enamel from the specimen was there are 13 sesamoids, 10 distals, 1 possibly proxi-
sent for dating by 14C AMS to ‘Centrum voor Iso- mal, and 2 fragments; 19 rib pairs; several unidenti-
topen Onderzoek’ (Groningen, The Netherlands), fiable fragments, and fragments that possibly are part
giving a date of 27,910 ± 270 years BP (GrA-23479), of a hyoid bone.
indicating a Late Pleistocene age for the deposit. Methodologically, we selected series of measure-
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ments and characters following those proposed by
TASSY (1983) and BOEUF (1983, 1992) for remains of
3. Material and methods
Anancus and Mammuthus from Chilhac (see ALBERDI
Fossil remains for this group in North America are et al. 2002 b). For the skull and mandible, the charac-
usually limited of few bones and isolated teeth, and it ters used are detailed in ALBERDI et al. (2002b: figs.
is quite rare to have an almost complete specimen 2-3).
such as the one on deposit at MPG. It has a partial The diagnostic elements for analysis of variation
skull with the tusks almost complete, although they are the molars, mainly M2/m2 and M3/m3. For the
were isolated and broken, and there is a broken Chapalan gomphothere there is only a M3 and a m3
M3; the mandible has fragments of both rami, with a that can be compared with the abundant data for
complete left m3 and part of the right m3. From the gomphotheres from South America (ALBERDI et al.
postcranial skeleton there are: atlas, partial axis, 28 2002 b, 2004; PRADO et al. 2002). For the molars, the
pre-sacral vertebrae, 7 caudal vertebrae, and the maximum length, and maximum width at each one
sacrum; the anterior limbs are represented by: left and of lophs were measured. ALBERDI et al. (2004) did
right scapulae, left humerus, distal portion of right several analyses of correlation between the dental
humerus, left and right radii and ulnae (broken ole- characters, showing a high correlation between maxi-
cranon of right ulna); carpal bones: right and left mum length and maximum width of the 4th loph(id)

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.
 The most complete skeleton of Stegomastodon recorded for the Mexican Late Pleistocene 243

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Fig. 3 (Legend see p. 242)

244 M. T. Alberdi et al.

for the last molars. Following those we undertook allocating the species ‘Mastodon’ waringi (= chimborazi,
bivariate diagrams among the maximum lengths and according to FICCARELLI et al.) from Eduador and Brazil
the widths of the 4th loph(id), comparing our study to the genus Haplomastodon, but we consider it a junior
synonym of Stegomastodon (ALBERDI et al. 2002 b, 2004;
material with those from other localities in North, PRADO et al. 2002, 2005).
Central, and South America. In some cases, multi-
variate analyses were performed to explore and con- Diagnosi s : Gomphothere with elephant-like skull, short
trast the differences and similarities between the few and high, less depressed than in Cuvieronius. Mandible
dental remains from Chapala, and those specimens brevirostrine with bunolophodont and trilophodont inter-
mediate teeth (d3, d4, m1, m2). Second molars usually have
known from other Americas localities by a discrimi- a strong lophid that some authors define as a fourth lophid.
nate analysis (DA) to identify specimens not included Moderate alternation of M3/m3 posterior loph(id)s (with a
in the original analysis which established the groups certain angular inclination, pretrite and posttrite cusps
(REYMENT 1991). For the postcranial elements, we opposite, and 5-51/2 lophs, or more). Molars have double
selected measurements that show size differences trefoils, and wear resulting in the presence of several
conules among the lophs, showing a very complicated
among them. occlusal pattern. Tusks slightly curved, upturned or nearly
straight, without enamel; enamel present only in some
4. Systematic paleontology juvenile individuals.

CABRERA (1929) proposed that the families Gompho-

theriidae, Mammutidae, and Elephantidae should be 5. Description of the Chapalan specimen
included within the superfamily Elephantoidea, with
all of the South American proboscideans being The Chapalan specimen is cataloged as MPG-PD-001.
allocated to the family Gomphotheriidae, and to two The skull is flattened and deformed in the dorsal or
subfamilies: Cuvieroniinae and Anancinae. The most external portion; ventrally it is crushed, and retains
recent taxonomic proposal by SHOSHANI & TASSY only a much worn right M3, isolated and partially
(2005) included all gomphotheres in the suprafamily broken. The cranium still keeps the central vertex, but
Elephantida. In this study we follow CABRERA’s otherwise it is flattened and deformed (Fig. 3).
(1929) proposal for including all those in the family The anterior portion of the maxillar region is en-
Gomphotheriidae since they all show the basic
eschweizerbartxxx ingenta

larged due to the skull flattening, and the tusk alveoli

general characters for the family. LAMBERT (1996) and are divergent but not curved. Such divergence is simi-
DUDLEY (1996) studied specimens of the family lar to that found in other Stegomastodon skulls from
Gomphotheriidae from North America, and concluded the Buenos Aires province (PRADO et al. 2002). The
that the members of the subfamily Anancinae are alveolar region at the anterior portion of the pre-
limited to the Old World. maxilla is almost straight in comparison with the
divergent pattern shown by Cuvieronius skulls from
Bolivia. For the Chapalan specimen, similar to skulls
Family Gomphotheriidae CABRERA, 1929 of Stegomastodon from Argentina, Brazil, and Ecua-
dor, the sagittal plane is characterized by a fissure or
Genus Stegomastodon POHLIG, 1912 central depression. The few mensural characters that
could be recorded from this fragmented skull indicate
For a complete synonymy, see CABRERA (1929) and
OSBORN (1936). a large animal, as seen in Table 1. The skull nuchal
portion is missing, apart from the nuchal fossae that is
Ty pe s p e c ie s : Mastodon mirificus LEIDY, 1858. broken, and nasal nares deformed.
Right M3 has four lophs and lacks, possibly, the last
S t r at i g r aphy : Stegomastodon has been recorded from the cusp and the heel; it is very worn and wide, with
Late Hemphillian to the Early Pleistocene in the central and
western regions of North America. In South America, it is
rugose enamel (ptychodonty) with some damage in
known from the Middle and Late Pleistocene. the first loph, covered with abundant cementum over
lophs and filling valleys; medium line is somehow
G e og r a p h ic d ist ri b ut ion : It is found in the central and erased and has multiple internal and central conules
western regions of North America, and in South America it providing a double trefoiled figure with the rugose
is known from Brazil, Ecuador, Argentina, Uruguay, and
enamel; medial and lateral cingula are strong and
Perú; also probably from Venezuela, Colombia, and Para-
guay. FICCARELLI et al. (1993, 1995) followed HOFFSTETTER reach up to the heel (Fig. 3). It has a massive aspect
(1950, 1952) and SIMPSON & PAULA COUTO (1957) in and its measurements are; tooth total length 206 mm
 The most complete skeleton of Stegomastodon recorded for the Mexican Late Pleistocene 245

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).
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(approximately), loph widths; 1st 105 mm, 2nd 105 rugose enamel and a much worn stage is similar to
mm, 3rd 98, 4th 84, and broken heel 40 mm (approxi- the cranial M3 and resembles those of the South
mately). Upper tusks are present, broken at different American gomphotheres. There is a medial line that is
levels, but long, large, straight, and without enamel somehow lost at the level of the first loph, with well
band; their lengths are 152 and 162 cm, and at the base developed mesial and external cingula, central conules
have diameters of 131 x 152 mm, and 153 x 163 mm, closing valleys, which due to the deep wear has
respectively; at the apex, the later tusk has a 140 x been transformed into a complicated trefoil that also
120 mm diameter. is being lost. Measurements are: right m3 length,
There are two mandibular fragments that represent 214 mm (approximately), lophids width 82, 86, 88.7,
both rami (Fig. 3, Table 2). The only measurements and 82 mm, corresponding respectively from the first
and characters of the robust mandible that could be to the fourth lophid, and a heel of 64 mm width. Width
assessed were in Table 2. The two mental foramina on of the distal fragment of left m3 is 81 mm on the
the left ramus are larger the anterior than the posterior, fourth lophid, and 66 mm at the heel.
and the contrary on the right ramus; anterior section of There are some fragments that pertain to the hyoid
the left ramus symphysis is 126 x 94 mm. bone as compared with the records provided by
Lower dental remains are few; on the right ramus SHOSHANI & MARCHANT (2001). Two fragments of the
there is the m3 alveolus, and an isolated right m3 with stylohyoid are preserved and a possible fragment of
four lophids and a large heel. On the left ramus there the thyrohyal bone. We put it in its possible anatomical
is a mark of the obliterated m2 alveolus, and m3 position in Figure 4. Due to the scarce number of
alveolus, and an isolated fragment that has the last two these kinds of bones known in the Gomphotheriidae
lophids of a left m3, both lophids showing much wear family we think it is interested to communicate their
(Fig. 3). The morphology of all those teeth, with 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 247

as new materials pertaining to hyoid bones become trapezoids, with the second one complete but broken
available. into two fragments, and dimensions of 52 x 88x
Measurements of the postcranial remains are pro- 33 mm.
vided in the text. From the vertebral column, there is a Among the metacarpals (MC), there are both right
complete, large-sized atlas, on which the facets for and left MCIV, with an almost complete left one with
the occipital condyles are 132.5 x 80 mm; maximum maximum length of 152 mm, a minimal width of
diameter of neural canal is 107 mm. The axis is 72 mm, proximal end 67 x 96.5 mm, and the maximum
incomplete and the other vertebrae are mentioned width at the distal end is 76 mm, right one measures
previously (see material section). 158/72/74 – x/87, respectively; both right and left
The left scapula is almost complete, with the MCIII, with the left maximum length of 174 mm,
glenoid cavity that articulates with the humerus head diaphysis minimal width 77 mm, proximal articu-
having a maximum diameter of approximately 200 lation of 94 x 99 mm, and maximum width at the distal
mm, conserving the cranial portion and the spinous end of 86.5 mm; right and left MCII, left one with the
process, but lacking the caudal angle portion. There maximum length of 152 mm, diaphysis minimal width
are isolated fragments of the right scapula. 79.6 mm, proximal articulation 80 x 92 mm, and maxi-
The available long bones of the front limb are the mal width of distal articulation 86.5 mm; right and left
left humerus, and the distal portion of the right one, MCI, left one with the maximum length of 129 mm,
the right and left radii, an incomplete right ulna while diaphysis minimal width 79 mm, proximal articu-
the left is complete. Left humerus has a maximum lation 79 x 96 mm, and maximal width of distal
length of 92 cm, minimum diameter of the diaphysis articulation 93.5 mm.
is 128 mm, diaphysis diameter at the level of the The pelvic bone is almost complete; right and left
maximum extension of the deltoid crest is 208 mm, ischia have the suture area and partially the ilium.
articular transverse diameter of the distal portion is Pelvis has very altered edges and acetabulum has a
250 mm, distance from the highest point of the deltoid maximum diameter of 198 mm.
crest to the proximal end is 390 mm, the distance from Long bones of hind leg are right femur and the
the highest point of the epicondyle crest to the distal distal articulation of left one; right and left patella, and
end is 383 mm, and maximum length of epicondyle eschweizerbartxxx ingenta
both complete tibia. Right femur has a maximum
crest is 265 mm. Left radius is 67.5 cm long. Right length of 113 cm; a maximum width at the proximal
ulna is 80 cm long, and left one (without olecranon) end of 386 mm, and a maximum distal width at
is 67 cm; diaphysis transverse diameter is 90 mm, the distal lateral end of 246 mm. Left tibia has a maxi-
sigmoid notch of the maximum transverse diameter is mum length of 68 cm, the proximal articulation has
260 mm, distal articulation transverse diameter is articular widths of approximately 260 x 168 mm, and
175 mm. the distal end 195 x 150 mm. It has both patellae, of
Of the carpus, there are both pisiforms, with which the right one is almost complete, with articular
tuberosity lengths of 139 mm right, and 135 mm left, facets of 130 x 140 mm, and a height of 120 mm.
articulation facets of 64 x52.3, and 63 x 58 mm, Of the tarsal bones, there are both right and left
respectively. Right and left cuneiform or pyramidal calcanea, with dimensions, respectively: maximum
bones have maximum dimensions of 187 x 123 x61, length 228 and 220 mm; minimum length 136.5 and
and 118x133x 51 (length, depth, height); right semi- 117.5 mm; maximum width of facets, 195 and 189
lunar, 140x148 x82 mm; both right and left scaphoids mm; right and left astragali, maximum length 170 and
with measurements of 61 x46 x62.5, and 60 x 42 x 171.6 mm, antero-posterior maximum depth 148.3
59.2 mm; right and left unciforms, the left one much and 153.5 mm; maximum height 105 and 83.5 mm.
better preserved with maximum dimensions of Naviculars, right and left, with maximum dimensions
110x 141x 110 mm; right and left magna, with right of 149 x 105 x 44 and 147.5 x 105 x 42 mm, respective-
measurements of 102 x140 x112 mm; right and left 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.
Fig. 5. Left metatarsals I to V from the Stegomastodon Of metatarsals (MT), there are right and left MTI,
skeleton from Chapala, right (up) and left (down). A –
the right one with a maximum length of 58 mm, dia-
Anterior view; B – latero-external view.
248 M. T. Alberdi et al.

physis minimum width 36.7; a proximal end of 37 x fused vertebrae, and holes in most of the bones
42 mm, and the maximum width of the distal end possible due to the action of boring insects. Morpho-
is broken; the left one has a maximum length of logical characters for skull and mandible, as well as
> 60 mm, diaphysis minimal width 30; a proximal end the few dental remains with the complex morphology,
of 33x35 mm, and the maximum width of the distal and also the straight tusks lacking an enamel band,
end is broken. Right and left MTII, the right one with suggest the presence of a gomphothere pertaining
a maximum length of 110 mm, diaphysis minimal to the genus Stegomastodon as stated elsewhere
width 60.3; a proximal end of 85 x78 mm, and the (ALBERDI et al. 2002a). Among the South America
maximum width of the distal end is 79 mm; the left Stegomastodon remains is present many times nearly
one with a maximum length at 110 mm, diaphysis straight tusks (FRASSINETTI & ALBERDI 2005).
minimal width, 59; a proximal end of 82 x75 mm, and The skull of the Chapalan specimen, although
maximum width of the distal end is 76 mm. Right and crushed, shows some morphological features similar
left MTIII, the right one with a maximum length at to Stegomastodon and distinct from Cuvieronius,
114 mm, diaphysis minimal width, 66; a proximal end mostly in the morphology of the anterior portion of
of 65.2x74 mm, and the maximum width of the distal the maxillar symphysis, where the tusk alveoli are
end is 77 mm; the left one as a maximum length at slightly divergent, but not tilted outwards as found in
114.3 mm, diaphysis minimal width, 68; a proximal the large sample of Cuvieronius skulls from Tarija
end of 68x 68 mm, and maximum width of the distal (Bolivia), and in the other species of Cuvieronius
end is 76 mm. Right and left MTIV, the right one with illustrated by OSBORN (1936) for North America. The
a maximum length at 97 mm, diaphysis minimal alveolar divergence in the Chapalan skull is likely
width, 54.2; a proximal end of 53 x68 mm, and the more due to the crushing. Its measurements are
maximum width of the distal end is 59 mm; the left similar to the Stegomastodon skulls from the Buenos
one with a maximum length at 96.3 mm, diaphysis Aires province (Argentina), Pains (Brazil), and La
minimal width, 56.6; a proximal end of 53 x 67 mm, Huaca (Perú) as shown in Table 1. Those observed
and maximum width of the distal end is 60.5 mm. differences are within the range of variability in
Right and left MTV, the right one with a maximum the group, and also may be attributed to the crushed
length at 69 mm, diaphysis minimal width, 67; a eschweizerbartxxx ingenta
condition of the skull (ALBERDI et al. 2002 b, 2004;
proximal end of 63 x68 mm, and the maximum width PRADO et al. 2002).
of the distal end is 74 mm; the left one as a maximum Mandibular remains also allow comparison with
length at 64 mm, diaphysis minimal width, 64; a those studied from South America, and yield a data-
proximal end of 57 x71 mm, and maximum width of base with metric and morphological data (Table 2). As
the distal end is 74 mm. Left hind foot is the most shown, the specimen is very similar to the mandibular
nearly complete (Fig. 5). remains of Stegomastodon from South America,
Among the phalanges, there are the first front although the available data points for the Chapalan
phalanges of the right fingers III, II, and I, and those specimen are few.
of the left IV, I, and possibly V; also there are 2 Third molars allow more detailed comparisons with
damaged third phalanges. Among the hind phalanges, remains having M3/m3 from South America that are
there are the first ones for left fingers II and III, and fairly available. Bivariate analyses for molar metric
for right fingers I and II; second phalanges for left characters for the complete sample available from
II and III, and right toes II. Their measurements are South America, show a high correlation between the
provided in Table 3 in mm. Also, there are available tooth lengths and the widths of the fourth loph(id) of
10 distal sesamoids, and possibly one proximal, with the same teeth (ALBERDI et al. 2004).
metrical data provided in Table 4. Bivariate analyses including the Chapala sample
Based on the apendicular skeleton, it can be in- with the South American Gomphotheriidae (ALBERDI
ferred that the height at the shoulder blade for this et al. 2002b, 2004; PRADO et al. 2002), showed that
animal was around 2.5-3 m (Fig. 6). 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
6. Discussion
associated with the extreme wear of the tooth; in the
The skeleton is well-preserved and pertained to an old diagram length/width for the fourth loph, the study
adult individual of great size, showing some exostosis, specimen lies among the largest for the proposed
 The most complete skeleton of Stegomastodon recorded for the Mexican Late Pleistocene 249

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 cision. OSBORN (1936) included eight species in the
locates both molars among those of the largest size genus Stegomastodon distributed from Nebraska
from South America, where there are one the best down to Texas. Among those, S. primitivus, from the
population of this group of gomphotheres (Fig. 8). The Early Pleistocene of Nebraska, although quite primi-
morphological characters, including those for the tive, shows tusk morphology and molars quite similar
skull and those for the molars, all point to the to the study gomphothere. Those characters are
relationship to the genus Stegomastodon, rather mainly found in the number of loph(id)s at the third
than to Cuvieronius. Furthermore, the large size and molar and the rugose enamel morphology of the same
straightness of the tusks, and the lack of an enamel tooth, which eventually provided the basis for a very
band, also suggests that the specimen can not be complex wear pattern on the studied molar. Also the
assigned to Cuvieronius. All those characters, and tusks are quite straight, and lack the enamel band,
the brevirostrine mandible and lack of lower tusks, and the anterior portion of the skull, where the tusk
separates the study specimen from the other two alveoli are located is slightly divergent. All of those
gomphothere genera known from México, Rhyncho- characters are also similar to those of Stegomastodon
therium, and Gomphotherium. remains from South America. Similar primitive con-
For the specific identification, our data were com- ditions were stated by WOODBURNE (1961) in de-
pared with those provided by OSBORN (1936) for scribing Stegomastodon rexroadensis for Pliocene
North America, but we have not reached a final de- deposits in Kansas.
250 M. T. Alberdi et al.

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
eschweizerbartxxx ingenta
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-
Fig. 7. Scatter diagrams of Stegomastodon M3 /m3 from stodon may have been a more specialized grazer than
Chapala, including the data for the third molars for
gomphotheres known from South American populations. other gomphotheres, they certainly have been re-
Top: m3; bottom: M3. Cuvieronius is present in Perú, Chile corded in some localities co-occurring with mam-
and Bolivia; Stegomastodon in Perú, Brazil, Uruguay, and moths. It seems that Stegomastodon mostly occurred
Argentina, in Chile it is only in Taguatagua. 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
As for the remaining seven species cited by of the Pleistocene (ARROYO-CABRALES et al. 2007;
OSBORN (1936) for North America: S. chapmani, S. MEAD et al. 2006).
mirificus (type species), S. successor, S. texanus, S. We think that the specimen from Chapala could be
arizonae, S. aftoniae and S. priestleyi, following representing a primitive form with the most recent
 The most complete skeleton of Stegomastodon recorded for the Mexican Late Pleistocene 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
eschweizerbartxxx ingenta
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.

References
ALBERDI, M. T., CARTELLE, C. & PRADO, J. L. (2002 b): El
registro de Stegomastodon (Mammalia, Gompho-
Fig. 8. Principal component analysis of the third molars theriidae) en el Pleistoceno superior de Brasil. – Revista
from South American population and the Chapalan Española de Paleontología, 17 (2): 217-235.
specimen. Top: m3; bottom: M3. ALBERDI, M. T. & CORONA-M., E. (2005): Revisión de los
gonfoterios en el Cenozoico tardío de México. – Revista
Mexicana de Ciencias Geológicas, 22 (2): 246-260.
ALBERDI, M. T., POLACO, O. J., JUÁREZ WOO, J. & ARROYO-
occurrence of the genus. It deserves further discussion CABRALES, J. (2002a): Presencia de Stegomastodon
(Mammalia: Gomphotheriidae) en el Pleistoceno final
in the future.
de Jalisco, México. – VIII Congreso Nacional de
As a separate note, we want to state that we com- Paleontología, Noviembre de 2002, Guadalajara, Ponen-
pletely disagree with WEBB (1992) and WEBB & cias: 22.
RANCY (1996) in regard to the use of Notiomastodon ALBERDI, M. T., PRADO, J. L. & SALAS, R. (2004): The
instead of Stegomastodon for the brevirostrine Pleistocene Gomphotheriidae (Proboscidea) from Peru.
gomphotheres with more than five lophids at M3 from – Neues Jahrbuch für Geologie und Paläontologie,
Abhandlungen, 231: 423-452.
South America since Notiomastodon has a doubtful ARROYO-CABRALES, J., POLACO, O. J., LAURITO, C., JOHNSON,
status as discussed in depth by SIMPSON & PAULA E., ALBERDI, M. T. & VALERIO ZAMORA, A. L. (2007):
COUTO (1957), PAULA COUTO (1979), and PRADO et The proboscideans (Mammalia) from Mesoamerica. –
al. (2005). Quaternary Internacional, 169-170: 17-23.
252 M. T. Alberdi et al.

BELL, C. J., LUNDELIUS Jr., E. L., BARNOSKY, A. D., LAMBERT, W. D. & SHOSHANI, J. (1996): Proboscidea. – In:
GRAHAM, R. W., LINDSAY, E. H., RUEZ Jr., D. R., JANIS, C. M., SCOTT, K. M. & JACONS, L. L. (Eds.):
SEMKEN Jr., H. A., WEBB, S. A. & ZAKRZEWSKI, R. J. Evolution of Tertiary Mammals of North America: I.
(2004): The Blancan, Irvingtonian, and Rancholabrean Terrestrial, Carnivores, Ungulates and Ungulate-like
Mammal Ages. – In: WOODBURNE, M. O. (Ed.): Late Mammals, 606-621; Cambridge (Cambridge University
Cretaceous and Cenozoic Mammals of North America. Press).
Biostratigraphy and Geochronology, 232-314; New York LEIDY, J. (1858): Notice of remains of extinct Vertebrata,
(Columbia University Press). from the valley of the Niobrara River, collected during
BOEUF, O. (1983): Le site Villafranchien de Chilhac (Haute- the exploring expedition of 1857, in Nebraska under the
Loire) France. Etude Paléontologique et Biochrono- command of Lieut. G. K. Warren, U. S. Top Eng., by Dr.
logique. – Thèse de Doctorat d’État, Université de Paris F. V. Hayden, geologist of the expedition. – Proceedings
VII. – 253 pp. of the Academy of Natural Sciences of Philadelphia, 10:
– (1992): Anancus arvernensis chilhiacensis nov. sub- 20-29.
sp. (Proboscidea, Mammalia), un Mastodonte du Plio- LUCAS, S. G. (2003): Pleistocene Proboscideans from Lago
Pléistocène de Haute-Loire, France. – Géobios, 14: de Chapala, Jalisco, Mexico. – Current Research in the
179-188. Pleistocene, 20: 105-107.
CABRERA, A. (1929): Una revisión de los Mastodontes MEAD, J. I., BAEZ, A., SWIFT, S. L., CARPENTER, M. C.,
Argentinos. – Revista del Museo de La Plata 32: 61-144. HOLLENSHEAD, M., CZAPLEWSKI, N. J., STEADMAN, D.
CORREAL URREGO, G. & HAMMEN, T. VAN DE (2003): W., JORDON, B. & ARROYO-CABRALES, J. (2006): Tro-
Supervivencia de Matodontes, Megaterios y presencia pical marsh and savanna of the Late Pleistocene in
del hombre en el Valle del Magdalena (Colombia) entre northeastern Sonora, Mexico. – The Southwestern
6000 y 5000 AP. – Revista de la Academia Colombiana Naturalist, 51: 226-239.
de Ciencias Exactas, Físicas y Naturales, 27 (103): OSBORN, H. F. (1936): Proboscidea. A monograph of the
159-164. discovery, evolution, migration and extinction of the
DUDLEY, J. P. (1996): Mammoths, gomphotheres, and the mastodons and elephants. Vol. 1 – Moeritherioidea,
Great American Faunal Interchange. – In: SHOSHANI, J. Deinotheiroidea, Mastodontoidea. – Bulletins of the
& TASSY, P. (Eds.): The Proboscidea. Evolution and American Museum of Natural History, 1: 1-802.
Palaeoecology of Elephants and their relatives, 289-295; PAULA COUTO, C. (1979): Tratado de Paleomastozoologia. –
Oxford (Oxford University Press). 590 pp.; Rio de Janeiro (Academia Brasileira de
FICCARELLI, G., BORSELLI, V., HERRERA, G., MORENO ESPI- Ciências).
NOSA, G. & TORRE, D. (1995): Taxonomic remarks on POHLIG, H. (1912): Sur une vieille mandibule de “Tetra-
the South American mastodons referred to and Cuviero- eschweizerbartxxx ingenta
caulodon ohioticum” BLUM., avec défense in situ. –
nius. – Géobios, 28: 745-756. Bulletin de la Société Belge de Géologie, 26: 187-193.
FICCARELLI, G., BORSELLI, V., MORENO ESPINOSA, M. & PRADO, J. L., ALBERDI, M. T., AZANZA, B., SÁNCHEZ, B. &
TORRE, D. (1993): New Haplomastodon finds from the FRASSINETTI, D. (2005): The Pleistocene Gompho-
Late Pleistocene of northern Ecuador. – Géobios, 26: theriidae (Probsocidea) from South America. – Quater-
231-240. nary International, 126/128: 21-30.
FICCARELLI, G., COLTORTI, M., MORENO-ESPINOSA, M., PRADO, J. L., ALBERDI, M. T. & GÓMEZ, G. N. (2002): Late
PIERUCCINI, P. L., ROOK, L. & TORRE, D. (2003): A Pleistocene gomphothere (Proboscidea) remains from
model for the Holocene extinction of the mammal the Arroyo Tapalqué locality (Buenos Aires, Argentina)
megafauna in Ecuador. – Journal of South American and their taxonomic and biogeographic implications. –
Earth Sciences, 15: 835-845. Neues Jahrbuch für Geologie und Paläontologie,
FRASSINETTI, D. & ALBERDI, M. T. (2005): Presencia del Abhandlungen, 225: 275-296.
género Stegomastodon entre los restos fósiles de masto- REYMENT, R. A. (1991): Multidimensional paleobiology. –
dontes de Chile (Gomphotheriidae), Pleistoceno 416 pp.; New York (Pergamon Press).
Superior. – Estudios Geológicos, 61: 101-107. SENDERS, W. J., KAPPELMAN, J. & RASMUSSEN, D. T. (2004):
HOFFSTETTER, R. (1950): Observaciones sobre los masto- New large-bodied mammals from the late Oligocene site
dontes de Sud América y especialmente del Ecuador. of Chilga, Ethiopia. – Acta Palaeontologica Polonica, 49
Haplomastodon, subgen. nov. de Stegomastodon. – (3): 365-392.
Publicaciones de la Escuela Politécnica Nacional, 1: SHOSHANI, J. & MARCHANT, G. H. (2001): Hyoid apparatus:
1-51. a little known complex of bones and its contribution to
– (1952): Les mammifères Pléistocènes de la République proboscidean evolution. – In: CAVARRETTA, G., GIOIA, P.,
de l’Equateur. – Mémoires de la Société géologique de MUSSI, M. & PALOMBO, M. R. (Eds.): La Terra degli
France, 66: 1-391. Elefanti. Atti del 1º Congresso Internazionale. Roma,
KURTÉN, B. & ANDERSON, E. (1980): Pleistocene Mammals 16-20 Ottobre 2001, 668-675; Roma (Consiglio Nazio-
of North America. – 443 pp.; New York (Columbia nale delle Ricerche).
University Press). SHOSHANI, J. & TASSY, P. (2005): Advances in probosci-
LAMBERT, W. D. (1996): The biogeography of the gompho- deans taxonomy & classification, anatomy & physio-
theriid proboscideans of North America. – In: SHA- logy, and ecology & behaviour. – Quaternary Inter-
SHANI, J. & TASSY, P. (Eds.): The Proboscidea. Evolution national, 126-128: 5-20.
and Palaeoecology of Elephants and their relatives,
143-148; Oxford (Oxford University Press).
 The most complete skeleton of Stegomastodon recorded for the Mexican Late Pleistocene 253

SIMPSON, G. G. (1945): The principles of classification and WOODBURNE, M. O., CIONE, A. L. & TONNI, E. P. (2006):
a classification of mammals. – Bulletin of the American Central American provincialism and the Great American
Museum of Natural History, 85: 1-350. Biotic Interchange. – In: CARRANZA-CASTANEDA, O. &
SIMPSON, G. G. & PAULA COUTO, C. (1957): The masto- LINDSAY, E. H. (Eds.): Advances in late Tertiary verte-
donts of Brazil. – Bulletin of the American Museum of brate paleontology in Mexico and the Great American
Natural History, 112: 125-190. Biotic Interchange. – Universidad Nacional Autónoma
TASSY, P. (1983): Les Elephantoidea miocènes du Plateau de México, Instituto de Geología y Centro de Geo-
du Potwar, Groupe de Siwalik, Pakistan. – Annales de ciencias, Publicación Especial, 4: 73-101.
Paléontologie, 69: 99-136, 235-297, 317-354.
WEBB, S. D. (1992): A brief history of New World Pro- Manuscript received: June 30th, 2008.
boscidea with emphasis on their adaptations and inter- Revised version accepted by the Stuttgart editor: September
actions with man. – In: FOX, J. W., SMITH, C. B. & 2nd, 2008.
WILKINS, K. T. (Eds.): Proboscidean and Paleoindian
Interactions, 15-34; Baylor, Texas (Baylor University Addresses of the authors:
Press).
WEBB, S. D. & RANCY, A. (1996): Late Cenozoic evolution MARÍA TERESA ALBERDI, Departamento de Paleobiología,
of the Neotropical mammal fauna. – In: JACKSON, J. B. Museo Nacional de Ciencias Naturales, CSIC, José
C, BUDD, A. F. & COATES, A. G. (Eds.): Evolution and Gutiérrez Abascal, 2, 28006 Madrid, Spain;
environment in tropical America, 335-358; Chicago e-mail: [email protected]
(University of Chicago Press). JAVIER JUÁREZ-WOO, Museo de Paleontología “Federico A.
WOODBURNE, M. D. (1961): Upper Pliocene geology and Solórzano Barreto”, Av. Dr. R. Michel 520, Col. San Carlos,
vertebrate paleontology of part of the Meade Basin, 44100 Guadalajara, Jalisco, Mexico;
Kansas. – Papers of the Michigan Academy of Science, e-mail: [email protected]
Arts, and Letters, 46: 61-101. 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: [email protected],
[email protected]

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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.

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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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