Hodgson Et Al. (2008) - Mastodon and Mammuthus Osteology PDF

Mastodon Paleobiology,
Taphonomy, and
Paleoenvironment in the Late
Pleistocene of New York State:
Studies on the Hyde Park,
Chemung, and North Java Sites
Edited by
Warren D. Allmon and Peter L. Nester
Palaeontographica Americana
Number 61, July 2008
301
COMPARATIVE OSTEOLOGY OF LATE PLEISTOCENE MAMMOTH AND MASTODON
REMAINS FROM THE WATKINS GLEN SITE, CHEMUNG COUNTY, NEW YORK
Jennifer A. Hodgson
Paleontological Research Institution, 1259 Trumansburg Road, Ithaca, New York 14850, U. S. A. Current
address: Department of Anthropology, City University of New York, 365 Fifth Avenue, New York, New York
10016-4309 U. S. A., email hodgson@nycep.org.
Warren D. Allmon
Paleontological Research Institution, 1259 Trumansburg Road, Ithaca, New York 14850, U. S. A., and
Department of Earth and Atmospheric Sciences, Cornell University, Ithaca, New York 14853, U.S.A.
Peter L. Nester
Department of Earth and Atmospheric Sciences, Cornell University, Ithaca, New York 14853, U. S. A. Current
address: Chevron Energy Technology Company, 1500 Louisiana Street, Houston, Texas 77002, U. S. A.
James M. Sherpa
Paleontological Research Institution, 1259 Trumansburg Road, Ithaca, New York 14850, U. S. A. Current
address: 302 Giles Street, Ithaca, New York 14850, U. S. A.
and John J. Chiment
Department of Earth and Atmospheric Sciences, Cornell University, Ithaca, New York 14853, U. S. A. Current
address: 5981 Route 228, Trumansburg, New York 14886, U. S. A.
ABSTRACT
The Watkins Glen site, at which partial skeletons of one mastodon [Mammut americanum (Kerr, 1792)] and one mammoth (likely Mammuthus
primigenius Blumenbach, 1799) were found, presents an opportunity for comparison of the osteology of these two genera. It is often quite
dicult to distinguish specic bones between these taxa in the absence of teeth. The descriptions of postcranial morphology here add to our
understanding of characteristics of the two taxa. In addition, this site presents evidence for Mammut and Mammuthus possibly sharing the same
habitat. We have not determined if the two animals represent actual cohabitation at the site, or were separated in time. Co-occurrence is not
well documented at other sites; the close association of the two taxa here raises issues of ecological interaction between these species in the Late
Pleistocene.
INTRODUCTION
The Watkins Glen site (a.k.a. Chemung, Cornell, or Gilbert),
at which remains of one mastodon and one mammoth
were found, presents an opportunity for comparison of the
osteology of these two taxa (Pls 1-2; Appendix 1).
It is of great interest that this site presents evidence of
Mammut and Mammuthus in such close association. Although
the geographic ranges of these two genera were partly
separate, they did overlap, most extensively in the Mississippi
River drainage basin (Shoshani & Tassy, 1996b), and New
York State was within the bounds of this overlapping area.
Chapter 17, in Mastodon Paleobiology, Taphonomy, and Paleoenvironment in the Late Pleistocene of New York State: Studies on the Hyde
Park, Chemung, and North Java Sites, edited by Warren D. Allmon and
Peter L. Nester, Palaeontographica Americana, 2008, (61): 301-367.
Bone collagen from the Watkins Glen mastodon specimen

has been radiometrically dated to 10,780 60 14C yr BP
(conventional 10,840 60 yr BP; Beta Analytic 176930), and
the mammoth specimen has been dated to 10,820 50 14C yr
BP (conventional 10,840 50 yr BP; Beta Analytic 176929).
The margins of error of these dates overlap, suggesting that
the species very likely coexisted at the site, or at least were not
separated by more than ca. 150 years. If the site does in fact
represent actual sympatry of the two species, which is not well
documented at other sites, this would raise important issues
about changing habitats and potential ecological competition
between these species in the Late Pleistocene.
Based on known distributions of Mammuthus spp. through
space and time, it is likely that the mammoth remains at the
Watkins Glen site (PRI 8830, formerly CU 42440 in part)
302
Palaeontographica Americana, No.
represent M. primigenius Blumenbach, 1799, the woolly

mammoth. Alternatively, they could represent M. columbi
(Falconer, 1857) [= M. jeersonii (Osborn, 1922)]. Mammuthus
columbi was a larger, more southern Late Pleistocene species
mainly occupying the midwestern United States (Graham,
2001). Although the ranges of these two mammoth species
did overlap in the New York State area, they are believed to
have had dierent habitat preferences (Agenbroad, 1984):
M. primigenius preferred a forest/woodland or tundra
environment, whereas M. columbi preferred a steppe/savanna/
parkland environment (Graham, 2001: 707). It is dicult to
distinguish the species in the absence of teeth, because their
postcranial anatomy is almost identical (Haynes, 1991).
The more complete mastodon skeleton from the Watkins
Glen site (PRI 8829, formerly CU 42440 in part) has been
identied positively as Mammut americanum (Kerr, 1792)
based on the dentition (Pl. 3). We are quite certain that
only two animals are represented at the site. The minimum
number of individuals for each species is one. Additionally, all
elements from each animal either articulate with one another
or appear to be from the same individual.
The geographic and stratigraphic ranges of Mammut
and Mammuthus did overlap, but at well-stratied sites, the
species almost never occur together (Graham, 2001). The two
animals are generally found in separate habitats, but could
have potentially become competitors due to degradation
of their environmental niches in the Late Pleistocene. The
environment at the Watkins Glen site was a shallow pond
surrounded by spruce-dominated forest (Griggs & Kromer,
2008). The site occurs at a period during which spruce forest
was gradually shifting toward pine-dominated forest (Griggs
& Kromer, 2008). It has been suggested, among other reasons,
that megafaunal extinctions in this area are linked to this
environmental change (King & Saunders, 1984; Robinson et
al., 2005). Spruce is thought to have been a preferred food of
mastodons and has been found associated with many other
mastodon sites (Miller, 2008). It is evident that spruce was
being consumed at the Watkins Glen site from the abundance
of possibly masticated spruce twigs found (Griggs & Kromer,
2008). It is therefore interesting that we nd the mammoth
at this site, where the environment appears similar to other
mastodon sites. It is possible that these animals were exploiting
separate resources at this same site. It is also possible that
the mammoth was simply migrating through. Hoppe et al.
(1999) described the migratory behavior of mammoths
using strontium isotope ratios from tooth enamel, which we
have not yet attempted on our specimen (no tooth enamel
was recovered for our mammoth specimen, and it is unclear
if this test would be successful on other bone material). A
cohabitation at the site might also provide evidence for
interspecic competition for dwindling resources during this
period leading up to the extinction of both species.
GENERAL DIFFERENCES BETWEEN TAXA

Proboscidean species in North America have generally been
described and taxonomically placed on the basis of their dental
characteristics. Henry Faireld Osborns Proboscidea (1936,
1942), which has long been treated as authoritative on the
subject, has descriptions on dentition. Subsequent literature
has continued this trend, often excluding detailed analyses of
postcranial morphology. Morphological dierences in teeth
are useful for identifying fossil species, because teeth are most
often preserved as fossils. The researcher encounters diculty,
however, if no teeth are present. Stanley J. Olsen (1972) has
compiled the most complete documentation of dierences
between mammoth and mastodon postcranial morphology.
His work remains one of the few reference tools for researchers
attempting to distinguish proboscidean species in the absence
of teeth, and has been extensively consulted in identication of
remains from the Watkins Glen site. This paper veries many
of Olsens observations and notes additional morphological
dierences observed between our specimens.
Dentition will be discussed only briey here (for more indepth descriptions, see Osborn, 1936, 1942; Maglio, 1973).
The tooth morphologies of both Mammut and Mammuthus
are quite distinct, making fossil nds with teeth immediately
identiable. These dierences are clearly related to the diering
diets and mode of chewing of the two genera. Mastodons
have retained the more primitive gomphotheriid tooth form
characterized by high crests of lophs on the occlusal surfaces
of the cheek teeth (Olsen, 1972: 3). This tooth form is
adapted for crushing twigs, leaves, and stems. Other evidence
also points toward a browsing adaptation in Mammut (see
Haynes, 1991). The cheek teeth of Mammuthus, on the
other hand, are more similar to modern elephants and are
characterized by a relatively at occlusal surface with parallel
ridges and plates (Olsen, 1972). This tooth morphology is
a highly specialized adaptation to grinding an abrasive diet
of grass (Haynes, 1991). Trends over time toward increasing
specialization for this adaptation have led to changes in skull
shape. The skull of Mammuthus is high and domed compared
with that of Mammut, which is a result of a shifting of the
skulls center of gravity related to this mode of mastication
(see Maglio, 1973).
Mammoths and mastodons are not taxonomically close
(Text-g. 1; in fact, their last common ancestor lived ca.
20 million years ago; Saunders, 1996), but because they
have similar body bulk, mode of walking, and method of
food gathering they logically have more similarities than
dierences (Olsen, 1972: 5). The overall body proportions
of Mammut and Mammuthus were somewhat dierent. In
general, the limbs of Mammut were shorter and thicker than
303
HODGSON ET AL.: COMPARATIVE OSTEOLOGY OF MAMMOTH AND MASTODON
Table 1. Number of ribs and vertebrae for each species (modied

from Haynes, 1991). C, cervical; T, thoracic, L, lumbar; S, sacral;
c, caudal.
Text-g. 1. Phyletic diagram illustrating relationships among families of the order Proboscidea (after Haynes, 1991; Maglio 1979).
those of Mammuthus, whereas those of Mammuthus were

longer and relatively more slender (Olsen, 1972). Mammut
was a more robust, heavyset animal; compared to modern
Elephas, it had a deeper chest, broader pelvis, shorter legs,
and longer back (Kurtn & Anderson, 1980: 345). Haynes
(1991) has suggested that this greater length and breadth of
the body are related to an increased gut capacity, which could
be a response to the greater quantity of food needed due to the
animals low quality diet of browse. The head of Mammut was
oriented similarly to Loxodonta, more horizontal in relation to
the body (Hayes, 1980) rather than domed as in Mammuthus.
The body prole of Mammuthus was more similar to modern
Asian elephants, with the shoulder as the highest point of the
prole. The longest vertebral spines were positioned at the
front of the shoulder and are sharply slanted back. This created
a hump at the shoulders and downward sloping back (Haynes,
1991). The prole of Mammut also had its highest point at
the front of the shoulders, but a more gradual rearward tilt
(Haynes, 1991).
POSTCRANIAL DESCRIPTIONS
VERTEBRAL COLUMN
The general outline of the vertebral column is quite dierent
in the two species. The prole of the vertebral column of
Mammuthus is humped and downward sloping. The longest
vertebral spines are positioned at the front of the shoulder and
slant sharply back. The length of the spines decreases abruptly
toward the rear of the back, creating a slope, which is more
exaggerated by the doming of the cranium. This is similar to
modern Asian elephants, but Mammuthus displays an even
greater degree of sloping (Osborn, 1942: 1229; Haynes,
1991).
The prole of the vertebral column of Mammut also has
its highest point at the front of the shoulder, but the spines
decrease in length more gradually, and increase again slightly
toward the rear, giving the back a dished prole. This is most
Ribs
Mastodon
18-20
18-20
4-5
27?
Mammoth
18-20
18-20
3-5
3-5
21-22?
similar to the prole of Loxodonta, but with slightly less

dishing (Haynes, 1991).
The number of ribs and vertebrae for these two species has
not been rmly documented in the literature, and it is likely
that there is variation among individuals (Haynes, 1991).
Table 1 gives the approximate ranges for these counts in several
skeletons; see Haynes (1991: 39) for a more complete list of
rib and vertebra counts for other specimens. For Mammut
there are usually 20 thoracics, whereas Mammuthus generally
has 19. A single individual with 18 thoracic vertebrae has been
reported in each taxon (see Haynes, 1991: 39). The reduction
of vertebral number in Mammuthus is a derived character
also present in modern elephants (Haynes, 1991; Shoshani,
1996).
In an attempt to determine how many ribs and vertebrae
were originally present in our specimens, we counted and
checked the articulations of these elements. Based on this
analysis, we conclude that the vertebral column of the Watkins
Glen mastodon originally consisted of 7 cervical vertebrae,
20 thoracics, 3 lumbar, and 5 sacral. Also recovered from
the site were 15 additional thoracic vertebrae belonging to
the mammoth, beginning with the fourth. Based on the rib
sequence, there appear to have been at least 19 ribs originally,
and thus, 19 thoracic vertebrae. Three caudal vertebrae are
present, but we are unable to associate them with one or the
other species, because these bones are not diagnostic.
Cervical Vertebrae
The rst ve cervical vertebrae, which articulate with one
another, as well as the seventh are present for our mastodon
specimen. There is then a gap in the vertebral sequence until
the third thoracic. Although the cervicals do not articulate with
the rest of the vertebral column, and the cranial articulation
is missing, their morphology appears diagnostically mastodon
as described below.
Atlas.When viewed from the posterior aspect (Pl. 4), the
shape of the dorsal margin appears rounded, where it would
be attened in a mammoth (Olsen, 1972). The vertebral
foramen also appears more oval in shape, whereas a mammoth
would show lateral constriction in this area.
304
Axis.The axis is dicult to distinguish between taxa. We have

assigned the axis present in the assemblage to our mastodon
specimen based on its articulation with the atlas and third
cervical vertebra, which we can identify as mastodon. Olsen
(1972: 9) described both animals as having high, pronounced
neural spines that are quite variable in outline when seen from
a lateral aspect. A notch or trough, running in an anteroposterior direction along the crest of the spine, may be preset
or absent in this vertebra of either animal. Our specimen
exhibits an acutely sloping neural spine and a well-dened
trough along the crest (Pl. 5).
Third Cervical.The third through the fth and also the
seventh cervical vertebrae are present for our mastodon
specimen. Olsen (1972) noted that there are few diagnostic
dierences between the mammoth and mastodon in these
vertebrae apart from the shape of the vertebral foramen.
The vertebral foramen in mastodons is more rounded
dorsally, whereas that of the mammoth is triangular in shape
(Olsen, 1972). Our third cervical specimen conforms to
the mastodon pattern, appearing more rounded (Pl. 6). The
cervical vertebrae of proboscideans in general are compressed
anteroposteriorly, giving them a disk-like appearance (Olsen,
1972). The cervicals of our specimen are unfused, but Olsen
noted that in both genera they can occasionally fuse together.
Thoracic Vertebrae
Sixteen thoracic vertebrae are present that have been identied
as belonging to a single mastodon. The third through the sixth
thoracics are present, followed by a gap in the sequence and
another sequence of 12 articulating vertebrae that are most
likely the ninth through the twentieth thoracics. As mentioned
above, we have based our estimation on articulations with
ribs. Some of the ribs in the sequence are missing and some of
the articular facets are missing on the middle ribs. The shape
of these ribs was dicult to compare on both sides in this
area because of several healed fractures on the right side. It is
possible that there were 19 thoracic vertebrae rather than 20
if we have overestimated the number of ribs missing in the
sequence.
The mammoth from this site had at least 19 thoracic
vertebrae. As mentioned above, we have a sequence of 15
articulating thoracics. Anterior and posterior thoracics are
missing, but we have anterior and posterior ribs, and have
determined from this sequence that there were originally at
least 19 sets of ribs, and thus 19 thoracic vertebrae with which
they articulated.
The posterior thoracics in both animals incline to a greater
degree than the anterior thoracics (Olsen, 1972), and those of
the mammoth incline to an even greater degree (Pl. 8). The
spinous processes on these vertebrae are most massive toward
the anterior. They sharply increase in length at the third and

fourth thoracics, which are located at the shoulder, and then
decrease more gradually toward the rear. The longest, most
massive spinous processes of the third and fourth thoracics
would have formed a hump behind the shoulders of the
animal. The spinous processes of our mastodon specimen
increase slightly in length again in the most posterior thoracics.
This would have created the slightly dished spinal prole that
is seen in other mastodon specimens as well as to a greater
degree in Loxodonta (Haynes, 1991). The spinous processes
appear to decrease in length through the vertebral column of
our mammoth specimen.
Slight diagnostic dierences in the thoracic vertebrae
observed here include a sharper spine on the anterior face
of the spinous process on the more anterior thoracics
of the mastodon. It is unclear if this is related to dierent
muscle development in these two individuals, gender, or is
a characteristic dierence between the genera. The concavity
on the posterior face of the spinous process is also deeper and
more dened in the mammoth in the anterior thoracics. We
are unable to compare the third thoracic vertebrae of the two
animals, because this element is missing in our mammoth
specimen. From a comparison of the fourth and fth thoracics
of the two animals (see Pls 7-8), it can be seen that the neural
arch is higher and thinner in the mastodon, and lower and
wider in the mammoth. The centra of these vertebrae are also
more robust in the mammoth. The spinous processes of these
vertebrae in our mammoth specimen are longer and incline
to a greater degree than in the mastodon. These characteristics
have been previously documented by Olsen (1972).
In the more posterior thoracics (see Pls 9-10), Olsen
(1972: 9) has noted that the ventral margin of the centrum is
more rounded in the mastodon and approaches a right angle
with the apex on the sagittal plane in the mammoth.
This character was not observed on our mammoth specimen,
which appears more similar to the mastodon with a rounded
form. Observed dierences in the posterior thoracics are the
signicantly longer spinous process in the mammoth, which
inclines to a greater degree than that of the mastodon (also
observed by Olsen, 1972), indicating a more gradual slope
in the lower back of the mammoth. The transverse processes
of these vertebrae also dier in their orientation. In the
mastodon, these project away from the centrum and are
oriented more dorsally, whereas in the mammoth, they have
a more lateral projection and are closer to the centrum. In
the most posterior thoracics of the mammoth, the transverse
process begins to orient more dorsally and to resemble
that of the mastodon. The vertebral foramen appears more
compressed dorsoventrally in the mammoth. The posterior
thoracics of our mammoth specimen also do not show well
dened facets for the ribs.

Lumbar Vertebrae
Three lumbar vertebrae are present for our mastodon
specimen, the last of which is fused to the sacrum. The
vertebrae articulate in sequence with the other mastodon
vertebrae, but can also be recognized independently as
mastodon by their rounded ventral margin (Pl. 11).
Sacrum
Olsen (1972) noted that there is little dierence between the
sacral and caudal vertebrae of mastodons and mammoths. The
sacrum is present for our mastodon specimen, and is fused to
the third lumbar vertebra. It articulates with the left and right
illium, but is not fused to them.
RIBS
Although Olsen (1972) did not describe dierences in the
ribs, some are noted here. Generally, the mammoth ribs
have a stronger arch at the proximal end than those of the
mastodon.
The anterior ribs are not readily distinguishable. The most
posterior ribs, however, are quite variable between the genera.
As seen in the nineteenth rib (Pl. 11), the posterior ribs in
the mastodon are much more short and broad, whereas the
mammoth ribs appear longer and thinner. The facet on the
tubercle of the rib is less pronounced in the mammoth and
absent in the middle and posterior ribs.
As discussed above, we believe that there were originally
20 sets of ribs in the mastodon and 19 in the mammoth. This
count conforms with most other data, although there is some
uncertainty in the literature on the number of these elements
(Haynes, 1991).
SCAPULA
The scapula has been recognized as one of the more important
bones in distinguishing between proboscidean taxa (Olsen,
1972). Our collection has both the right and left scapulae of
Mammut americanum (Pl. 12), identied as such based on
several diagnostic characteristics described by Olsen (1972).
Olsen (1972) described the midspinous process as positioned
midway along the scapular spine in Mammuthus primigenius
and closer to the acromion in Mammut americanum. This
process is broken in both scapulae, but the position of the
break is more proximal and nearer to the acromion process.
The vertebral border of the scapula is straight in both of our
specimens, which Olsen (1972) noted is a characteristic of
the mastodon, whereas that of the mammoth is concave. The
scapular neck is more constricted in M. primigenius than in
M. americanum according to Olsen (1972), but neither of our
specimens shows a high degree of constriction in this area.
Olsen (1972) described the glenoid cavity as more rectangular
in the mammoth and more irregular in the mastodon. We
305
Table 2. Measurements (in mm) of Watkins Glen mastodon scapula

(after Agenbroad, 1994).
Right
Left
763
775
632
642
779
783
276
275
266
264
220
226
141
145
would describe the glenoid cavity as more oval in shape in

our specimen, as well as in Olsens (1972) photographs, rather
than as irregular. See Table 2 for measurements.
PELVIS
A complete pelvis, which we have identied as belonging to
our mastodon specimen, is present in this assemblage (Pl. 13).
Olsen (1972) did not describe dierences in the pelvis of the
mastodon and mammoth, except in noting that the mastodon
pelvis is larger. We have identied this element as mastodon
based on its articulation with the vertebral column and hind
limbs. The pelvis is an important element for determining
gender in many animals. Basic dierences between male and
female proboscidean pelves include a smaller pelvic aperture
in males as well as a broader ilium.
Based on the size of the tusk (see Fisher, 2008) as well
as pelvic morphology, the mastodon appears be a male.
Sexual dimorphism in pelves has been used to determine
gender in Mammuthus and Loxodonta (Haynes, 1991; Lister
& Agenbroad, 1994; Lister, 1996). Although no data for
comparison are available for Mammut (but see Fisher, 2008;
Fisher et al., 2008), our measurements fall into the male range
of variation for Mammuthus, as dened by Lister (1996). It can
be expected that measurements for Mammut would follow the
same pattern. Lister (1996) has shown that the most clearly
distinguishing measurement for the pelvis of Mammuthus is
the ratio between the maximum horizontal aperture width
(see Table 3, measurement 3) and the minimum width of the
ilial shaft taken perpendicular to the long axis of the ilium
(Table 3, measurement 5). Lister (1996) observed that there
is almost complete separation between males and females for
this value, with males having a ratio of less than 2.6-2.7 and
females having a ratio greater than this value. This ratio for
306
Table 3. Measurements (in cm) of Watkins Glen mastodon pelvis

(after Lister, 1996).
159
42
41
4
5
Table 4. Measurements (in mm) of Watkins Glen mastodon humerus (after Agenbroad, 1994).
Right
Left
937
65
599
24
916
132
267
264
231
the Watkins Glen Mastodon is 1.75 (apertural width = 41

cm; ilial shaft width = 24 cm). If pelvic morphology follows a
similar pattern to that of Mammuthus, this ratio clearly places
our specimen in the male range.
GENERAL COMMENTS ON THE LIMB BONES
The limb bones of Mammut are shorter and more robust
than those of Mammuthus (as well as modern elephants).
Lengthening of the forelimbs might be an evolutionary trend
within the Elephantidae, whereas Mammut might retain the
more primitive state. Kubiak (1982) suggested that this could
be related to feeding adaptations. Mammoths and modern
elephants are browsers, which requires them to reach up
to higher tree branches. Haynes (1991) suggested that the
dierence could also be related to locomotor eciency, tusk
size, or some other selective pressure. Tables 4-9 provide metric
data for the limb bones present in our collection. Agenbroad
(1994) documented these measurements extensively for
Mammuthus from the Mammoth Hot Springs Site, but metric
data are not well documented for Mammut.
Humerus
Our collection contains one complete left humerus that has
been identied as mastodon (Pl. 14). In Mammuthus, this bone
appears longer and thinner, with a more gracefully sloping
shaft (Olsen, 1972: 16; Table 4). Our specimen is more robust
and has a more prominent supracondyloid ridge, which are
characteristics that Olsen (1972) attributed to Mammut. A
broken proximal portion of a right humerus is also present in
the collection and appears to be from the same individual.
Radius
The radii of both genera are similar (Olson, 1972: 16; Table
5). As with most of the other long bones, however, this bone is
more robust in the mastodon, especially at the distal end. We
have identied this bone in our collection as mastodon based
on its articulation with the ulna (Pl. 15).
Table 5. Measurements (in mm) of Watkins Glen mastodon radius

69
50
120
189
Table 6. Measurements (in mm) of Watkins Glen mastodon ulna

782
564
109
635
150
350

Ulna
The ulnae of both animals are quite similar as well, and were
described in detail by Olsen (1972). The ulna of the mastodon
has a slightly more developed olecranon process and deeper
semilunar notch, which is apparent in our specimen from
comparisons of photos (Table 6; Pl. 15). Olsen (1972) also
described the mastodon as having a greater overhang of the
olecranon process at the proximal end. This feature is dicult
to observe without comparative material.
Femur
The femur is quite similar in both animals, but the mastodon
femur is somewhat shorter and thicker than that in the
mammoth. The distal condyles appear more expanded medially
and laterally in the mastodon. Olsen (1972) noted that this
is a subtle dierence, but it is quite distinctly observed in our
specimen, which we have determined to be mastodon based
also on its articulation with the pelvis.
Haynes (1991) discussed sexual dimorphism in living and
extinct proboscideans, and argued that bone measurements,
especially femur length, show that extinct species were also
sexually dimorphic. A comparison of the femur of the Watkins
Glen specimen (which has been determined to be male) and
the North Java mastodon specimen (PRI 49618; which is
likely female) appears to follow this pattern (Pl. 16).
Modern adult male African elephants have femoral and
humeral lengths ca. 17-25% longer than those of females
(Haynes, 1991). This dimorphism is also seen in fossil
proboscidean bones, and is presumed to represent size
dierences between the sexes. The Berelyokh mammoth
bones at the Zoological Institute of the Academy of Sciences
in Saint Petersburg have a 14-26% dierence in femur length
(Haynes 1991). Haynes (1991) also reported that the Denver
mastodon (on display at the Milwaukee Public Museum) and
the mastodon from the John Neath site (in the collections of
the University of Wisconsin, Madison, Museum of Zoology)
have a dierence in limb bone length of 15-27%. This
dierence in size has been observed in other mastodon limb
bones as well (Haynes, 1991). The femur of our Watkins Glen
mastodon is 13.16% longer than the femur of our North Java
specimen (Hodgson et al., 2008; Table 7). The sex of the
North Java specimen has not been rmly established, but has
been determined to likely represent a female based on this size
dierence (Fisher, 2008). This percentage, however, is on the
border of dierences that have been observed.
Patella
The right patella of both specimens is present in our collection
(Pl. 17). The mammoth patella has been heavily gnawed by
a carnivore, but the diagnostic features are still visible. Olsen
(1972: 11) noted that the mastodon patella has an expanded
307
Table 7. Measurements (in mm) of Watkins Glen and North Java

mastodon femora (after Agenbroad, 1994). *See Hodson et al.
(2008).
Watkins
Java*
1,075
950
160
146
389
317
275
240
160
144
156
132
lip on the margin of the articular surface and a prominent

apex, whereas the mammoth patella lacks these features. Our
mastodon specimen displays the features described by Olsen
(1972) for Mammut. The caudal border of the mammoth
specimen is missing due to carnivore gnawing, but the anterior
surface does not appear to slope enough to form a lip. In
addition, the articular facet on the mastodon appears concave,
whereas this face on the mammoth patella has a atter surface.
This is to be expected because the condyles of the femur where
they articulate are more developed in the mastodon.
Table 8. Measurements (in mm) of Watkins Glen mastodon tibia

662
104
260
152
214
308
Tibia
The tibiae of Mammut and Mammuthus are dicult to
distinguish (Olsen, 1972; Table 8). No notable dierences
are observed here, but the element has been photographed
for reference (Pl. 17). There is both a right and left tibia in
our collection from the same individual. The tibiae in our
collection articulate with the rest of the mastodon skeleton.
Fibula
A left mastodon bula (Pl. 17) and a probable right mammoth
bula are present in our collection. The right bula consists
only of the shaft portion, and so is dicult to describe, but it
appears to be from a dierent individual than the left bula.
The left bula appears to be mastodon based on its articulation
with the mastodon tibia. It also appears similar in form to the
bula of the North Java mastodon specimen (Hodgson et al.,
2008; Table 9). Olsen (1972: 22) described the mammoth
bula as considerably more elongated, narrower, and more
pointed at the proximal end. These dierences are quite
subtle, however, and are dicult to recognize in the absence
of comparative material.
Fore Foot
All foot bones in our collection have been identied on
the basis of their morphology and articulations. The most
diagnostic faces of the bones have been photographed.
The reader is referred to Olsens work (1972) for complete
illustrations of all faces of these bones.
Table 9. Measurements (in mm) of Watkins Glen and North Java

mastodon bulae (after Agenbroad, 1994).
Watkins
Java
621
587
451
525
52
41
89
87
142
116
Pisiform.Olsen (1972) noted there is little dierence

between the pisiforms of the mammoth and mastodon.
Our assemblages has one complete right and one partial left
mammoth pisiform, identied on the basis of its articulation
with the other mammoth foot bones. None is present for
the Watkins Glen mastodon, although we do have a partial
right pisiform from our North Java mastodon specimen for
comparison (Pl. 19). Slight dierences between the genera
were noted in this bone. The mastodon bone appears to
be a more broad, compact bone, whereas the mammoth
pisiform appears slightly longer and thinner. The dorsomedial
surface around the articular facet is somewhat broader in the
mastodon.
Cuneiform.Our assemblage contains the left mastodon
cuneiform and the right mammoth cuneiform. The lateral
projection of the cuneiform is somewhat longer in the
mastodon. Dorsoventrally, the bone is somewhat thicker in
the mammoth (Pl. 20).
Lunar.The lunar is present from both the mastodon (left
side) and mammoth (right side) in our asssemblage. Olsen
(1972: 22) noted that this bone, when viewed ventrally in the
mastodon has a dished articular surface on the lateral face,
while this same surface in the mammoth forms a nearly right
angle with the straight articular face of the lateral surface.
Although the margin of the lateral and anterior faces do form
a near-right angle in the mammoth, this dishing was not
observed in our mastodon specimen, nor in the North Java
mastodon specimen. Rather, this margin in the mastodon
lunar appears to form a more acute angle when viewed
ventrally. General dishing of the ventral articular surface itself
was observed in all specimens (Pl. 21).
Scaphoid.A scaphoid from both our mastodon and
mammoth specimens is present in our collection. Olsen
(1972) noted no discernible dierences between the two taxa
in this element, save for slight dorsoventral compression in
the mastodon when viewed laterally (or medially). We were
unable to observe this due to fracturing in our specimen.
Unciform.The left mastodon and right mammoth unciforms
are present in our collection. No notable dierences are
observed between taxa in this element. We have photographed
them for identication purposes (Pl. 22).
Trapezoid.The left mastodon and right mammoth trapezoids
are present in our collection. The dorsal and ventral articular
surfaces are somewhat broader in that of the mastodon. No
other signicant dierences between taxa for this element
were observed (Pl. 23).
309

Trapezium.Our collection contains a trapezium from both
our mastodon and mammoth specimens. Olsen (1972) noted
considerable dorsoventral compression in the trapezium,
which is also observed on our specimens. This bone has
been photographed from all angles because of the signicant
variation in shape between the species (Pl. 24).
specimen. The mastodon naviculars have been identied by

their articulations with the mastodon astragali. Olsen (1972)
did not note any dierences between the naviculars of the
two genera, but two subtle dierences are observed here. In
the mastodon, this bone is thinner anteroposteriorly, and
the concave curvature on the ventral surface is deeper. In the
mammoth, this bone is thicker anteroposteriorly and less
curved on the ventral surface (Pl. 25). These observations
appear to also hold true for Olsons (1972: 40) published
photographs of the two genera as well as the navicular of the
North Java mastodon (Hodgson et al., 2008).
Metacarpals.The metacarpals of proboscideans are unique

from those of other animals such as equids or bovids in that
they are longer and more robust than the metatarsals. This is
because the majority of the animals weight is carried by the
front limbs (Mol & Agenbroad, 1994). The metacarpals of
Mammuthus appear longer than those in Mammut, whereas
those of Mammut appear a bit more robust. No signicant
morphological dierences on the proximal articular surfaces
were noted between the mammoth and mastodon, however,
they were photographed here to aid in identication, because
these bones are often overlooked (Pl. 18).
Mol & Agenbroad (1994) extensively described the
metapodials of Mammuthus columbi. Their measurements
are reproduced below, along with the measurements of our
specimens for comparison (see Table 10).
Calcaneum.Both the right and left calcanea of the mastodon

are present and were identied on the basis of their articulations
with the right and left astragalus (Pl. 26). In addition, they
possess diagnostically mastodon characteristics. Olsen (1972:
36) described the tuber calcis as expanded in the mastodon
and postlike in the mammoth. Our specimens display the
expanded tuber calcis. When viewed from the posterior, the
sustentaculum is not oriented dorsally as in the mammoth,
but more ventrally. This process is also notched on the ventral
border, which Olsen (1972) noted is a characteristic of the
mastodon calcaneum.
Hind Foot
The hind foot of each genus is relatively smaller and more
narrow than the fore foot. As discussed above, this is because
the front foot bears more of the animals weight. Some
diagnostic elements that were preserved in our collection are
discussed below. Elements were identied to genus based on
their morphology and articulations.
Astragalus.Both the right and left astragali of the mastodon

are present, and were identied by their articulations with
the right and left tibiae of the mastodon. A pronounced
condyle is present on the posterior surface of the bone, which
Olsen (1972) noted is only present in the mastodon. Olsen
(1972) also noted that the bone is considerably compressed
dorsoventrally in the mammoth and more expanded in
the mastodon when viewed anteriorly. More compression,
however, is observed in our specimen than in Olsens (1972)
Navicular.Our collection contains both naviculars from

our mastodon and the right navicular of our mammoth
Table 10. Comparative metacarpal measurements of Hot Springs Mammoth MSL 033 (HS), Watkins Glen Mammoth (MA) and Watkins Glen
Mastodon (MS) (after Mol & Agenbroad, 1994). All measurements in mm; n/d = no data.
MCI
MCII
MCIII
MCIV
MCV
HS
MA
MS
HS
MA
MS
HS
MA
MS HS
MA
MS
HS
MA
MS
123
n/d
n/d
208
n/d
160 237
204
171 202
193
157
181
170
126
Proximal epiphysis
62
n/d
n/d
90
78
80
93
78
78
71
96
93
93
80
104
Distal epiphysis
67
n/d
n/d
109
91
80
115
102
90
105
100
97
100
94
89
Mid-shaft
49
n/d
n/d
76
72
72
79
76
75
83
80
100
71
68
79
Proximal epiphysis
98
n/d
n/d
100
128
101 128
123
102 115
116
107
108
106
99
Distal epiphysis
60
n/d
n/d
103
96
93
110
92
94
100
96
97
99
100
88
Mid-shaft
69
n/d
n/d
74
65
51
72
61
60
72
64
61
101
95
91
Greatest length
Transverse diameter:
Anteroposterior diameter:
310
mastodon specimen (Pl. 27).

EPIPHYSEAL FUSION AND AGING
There is some uncertainty in the literature on the lifespan of
modern elephants, but an upper age limit of 60-65 yr seems
to be most accepted (Haynes, 1991). Modern elephants have
a long growth period, which extends into middle age (Lister,
1994). The timing of epiphyseal fusion can been used as a
tool for determining individual animal ages. Roth (1984)
compiled an epiphyseal fusion sequence for modern African
and Asian elephants. Epiphyseal fusion states for elephants
seem to be regular, with minor dierences between species
and sexes (Lister, 1994). Haynes (1991) and Lister (1994)
argued that epiphyseal fusion in fossil proboscideans appears
to follow tooth progression and wear in a similar pattern to
that of modern elephants, implying that life span was probably
also similar in these fossil species.
Bone fusion in the Watkins Glen specimens is detailed in
Table 11 and is compared to the fusion schedules of Haynes
(1991) and Lister (1994). Most bones with diagnostic
epiphyses were not present for our mammoth specimen, but
the age of this individual can be estimated by other means.

All the centra of the vertebral column are fused to the arches,
and the plates are in the process of fusing to the centra.
Approximately half of these are fused and the suture lines are
still visible. This would place the individual at at least 20 yr
of age, but more likely quite a bit older. Haynes (1991) noted
that fusion of centra to plates could start around the age of
20, but continues into the 50s. Considering that the plates
are approximately half fused, the animal was perhaps around
the age of 40. Evidence of arthritis in the ribs could place it
a bit older.
Evidence of epiphyseal fusion in our mastodon specimen
place the animal approximately in its 40s or 50s. The distal
radius is fused, which probably occurs during the late 40s
(Haynes, 1991). Also, the vertebral centra are all fused, which
Haynes (1991) noted normally is not completed until the 50s.
However, the sacrum is not fused to the pelvis. Haynes (1991)
noted that this bone can remain unfused in a male into the
50s. Based on this information, it appears that the animal was
probably in its late 40s or early 50s .
Table 11. Epiphyseal fusion in Watkins Glen mastodon and mammoth. f, fused (fusion schedules after Haynes, 1991, and Lister, 1994); n/d,
no data.
Age at which epiphyses have fully fused (yr)
Element
Female
Male
Mastodon
Mammoth
Femur
Distal
17-23
26-29
n/d
Proximal
25-32
> 29 (often unfused

into late 30s)
n/d
ca. 19 (18-20)
ca. 32?
n/d
17-24 (probably > 20)
28-32 (possibly earlier)
n/d
Tibia
Distal
Proximal
Humerus
Distal
< 19 (ca. 13-14)
> 18
n/d
fusing 19-26
ca. 32 (often unfused

into late 30s)
n/d
> 24
> 32 (probably late

40s)
f/f
n/d
fusing at 19
< 32 (probably by 20s)
f/f
n/d
some centra fused to

arches, plates attached
middle vertebrae only (<
19)
lengthy fusion
all arches and centra

fused
all centra fused to

arches, some plates
fused to centra
Ilium, ischium, pubis
fusing at ca. 8
fused at 8
n/d
n/d
Innominate/Sacrum
< 19
> 32
sacrum unfused
n/d
Proximal
Radius/Ulna
Distal
Proximal
Vertebral centra and plates
PATHOLOGIES AND MODIFICATIONS

Analysis of pathologies and modications on the Watkins
Glen mastodon and mammoth were beyond the scope of
this project (see Erwin Grundmann, 2008). Several general
observations, however, are made here.
Our mastodon specimen displays quite a severe break on
its right side in ribs 6, 7, and 9 (rib 8 is missing). This break
has healed, but did not heal well. False joints that have formed
at the healed break and extra bone growth are possibly signs of
chronic infection after receiving the wound or of rebreaking.
Muscle attachments on the left side of the vertebrae that
articulate with these ribs are slightly more developed, which
might suggest that the mastodon was carrying its weight on
the side opposite its wound for some time. No signicant
pathologies were noted in our mammoth specimen apart from
some arthritis in the ribs, which is not likely to be abnormal
given its estimated age.
Carnivore gnawing has been observed on both specimens,
but has not been thoroughly studied or quantied.
CONCLUSIONS
The Watkins Glen Site presents an interesting opportunity for
a comparison of the postcranial morphology of Mammut and
Mammuthus. From the descriptions above, it can be seen that
although strikingly similar, postcranial elements of these two
genera can be dierentiated. Subtle morphological dierences
between these genera are likely related to their adaptations
to diering habitats. For this reason, it is equally interesting
that we nd these two genera at the same site. Radiocarbon
dates suggest their occurrence at the site was not separated by
a great amount of time and could represent a co-occurrence.
The descriptions of postcranial morphology above add to the
growing body of knowledge on morphological dierences
between these taxa, and we hope can be a valuable aid to the
researcher in identifying these genera and distinguishing them
from one another.
ACKNOWLEDGMENTS
We would like to thank the many volunteers who participated
in the Watkins Glen excavation, as well as museum volunteers
who participated in this project (especially Francesca Allaby
and Verity Whalen). Also many thanks for helpful comments
from reviewers of this manuscript
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313
Appendix 1. Element representation (with bone numbers) for Watkins Glen mastodon (PRI 8829) and mammoth (PRI 8830) (see Haynes,
1991: 38-39; Shoshani, 1996: 11). L = left; R = right. Note: Some bones exist in more than one piece, thus more than one bone number.
Element
No. in Full Skeleton
Mastodon
Mammoth
Unidentied
Skull
37 fragments
Mandible
1 (R 207, L 208)
Basihyoid
Stylohyoid
2 (259, 262)
1 (263)
Thyrohyoid
2 (260, 261)
2 (124, 289, 290)
Cervical vertebrae
5 (173, 230, 231, 233, 234, 236)
Thoracic vertebrae
18-20
17 A
15 B
Lumbar vertebrae
3-5
3 (172, 210, 251)
Sacral vertebrae
3-5
5 (209, 210)
Caudal vertebrae
21+
4 (87, 103, 244, 274)
Ribs
18-20 pairs
16 L, 17 R C
14 L, 12 R D
Sternum
2 (107, 175)
Scapula (L/R)
2 (L 200, R 222)
Humerus (L/R)
2 (R 185, L 204)
Radius (L/R)
1 (L 76)
Ulna (L/R)
1 (L 78)
Scaphoid (L/R)
1 (L 97)
1 (R 205)
Lunar (L/R)
2 (L 218, R 224)
1 (R 174)
Cuneiform (L/R)
1 (L 128)
1 (R 70)
Pisiform (L/R)
2 (L 125, R 104)
Trapezium (L/R)
1 (L 130)
1 (R 123)
Trapezoid (L/R)
1 (L 129)
1 (R 176)
Magnum (L/R)
1 (R 183)
Unciform (L/R)
1 (L 106)
1 (R 187)
Metacarpals
5 pairs
4 (L 39, 68, 213, 226)
4 (R 67, 80, 84, 86)
Phalanges (anterior)
28
1 (229)
3 (89, 181, 228)
Innominate (L/R)
2 (L 206, R 297)
Femur
2 (L 296, R 203)
Patella
1 (R 98)
1 (R 164)
Tibia
2 (L 79, R 72)
Fibula
1 (L 81)
1 (R 96)
Astragalus (L/R)
2 (L 177, R 122)
Calcaneum (L/R)
2 (L 227, R 37)
Navicular (L/R)
2 (L 33, R 132)
1 (R 105)
Cuboid (L/R)
Internal cuneiform (L/R)
1 (R 165)
External cuneiform (L/R)
1 (R 134)
Metatarsals
5 pairs
3 (L 223)
2 (115, 142)
Phalanges (posterior)
28
Tusks
1 (303)
47, 102, 201, 212, 232, 238-240, 242-243, 246, 249-250, 252-253, 257, 293.
28, 30, 48, 56, 71, 83, 88, 116, 157, 202, 212, 235, 237, 245, 247-248, 254-256, 258, 294.
C
L 6, 11, 18, 26, 29, 40, 43, 49-50, 55, 74, 82, 101, 152, 279, 281, 299; R 3, 7, 10, 12, 14, 16-17, 20, 31, 38, 45, 51, 57-58, 61, 63, 75, 100, 137, 211, 282, 295.
D
L 13, 21, 35, 42, 53-54, 62, 90, 109-112, 298; R 5, 8-9, 15, 19, 23, 25, 34, 44, 52, 60, 99.
B
314
Plate
Complete skeleton of the Watkins Glen mastodon (PRI 8829).
315
316
Plate
Complete skeleton of the Watkins Glen mammoth (PRI 8830).
317
318
Plate
Figure
1-2
Mastodon mandible (PRI 8829, bones 207, 208).

1. Occlusal view.
2. Lateral view of left side.
319
320
Plate
Figure
1-2
Mastodon atlas (PRI 8829, bone 173).

1. Anterior view.
2. Posterior view.
321
322
Plate
Figure
1-2
Mastodon axis (PRI 8829, bone 234).

1. Anterior view.
2. Lateral view.
323
324
Plate
Figure
1-2
Mastodon third cervical vertebra (PRI 8829, bone 231).

1. Anterior view.
2. Lateral view.
325
326
Plate
Figure
1-2
3-4
Mastodon fourth thoracic vertebra (PRI 8829, bone 246).

1. Anterior view.
2. Posterior view.
Mammoth fourth thoracic vertebra (PRI 8830, bones 235).
3. Anterior view.
4. Posterior view.
327
328
Plate
Figure
1-3
4-6
Mastodon fth thoracic vertebra (PRI 8829, bone 293).

1. Anterior view.
2. Posterior view.
3. Lateral view.
Mammoth fth thoracic vertebra (PRI 8830, bone 202).
4. Anterior view.
5. Posterior view.
6. Lateral view.
329
330
Plate
Figure
1-2
3-4
Mastodon ninth thoracic vertebra (PRI 8829, bone 249).

1. Anterior view.
2. Lateral view.
Mammoth ninth thoracic vertebra (PRI 8830, bones 30 and 48).
1. Anterior view.
2. Lateral view.
331
332
Plate
Figure
1-2
3-4
Mastodon thirteenth thoracic vertebra (PRI 8829, bone 252).

1. Anterior view.
2. Lateral view.
Mammoth thirteenth thoracic vertebra (PRI 8830, bone 294).
3. Anterior view.
4. Lateral view.
333
334
Plate
Figure
1-2
3
4
Mastodon rst lumbar vertebra (PRI 8829, bone 251).

1. Anterior view.
2. Lateral view.
Mastodon nineteenth right rib (PRI 8829, bone 38).
Mammoth nineteenth right rib (PRI 8830, bone 60).
335
336
Plate
Figure
1-2
Mastodon right scapula (PRI 8829, bone 222).

1. Lateral view.
2. Glenoid cavity.
337
338
Plate
Mastodon pelvis (PRI 8829, bones 206, 297).
339
340
Plate
Figure
1-3
Mastodon left humerus (PRI 8829, bone 204).

1. Anterior view.
2. Posterior view.
3. Proximal view.
341
342
Plate
Figure
1-2
3-4
Mastodon left radius (PRI 8829, bone 76).

1. Lateral view.
2. Medial view.
Mastodon left ulna (PRI 8829, bone 78).
3. Lateral view.
4. Medial view.
343
344
Plate
Figure
1-3
4-6
Mastodon left femur (PRI 8829, bone 203).

1. Anterior view.
2. Posterior view.
3. Distal view.
North Java mastodon right femur (PRI 49618, bone 141).
4. Anterior view.
5. Posterior view.
6. Distal view.
345
346
Plate
Figure
1-2
3-4
5-6
7-8
Mastodon riht tibia (PRI 8829, bone 72).

1. Anterior view.
2. Posterior view.
Mastodon left bula (PRI 8829, bone 81).
3. Medial view.
4. Lateral view.
Mastodon right patella (PRI 8829, bone 98).
5. Ventral view.
6. Lateral view.
Mammoth right patella (PRI 8830, bone 164).
7. Ventral view (incomplete due to carnivore gnawing).
8. Lateral view.
347
348
Plate
Figure
1
2
3a-h
Mastodon left fore foot (PRI 8829).

Mastodon left metacarpals, proximal view (PRI 8829, bones 213, 226, 39, 68).
Mammoth right fore foot (PRI 8830).
a. Pisiform (hidden) (104).
b. Cuneiform (7).
c. Lunar (174).
d. Scaphoid (205).
e. Unciform (187).
f. Magnum (183).
g. Trapezoid (176).
h. Trapezium (123).
Mammoth right metacarpals, proximal view (PRI 8830, bones 84, 67, 80, 86).
349
350
Plate
Figure
1-3
4-5
Mammoth right pisiform (PRI 8830, bone 104).

1. Anterior view.
2. Medial view.
3. Lateral view.
North Java mastodon left pisiform (PRI 49618, bone 39).
4. Medial view.
5. Lateral view.
351
352
Plate
Figure
1-4
5-8
Mastodon left cuneiform (PRI 8829, bone 128).

1. Anterior view.
2. Posterior view.
3. Dorsal view.
4. Ventral view.
Mammoth right cuneiform (PRI 8830, bone 7).
5. Anterior view.
6. Posterior view.
7. Dorsal view.
8. Ventral view.
353
354
Plate
Figure
1-2
3-4
Mastodon left lunar (PRI 8829, bone 218).

1. Ventral view.
2. Medial view.
Mammoth right lunar (PRI 8830, bone 174).
3. Ventral view.
4. Medial view.
355
356
Plate
Figure
1-3
4-6
Mastodon left unciform (PRI 8829, bone 106).

1. Dorsal view.
2. Ventral view.
3. Medial view.
Mammoth right unciform (PRI 8830, bone 187).
4. Dorsal view.
5. Ventral view.
6. Medial view.
357
358
Plate
Figure
1-2
3-4
Mastodon left trapezoid (PRI 8829, bone 129).

1. Dorsal view.
2. Ventral view.
Mammoth right trapezoid (PRI 8830, bone 176).
3. Dorsal view.
4. Ventral view.
359
360
Plate
Figure
1-6
7-12
Mastodon left trapezium (PRI 8829, bone 130).

1. Anterior view.
2. Posterior view.
3. Dorsal view.
4. Ventral view.
5. Lateral view.
6. Medial view.
Mammoth right trapezium (PRI 8830, bone 123).
7. Anterior view.
8. Posterior view.
9. Dorsal view.
10. Ventral view.
11. Lateral view.
12. Medial view.
361
362
Plate
Figure
1-2
3-4
Mastodon right navicular (PRI 8829, bone 123).

1. Anterior view.
2. Dorsal view.
Mammoth right navicular (PRI 8830, bone 105).
3. Anterior view.
4. Dorsal view.
363
364
Plate
Figure
1-4
Mastodon right calcaneum (PRI 8829, bone 37).

1. Anterior view.
2. Posterior view.
3. Lateral view.
4. Medial view.
365
366
Plate
Figure
1-4
Mastodon right astragalus (PRI 8829, bone 122).

1. Anterior view.
2. Posterior view.
3. Dorsal view.
4. Ventral view.
367

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Mastodon Paleobiology,

Bone collagen from the Watkins Glen mastodon specimen

Palaeontographica Americana, No.

represent M. primigenius Blumenbach, 1799, the woolly

period leading up to the extinction of both species.

GENERAL DIFFERENCES BETWEEN TAXA

HODGSON ET AL.: COMPARATIVE OSTEOLOGY OF MAMMOTH AND MASTODON

Table 1. Number of ribs and vertebrae for each species (modied

those of Mammuthus, whereas those of Mammuthus were

similar to the prole of Loxodonta, but with slightly less

Palaeontographica Americana, No.

Axis.The axis is dicult to distinguish between taxa. We have

the anterior. They sharply increase in length at the third and

HODGSON ET AL.: COMPARATIVE OSTEOLOGY OF MAMMOTH AND MASTODON

Table 2. Measurements (in mm) of Watkins Glen mastodon scapula

would describe the glenoid cavity as more oval in shape in

Palaeontographica Americana, No.

Table 3. Measurements (in cm) of Watkins Glen mastodon pelvis

the Watkins Glen Mastodon is 1.75 (apertural width = 41

Table 5. Measurements (in mm) of Watkins Glen mastodon radius

Table 6. Measurements (in mm) of Watkins Glen mastodon ulna

HODGSON ET AL.: COMPARATIVE OSTEOLOGY OF MAMMOTH AND MASTODON

Table 7. Measurements (in mm) of Watkins Glen and North Java

lip on the margin of the articular surface and a prominent

Table 8. Measurements (in mm) of Watkins Glen mastodon tibia

Palaeontographica Americana, No.

Table 9. Measurements (in mm) of Watkins Glen and North Java

Pisiform.Olsen (1972) noted there is little dierence

HODGSON ET AL.: COMPARATIVE OSTEOLOGY OF MAMMOTH AND MASTODON

specimen. The mastodon naviculars have been identied by

Metacarpals.The metacarpals of proboscideans are unique

Calcaneum.Both the right and left calcanea of the mastodon

Astragalus.Both the right and left astragali of the mastodon

Navicular.Our collection contains both naviculars from

Palaeontographica Americana, No.

mastodon specimen (Pl. 27).

the age of this individual can be estimated by other means.

> 29 (often unfused

17-24 (probably > 20)

28-32 (possibly earlier)

< 19 (ca. 13-14)

ca. 32 (often unfused

> 32 (probably late

< 32 (probably by 20s)

some centra fused to

all arches and centra

all centra fused to

Ilium, ischium, pubis

HODGSON ET AL.: COMPARATIVE OSTEOLOGY OF MAMMOTH AND MASTODON

PATHOLOGIES AND MODIFICATIONS

in: The Hot Springs Mammoth Site: a Decade of Field and

Palaeontographica Americana, No.

mammoths and mastodons: reconstruction of migratory behavior

Site: a Decade of Field and Laboratory Research in Paleontology,

HODGSON ET AL.: COMPARATIVE OSTEOLOGY OF MAMMOTH AND MASTODON

No. in Full Skeleton

2 (124, 289, 290)

5 (173, 230, 231, 233, 234, 236)

3 (172, 210, 251)

4 (87, 103, 244, 274)

4 (L 39, 68, 213, 226)

4 (R 67, 80, 84, 86)