Hodgson et al. (2008) -- Mastodon and Mammuthus osteology.pdf

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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


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INRODUCIONTe 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 theosteology 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. Althoughthe geographic ranges of these two genera were partlyseparate, they did overlap, most extensively in the MississippiRiver drainage basin (Shoshani & assy, 1996b), and New

York State was within the bounds of this overlapping area.

Bone collagen from the Watkins Glen mastodon specimenhas been radiometrically dated to 10,780 ± 60 14C yr BP(conventional 10,840 ± 60 yr BP; Beta Analytic 176930), andthe mammoth specimen has been dated to 10,820 ± 50 14C yrBP (conventional 10,840 ± 50 yr BP; Beta Analytic 176929).

Te margins of error of these dates overlap, suggesting thatthe species very likely coexisted at the site, or at least were notseparated by more than ca. 150 years. If the site does in factrepresent actual sympatry of the two species, which is not welldocumented at other sites, this would raise important issuesabout changing habitats and potential ecological competitionbetween these species in the Late Pleistocene.

Based on known distributions of Mammuthus spp. throughspace and time, it is likely that the mammoth remains at the

Watkins Glen site (PRI 8830, formerly CU 42440 in part)

COMPARAIVE OSEOLOGY OF LAE PLEISOCENE MAMMOH AND MASODONREMAINS FROM HE WAKINS GLEN SIE, CHEMUNG COUNY, NEW YORK

J A. H

Paleontological Research Institution, 1259 rumansburg Road, Ithaca, New York 14850, U. S. A. Currentaddress: Department of Anthropology, City University of New York, 365 Fifth Avenue, New York, New York

10016-4309 U. S. A., email [email protected].

W D. APaleontological Research Institution, 1259 rumansburg Road, Ithaca, New York 14850, U. S. A., andDepartment of Earth and Atmospheric Sciences, Cornell University, Ithaca, New York 14853, U.S.A.

P L. NDepartment of Earth and Atmospheric Sciences, Cornell University, Ithaca, New York 14853, U. S. A. Current

address: Chevron Energy echnology Company, 1500 Louisiana Street, Houston, exas 77002, U. S. A.

J M. S Paleontological Research Institution, 1259 rumansburg Road, Ithaca, New York 14850, U. S. A . Current

address: 302 Giles Street, Ithaca, New York 14850, U. S. A.

and J J. CDepartment of Earth and Atmospheric Sciences, Cornell University, Ithaca, New York 14853, U. S. A. Current

address: 5981 Route 228, rumansburg, New York 14886, U. S. A.

ABSRAC Te 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 quitediffi cult to distinguish specific bones between these taxa in the absence of teeth. Te descriptions of postcranial morphology here add to ourunderstanding of characteristics of the two taxa. In addition, this site presents evidence for Mammut and Mammuthus possibly sharing the samehabitat. 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 LatePleistocene.

Chapter 17, in Mastodon Paleobiology, Taphonomy, and Paleoenviron-

ment 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.


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represent M. primigenius Blumenbach, 1799, the woollymammoth. Alternatively, they could represent M. columbi (Falconer, 1857) [= M. jeffersonii (Osborn, 1922)]. Mammuthuscolumbi was a larger, more southern Late Pleistocene species

mainly occupying the midwestern United States (Graham,2001). Although the ranges of these two mammoth speciesdid overlap in the New York State area, they are believed tohave had different habitat preferences (Agenbroad, 1984):

M. primigenius preferred a forest/woodland or tundraenvironment, whereas M. columbipreferred a “steppe/savanna/parkland” environment (Graham, 2001: 707). It is diffi cult todistinguish the species in the absence of teeth, because theirpostcranial anatomy is almost identical (Haynes, 1991).

Te more complete mastodon skeleton from the WatkinsGlen site (PRI 8829, formerly CU 42440 in part) has beenidentified 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. Te minimumnumber of individuals for each species is one. Additionally, allelements from each animal either articulate with one anotheror appear to be from the same individual.

Te geographic and stratigraphic ranges of Mammut and Mammuthus did overlap, but at well-stratified sites, thespecies almost never occur together (Graham, 2001). Te twoanimals are generally found in separate habitats, but couldhave potentially become competitors due to degradationof their environmental niches in the Late Pleistocene. Teenvironment at the Watkins Glen site was a shallow pondsurrounded by spruce-dominated forest (Griggs & Kromer,

2008). Te 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 thisenvironmental change (King & Saunders, 1984; Robinson etal., 2005). Spruce is thought to have been a preferred food ofmastodons and has been found associated with many othermastodon sites (Miller, 2008). It is evident that spruce wasbeing consumed at the Watkins Glen site from the abundanceof possibly masticated spruce twigs found (Griggs & Kromer,2008). It is therefore interesting that we find the mammothat this site, where the environment appears similar to othermastodon sites. It is possible that these animals were exploiting

separate resources at this same site. It is also possible thatthe mammoth was simply migrating through. Hoppe et al. (1999) described the migratory behavior of mammothsusing strontium isotope ratios from tooth enamel, which wehave not yet attempted on our specimen (no tooth enamel

was recovered for our mammoth specimen, and it is unclearif this test would be successful on other bone material). Acohabitation at the site might also provide evidence forinterspecific competition for dwindling resources during this

period leading up to the extinction of both species.

GENERAL DIFFERENCES BEWEEN AXA Proboscidean species in North America have generally been

described and taxonomically placed on the basis of their dentalcharacteristics. Henry Fairfield Osborn’s Proboscidea (1936,1942), which has long been treated as authoritative on thesubject, has descriptions on dentition. Subsequent literaturehas continued this trend, often excluding detailed analyses ofpostcranial morphology. Morphological differences in teethare useful for identifying fossil species, because teeth are mostoften preserved as fossils. Te researcher encounters diffi culty,however, if no teeth are present. Stanley J. Olsen (1972) hascompiled the most complete documentation of differencesbetween mammoth and mastodon postcranial morphology.His work remains one of the few reference tools for researchersattempting to distinguish proboscidean species in the absence

of teeth, and has been extensively consulted in identification ofremains from the Watkins Glen site. Tis paper verifies manyof Olsen’s observations and notes additional morphologicaldifferences observed between our specimens.

Dentition will be discussed only briefly here (for more in-depth descriptions, see Osborn, 1936, 1942; Maglio, 1973).Te tooth morphologies of both Mammut and Mammuthus are quite distinct, making fossil finds with teeth immediatelyidentifiable. Tese differences are clearly related to the differingdiets and mode of chewing of the two genera. Mastodonshave retained the more primitive gomphotheriid tooth formcharacterized by “high crests of lophs on the occlusal surfaces

of the cheek teeth” (Olsen, 1972: 3). Tis tooth form isadapted for crushing twigs, leaves, and stems. Other evidencealso points toward a browsing adaptation in Mammut (seeHaynes, 1991). Te cheek teeth of Mammuthus , on theother hand, are more similar to modern elephants and arecharacterized by a relatively flat occlusal surface with parallelridges and plates (Olsen, 1972). Tis tooth morphology isa highly specialized adaptation to grinding an abrasive dietof grass (Haynes, 1991). rends over time toward increasingspecialization for this adaptation have led to changes in skullshape. Te skull of Mammuthus is high and domed compared

with that of Mammut , which is a result of a shifting of theskull’s center of gravity related to this mode of mastication

(see Maglio, 1973).Mammoths and mastodons are not taxonomically close

(ext-fig. 1; in fact, their last common ancestor lived ca.20 million years ago; Saunders, 1996), but because theyhave similar “body bulk, mode of walking, and method offood gathering” they logically have more similarities thandifferences (Olsen, 1972: 5). Te overall body proportionsof Mammut and Mammuthus were somewhat different. Ingeneral, the limbs of Mammut were shorter and thicker than


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those of Mammuthus , whereas those of Mammuthus werelonger and relatively more slender (Olsen, 1972). Mammut

was a more robust, heavyset animal; compared to modernElephas , it “had a deeper chest, broader pelvis, shorter legs,and longer back” (Kurtén & Anderson, 1980: 345). Haynes(1991) has suggested that this greater length and breadth ofthe body are related to an increased gut capacity, which couldbe a response to the greater quantity of food needed due to theanimal’s low quality diet of browse. Te head of Mammut wasoriented similarly to Loxodonta , more horizontal in relation tothe body (Hayes, 1980) rather than domed as in Mammuthus .Te body profile of Mammuthus was more similar to modern

Asian elephants, with the shoulder as the highest point of theprofile. Te longest vertebral spines were positioned at thefront of the shoulder and are sharply slanted back. Tis created

a hump at the shoulders and downward sloping back (Haynes,1991). Te profile of Mammut also had its highest point atthe front of the shoulders, but a more gradual rearward tilt(Haynes, 1991).

POSCRANIAL DESCRIPIONS

V EREBRAL COLUMNTe general outline of the vertebral column is quite differentin the two species. Te profile of the vertebral column of

Mammuthus is humped and downward sloping. Te longestvertebral spines are positioned at the front of the shoulder andslant sharply back. Te length of the spines decreases abruptlytoward the rear of the back, creating a slope, which is moreexaggerated by the doming of the cranium. Tis is similar tomodern Asian elephants, but Mammuthus displays an evengreater degree of sloping (Osborn, 1942: 1229; Haynes,1991).

Te profile of the vertebral column of Mammut also hasits highest point at the front of the shoulder, but the spinesdecrease in length more gradually, and increase again slightlytoward the rear, giving the back a dished profile. Tis is most

similar to the profile of Loxodonta , but with slightly lessdishing (Haynes, 1991).

Te number of ribs and vertebrae for these two species hasnot been firmly documented in the literature, and it is likelythat there is variation among individuals (Haynes, 1991).able 1 gives the approximate ranges for these counts in severalskeletons; 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 generallyhas 19. A single individual with 18 thoracic vertebrae has beenreported in each taxon (see Haynes, 1991: 39). Te reductionof vertebral number in Mammuthus is a derived characteralso 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 andchecked the articulations of these elements. Based on thisanalysis, we conclude that the vertebral column of the WatkinsGlen mastodon originally consisted of 7 cervical vertebrae,

20 thoracics, 3 lumbar, and 5 sacral. Also recovered fromthe site were 15 additional thoracic vertebrae belonging tothe mammoth, beginning with the fourth. Based on the ribsequence, there appear to have been at least 19 ribs originally,and thus, 19 thoracic vertebrae. Tree caudal vertebrae arepresent, but we are unable to associate them with one or theother species, because these bones are not diagnostic.

Cervical VertebraeTe first five cervical vertebrae, which articulate with oneanother, as well as the seventh are present for our mastodonspecimen. Tere is then a gap in the vertebral sequence untilthe third thoracic. Although the cervicals do not articulate with

the rest of the vertebral column, and the cranial articulationis missing, their morphology appears diagnostically mastodonas described below.

Atlas.– When viewed from the posterior aspect (Pl. 4), theshape of the dorsal margin appears rounded, where it wouldbe flattened in a mammoth (Olsen, 1972). Te vertebralforamen also appears more oval in shape, whereas a mammoth

would show lateral constriction in this area.

ext-fig. 1. Phyletic diagram illustrating relationships among fami-lies of the order Proboscidea (after Haynes, 1991; Maglio 1979).

able 1. Number of ribs and vertebrae for each species (modifiedfrom Haynes, 1991). C, cervical; , thoracic, L, lumbar; S, sacral;c, caudal.

Ribs C L S cMastodon 18-20 7 18-20 3 4-5 27?

Mammoth 18-20 7 18-20 3-5 3-5 21-22?

HODGSON ET AL.: COMPARATIVE OSTEOLOGY OF MAMMOTH AND MASTODON


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Axis.–Te axis is diffi cult to distinguish between taxa. We haveassigned the axis present in the assemblage to our mastodonspecimen based on its articulation with the atlas and thirdcervical vertebra, which we can identify as mastodon. Olsen

(1972: 9) described both animals as having “high, pronouncedneural spines that are quite variable in outline when seen froma lateral aspect. A notch or trough, running in an antero-posterior direction along the crest of the spine, may be presetor absent in this vertebra of either animal.” Our specimenexhibits an acutely sloping neural spine and a well-definedtrough along the crest (Pl. 5).

Tird Cervical.–Te third through the fifth and also theseventh cervical vertebrae are present for our mastodonspecimen. Olsen (1972) noted that there are few diagnosticdifferences between the mammoth and mastodon in thesevertebrae apart from the shape of the vertebral foramen.

Te vertebral foramen in mastodons is more roundeddorsally, whereas that of the mammoth is triangular in shape(Olsen, 1972). Our third cervical specimen conforms tothe mastodon pattern, appearing more rounded (Pl. 6). Tecervical vertebrae of proboscideans in general are compressedanteroposteriorly, giving them a disk-like appearance (Olsen,1972). Te cervicals of our specimen are unfused, but Olsennoted that in both genera they can occasionally fuse together.

Toracic VertebraeSixteen thoracic vertebrae are present that have been identifiedas belonging to a single mastodon. Te third through the sixth

thoracics are present, followed by a gap in the sequence andanother sequence of 12 articulating vertebrae that are mostlikely the ninth through the twentieth thoracics. As mentionedabove, we have based our estimation on articulations withribs. Some of the ribs in the sequence are missing and some ofthe articular facets are missing on the middle ribs. Te shapeof these ribs was diffi cult to compare on both sides in thisarea because of several healed fractures on the right side. It ispossible that there were 19 thoracic vertebrae rather than 20if we have overestimated the number of ribs missing in thesequence.

Te mammoth from this site had at least 19 thoracicvertebrae. As mentioned above, we have a sequence of 15

articulating thoracics. Anterior and posterior thoracics aremissing, but we have anterior and posterior ribs, and havedetermined from this sequence that there were originally atleast 19 sets of ribs, and thus 19 thoracic vertebrae with whichthey articulated.

Te posterior thoracics in both animals incline to a greaterdegree than the anterior thoracics (Olsen, 1972), and those ofthe mammoth incline to an even greater degree (Pl. 8). Tespinous processes on these vertebrae are most massive toward

the anterior. Tey sharply increase in length at the third andfourth thoracics, which are located at the shoulder, and thendecrease more gradually toward the rear. Te longest, mostmassive spinous processes of the third and fourth thoracics

would have formed a hump behind the shoulders of theanimal. Te spinous processes of our mastodon specimenincrease slightly in length again in the most posterior thoracics.Tis would have created the slightly ‘dished’ spinal profile thatis seen in other mastodon specimens as well as to a greaterdegree in Loxodonta (Haynes, 1991). Te spinous processesappear to decrease in length through the vertebral column ofour mammoth specimen.

Slight diagnostic differences in the thoracic vertebraeobserved here include a sharper spine on the anterior faceof the spinous process on the more anterior thoracicsof the mastodon. It is unclear if this is related to differentmuscle development in these two individuals, gender, or is

a characteristic difference between the genera. Te concavityon the posterior face of the spinous process is also deeper andmore defined in the mammoth in the anterior thoracics. Weare unable to compare the third thoracic vertebrae of the twoanimals, because this element is missing in our mammothspecimen. From a comparison of the fourth and fifth thoracicsof the two animals (see Pls 7-8), it can be seen that the neuralarch is higher and thinner in the mastodon, and lower and

wider in the mammoth. Te centra of these vertebrae are alsomore robust in the mammoth. Te spinous processes of thesevertebrae in our mammoth specimen are longer and inclineto a greater degree than in the mastodon. Tese 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 ismore rounded in the mastodon and “approaches a right angle

with the apex on the … sagittal plane in the mammoth.”Tis character was not observed on our mammoth specimen,

which appears more similar to the mastodon with a roundedform. Observed differences in the posterior thoracics are thesignificantly longer spinous process in the mammoth, whichinclines to a greater degree than that of the mastodon (alsoobserved by Olsen, 1972), indicating a more gradual slopein the lower back of the mammoth. Te transverse processesof these vertebrae also differ in their orientation. In the

mastodon, these project away from the centrum and areoriented more dorsally, whereas in the mammoth, they havea more lateral projection and are closer to the centrum. Inthe most posterior thoracics of the mammoth, the transverseprocess begins to orient more dorsally and to resemblethat of the mastodon. Te vertebral foramen appears morecompressed dorsoventrally in the mammoth. Te posteriorthoracics of our mammoth specimen also do not show welldefined facets for the ribs.


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Lumbar VertebraeTree lumbar vertebrae are present for our mastodonspecimen, the last of which is fused to the sacrum. Tevertebrae articulate in sequence with the other mastodon

vertebrae, but can also be recognized independently asmastodon by their rounded ventral margin (Pl. 11).

SacrumOlsen (1972) noted that there is little difference between thesacral and caudal vertebrae of mastodons and mammoths. Tesacrum is present for our mastodon specimen, and is fused tothe third lumbar vertebra. It articulates with the left and rightillium, but is not fused to them.

R IBS Although Olsen (1972) did not describe differences in theribs, some are noted here. Generally, the mammoth ribs

have a stronger arch at the proximal end than those of themastodon.

Te anterior ribs are not readily distinguishable. Te mostposterior ribs, however, are quite variable between the genera.

As seen in the nineteenth rib (Pl. 11), the posterior ribs inthe mastodon are much more short and broad, whereas themammoth ribs appear longer and thinner. Te facet on thetubercle of the rib is less pronounced in the mammoth andabsent in the middle and posterior ribs.

As discussed above, we believe that there were originally20 sets of ribs in the mastodon and 19 in the mammoth. Tiscount conforms with most other data, although there is some

uncertainty in the literature on the number of these elements(Haynes, 1991).

SCAPULA Te scapula has been recognized as one of the more importantbones in distinguishing between proboscidean taxa (Olsen,1972). Our collection has both the right and left scapulae of

Mammut americanum (Pl. 12), identified as such based onseveral diagnostic characteristics described by Olsen (1972).

Olsen (1972) described the midspinous process as positionedmidway along the scapular spine in Mammuthus primigenius and closer to the acromion in Mammut americanum. Tisprocess is broken in both scapulae, but the position of the

break is more proximal and nearer to the acromion process.Te vertebral border of the scapula is straight in both of ourspecimens, which Olsen (1972) noted is a characteristic ofthe mastodon, whereas that of the mammoth is concave. Tescapular neck is more constricted in M. primigenius than in

M. americanum according to Olsen (1972), but neither of ourspecimens shows a high degree of constriction in this area.Olsen (1972) described the glenoid cavity as more rectangularin the mammoth and more “irregular” in the mastodon. We

would describe the glenoid cavity as more oval in shape inour specimen, as well as in Olsen’s (1972) photographs, ratherthan as irregular. See able 2 for measurements.

PELVIS A complete pelvis, which we have identified as belonging toour mastodon specimen, is present in this assemblage (Pl. 13).Olsen (1972) did not describe differences in the pelvis of themastodon and mammoth, except in noting that the mastodonpelvis is larger. We have identified this element as mastodonbased on its articulation with the vertebral column and hind

limbs. Te pelvis is an important element for determininggender in many animals. Basic differences between male andfemale proboscidean pelves include a smaller pelvic aperturein males as well as a broader ilium.

Based on the size of the tusk (see Fisher, 2008) as wellas pelvic morphology, the mastodon appears be a male.Sexual dimorphism in pelves has been used to determinegender in Mammuthus and Loxodonta (Haynes, 1991; Lister& Agenbroad, 1994; Lister, 1996). Although no data forcomparison are available for Mammut (but see Fisher, 2008;Fisher et al., 2008), our measurements fall into the male rangeof variation for Mammuthus , as defined by Lister (1996). It canbe expected that measurements for Mammut would follow the

same pattern. Lister (1996) has shown that the most clearlydistinguishing measurement for the pelvis of Mammuthus isthe ratio between the maximum horizontal aperture width(see able 3, measurement 3) and the minimum width of theilial shaft taken perpendicular to the long axis of the ilium(able 3, measurement 5). Lister (1996) observed that thereis almost complete separation between males and females forthis value, with males having a ratio of less than 2.6-2.7 andfemales having a ratio greater than this value. Tis ratio for

able 2. Measurements (in mm) of Watkins Glen mastodon scapula(after Agenbroad, 1994).

Right Left

1 763 775

2 632 642

3 779 783

4 276 275

5 266 264

6 220 226

7 141 145



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the Watkins Glen Mastodon is 1.75 (apertural width = 41cm; ilial shaft width = 24 cm). If pelvic morphology follows asimilar pattern to that of Mammuthus , this ratio clearly places

our specimen in the male range.GENERAL COMMENS ON HE LIMB BONES

Te limb bones of Mammut are shorter and more robustthan those of Mammuthus (as well as modern elephants).Lengthening of the forelimbs might be an evolutionary trend

within the Elephantidae, whereas Mammut might retain themore primitive state. Kubiak (1982) suggested that this couldbe related to feeding adaptations. Mammoths and modernelephants are browsers, which requires them to reach upto higher tree branches. Haynes (1991) suggested that thedifference could also be related to locomotor effi ciency, tusksize, or some other selective pressure. ables 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 metricdata are not well documented for Mammut .

HumerusOur collection contains one complete left humerus that hasbeen identified as mastodon (Pl. 14). In Mammuthus , this boneappears longer and thinner, with a more gracefully slopingshaft (Olsen, 1972: 16; able 4). Our specimen is more robustand has a more prominent supracondyloid ridge, which arecharacteristics that Olsen (1972) attributed to Mammut . Abroken proximal portion of a right humerus is also present in

the collection and appears to be from the same individual.

RadiusTe radii of both genera are similar (Olson, 1972: 16; able5). As with most of the other long bones, however, this bone ismore robust in the mastodon, especially at the distal end. Wehave identified this bone in our collection as mastodon basedon its articulation with the ulna (Pl. 15).

able 3. Measurements (in cm) of Watkins Glen mastodon pelvis (after Lister, 1996).

1 1592 42

3 41

4 65

5 24

able 4. Measurements (in mm) of Watkins Glen mastodon humer-us (after Agenbroad, 1994).

Right Left

1 - 937

2 - 599

3 - 916

4 - 132

5 267 264

6 - 231

able 5. Measurements (in mm) of Watkins Glen mastodon radius (after Agenbroad, 1994).

1 69

2 50

3 120

4 189

able 6. Measurements (in mm) of Watkins Glen mastodon ulna (after Agenbroad, 1994).

1 782

2 564

3 109

4 635

5 150

6 350


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Ulna Te ulnae of both animals are quite similar as well, and weredescribed in detail by Olsen (1972). Te ulna of the mastodonhas a slightly more developed olecranon process and deeper

semilunar notch, which is apparent in our specimen fromcomparisons of photos (able 6; Pl. 15). Olsen (1972) alsodescribed the mastodon as having a greater overhang of theolecranon process at the proximal end. Tis feature is diffi cultto observe without comparative material.

FemurTe femur is quite similar in both animals, but the mastodonfemur is somewhat shorter and thicker than that in themammoth. Te distal condyles appear more expanded mediallyand laterally in the mastodon. Olsen (1972) noted that thisis a subtle difference, but it is quite distinctly observed in ourspecimen, 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 alsosexually dimorphic. A comparison of the femur of the WatkinsGlen specimen (which has been determined to be male) andthe North Java mastodon specimen (PRI 49618; which islikely female) appears to follow this pattern (Pl. 16).

Modern adult male African elephants have femoral andhumeral lengths ca. 17-25% longer than those of females(Haynes, 1991). Tis dimorphism is also seen in fossilproboscidean bones, and is presumed to represent size

differences between the sexes. Te Berelyokh mammothbones at the Zoological Institute of the Academy of Sciencesin Saint Petersburg have a 14-26% difference in femur length(Haynes 1991). Haynes (1991) also reported that the Denvermastodon (on display at the Milwaukee Public Museum) andthe mastodon from the John Neath site (in the collections ofthe University of Wisconsin, Madison, Museum of Zoology)have a difference in limb bone length of 15-27%. Tisdifference in size has been observed in other mastodon limbbones as well (Haynes, 1991). Te femur of our Watkins Glenmastodon is 13.16% longer than the femur of our North Javaspecimen (Hodgson et al., 2008; able 7). Te sex of theNorth Java specimen has not been firmly established, but has

been determined to likely represent a female based on this sizedifference (Fisher, 2008). Tis percentage, however, is on theborder of differences that have been observed.

Patella

Te right patella of both specimens is present in our collection(Pl. 17). Te mammoth patella has been heavily gnawed bya carnivore, but the diagnostic features are still visible. Olsen(1972: 11) noted that the mastodon patella has “an expanded

lip on the margin of the articular surface and a prominentapex,” whereas the mammoth patella lacks these features. Ourmastodon specimen displays the features described by Olsen(1972) for Mammut . Te caudal border of the mammothspecimen is missing due to carnivore gnawing, but the anteriorsurface does not appear to slope enough to form a lip. Inaddition, the articular facet on the mastodon appears concave,

whereas this face on the mammoth patella has a flatter surface.

Tis is to be expected because the condyles of the femur wherethey articulate are more developed in the mastodon.

able 7. Measurements (in mm) of Watkins Glen and North Javamastodon femora (after Agenbroad, 1994). *See Hodson et al.(2008).

Watkins Java*

1 1,075 950

2 160 146

3 389 317

4 275 240

5 160 144

6 156 132

able 8. Measurements (in mm) of Watkins Glen mastodon tibia (after Agenbroad, 1994).

1 662

2 1043 260

4 152

5 214



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ibia Te tibiae of Mammut and Mammuthus are diffi cult todistinguish (Olsen, 1972; able 8). No notable differencesare observed here, but the element has been photographed

for reference (Pl. 17). Tere is both a right and left tibia inour collection from the same individual. Te tibiae in ourcollection articulate with the rest of the mastodon skeleton.

Fibula A left mastodon fibula (Pl. 17) and a probable right mammothfibula are present in our collection. Te right fibula consistsonly of the shaft portion, and so is diffi cult to describe, but itappears to be from a different individual than the left fibula.Te left fibula appears to be mastodon based on its articulation

with the mastodon tibia. It also appears similar in form to thefibula of the North Java mastodon specimen (Hodgson et al.,2008; able 9). Olsen (1972: 22) described the mammoth

fibula as “considerably more elongated, narrower, and morepointed at the proximal end.” Tese differences are quitesubtle, however, and are diffi cult to recognize in the absenceof comparative material.

Fore Foot All foot bones in our collection have been identified onthe basis of their morphology and articulations. Te mostdiagnostic faces of the bones have been photographed.Te reader is referred to Olsen’s work (1972) for completeillustrations of all faces of these bones.

Pisiform.–Olsen (1972) noted there is little differencebetween the pisiforms of the mammoth and mastodon.Our assemblages has one complete right and one partial leftmammoth pisiform, identified on the basis of its articulation

with the other mammoth foot bones. None is present forthe Watkins Glen mastodon, although we do have a partialright pisiform from our North Java mastodon specimen forcomparison (Pl. 19). Slight differences between the genera

were noted in this bone. Te mastodon bone appears tobe a more broad, compact bone, whereas the mammothpisiform appears slightly longer and thinner. Te dorsomedialsurface around the articular facet is somewhat broader in themastodon.

Cuneiform.–Our assemblage contains the left mastodoncuneiform and the right mammoth cuneiform. Te lateralprojection of the cuneiform is somewhat longer in the

mastodon. Dorsoventrally, the bone is somewhat thicker inthe mammoth (Pl. 20).

Lunar.–Te lunar is present from both the mastodon (leftside) and mammoth (right side) in our asssemblage. Olsen(1972: 22) noted that this bone, when viewed ventrally in themastodon “has a dished articular surface on the lateral face,

while this same surface in the mammoth forms a nearly rightangle with the straight articular face of the lateral surface.”

Although the margin of the lateral and anterior faces do forma near-right angle in the mammoth, this “dishing” was notobserved in our mastodon specimen, nor in the North Java

mastodon specimen. Rather, this margin in the mastodonlunar appears to form a more acute angle when viewedventrally. General dishing of the ventral articular surface itself

was observed in all specimens (Pl. 21).

Scaphoid.– A scaphoid from both our mastodon andmammoth specimens is present in our collection. Olsen(1972) noted no discernible differences between the two taxain this element, save for slight dorsoventral compression inthe mastodon when viewed laterally (or medially). We wereunable to observe this due to fracturing in our specimen.

Unciform.–Te left mastodon and right mammoth unciforms

are present in our collection. No notable differences areobserved between taxa in this element. We have photographedthem for identification purposes (Pl. 22).

rapezoid.–Te left mastodon and right mammoth trapezoidsare present in our collection. Te dorsal and ventral articularsurfaces are somewhat broader in that of the mastodon. Noother significant differences between taxa for this element

were observed (Pl. 23).

able 9. Measurements (in mm) of Watkins Glen and North Javamastodon fibulae (after Agenbroad, 1994).

Watkins Java

1 621 587

2 451 525

3 52 41

4 89 87

5 142 116


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rapezium.–Our collection contains a trapezium from bothour mastodon and mammoth specimens. Olsen (1972) notedconsiderable dorsoventral compression in the trapezium,

which is also observed on our specimens. Tis bone has

been photographed from all angles because of the significantvariation in shape between the species (Pl. 24).

Metacarpals.–Te metacarpals of proboscideans are uniquefrom those of other animals such as equids or bovids in thatthey are longer and more robust than the metatarsals. Tis isbecause the majority of the animal’s weight is carried by thefront limbs (Mol & Agenbroad, 1994). Te metacarpals of

Mammuthus appear longer than those in Mammut , whereasthose of Mammut appear a bit more robust. No significantmorphological differences on the proximal articular surfaces

were noted between the mammoth and mastodon, however,they were photographed here to aid in identification, because

these bones are often overlooked (Pl. 18).Mol & Agenbroad (1994) extensively described the

metapodials of Mammuthus columbi . Teir measurementsare reproduced below, along with the measurements of ourspecimens for comparison (see able 10).

Hind Foot Te hind foot of each genus is relatively smaller and morenarrow than the fore foot. As discussed above, this is becausethe front foot bears more of the animal’s weight. Somediagnostic elements that were preserved in our collection arediscussed below. Elements were identified to genus based on

their morphology and articulations.

Navicular.–Our collection contains both naviculars fromour mastodon and the right navicular of our mammoth

specimen. Te mastodon naviculars have been identified bytheir articulations with the mastodon astragali. Olsen (1972)did not note any differences between the naviculars of thetwo genera, but two subtle differences are observed here. In

the mastodon, this bone is thinner anteroposteriorly, andthe concave curvature on the ventral surface is deeper. In themammoth, this bone is thicker anteroposteriorly and lesscurved on the ventral surface (Pl. 25). Tese observationsappear to also hold true for Olson’s (1972: 40) publishedphotographs of the two genera as well as the navicular of theNorth Java mastodon (Hodgson et al., 2008).

Calcaneum.–Both the right and left calcanea of the mastodonare present and were identified on the basis of their articulations

with the right and left astragalus (Pl. 26). In addition, theypossess diagnostically mastodon characteristics. Olsen (1972:36) described the tuber calcis as expanded in the mastodon

and “postlike” in the mammoth. Our specimens display theexpanded tuber calcis. When viewed from the posterior, thesustentaculum is not oriented dorsally as in the mammoth,but more ventrally. Tis process is also notched on the ventralborder, which Olsen (1972) noted is a characteristic of themastodon calcaneum.

Astragalus.–Both the right and left astragali of the mastodonare present, and were identified by their articulations withthe right and left tibiae of the mastodon. A pronouncedcondyle is present on the posterior surface of the bone, whichOlsen (1972) noted is only present in the mastodon. Olsen

(1972) also noted that the bone is considerably compresseddorsoventrally in the mammoth and more expanded inthe mastodon when viewed anteriorly. More compression,however, is observed in our specimen than in Olsen’s (1972)

able 10. Comparative metacarpal measurements of Hot Springs Mammoth MSL 033 (HS), Watkins Glen Mammoth (MA) and Watkins GlenMastodon (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

Greatest length 123 n/d n/d 208 n/d 160 237 204 171 202 193 157 181 170 126

ransverse diameter:

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

Anteroposterior diameter:

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



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mastodon specimen (Pl. 27).

EPIPHYSEAL FUSION AND A GINGTere is some uncertainty in the literature on the lifespan of

modern elephants, but an upper age limit of 60-65 yr seemsto be most accepted (Haynes, 1991). Modern elephants havea long growth period, which extends into middle age (Lister,1994). Te timing of epiphyseal fusion can been used as atool for determining individual animal ages. Roth (1984)compiled an epiphyseal fusion sequence for modern Africanand Asian elephants. Epiphyseal fusion states for elephantsseem to be regular, with minor differences between speciesand sexes (Lister, 1994). Haynes (1991) and Lister (1994)argued that epiphyseal fusion in fossil proboscideans appearsto follow tooth progression and wear in a similar pattern tothat of modern elephants, implying that life span was probablyalso similar in these fossil species.

Bone fusion in the Watkins Glen specimens is detailed inable 11 and is compared to the fusion schedules of Haynes(1991) and Lister (1994). Most bones with diagnosticepiphyses 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. Tis would place the individual at at least 20 yrof age, but more likely quite a bit older. Haynes (1991) notedthat fusion of centra to plates could start around the age of20, but continues into the 50s. Considering that the platesare approximately half fused, the animal was perhaps aroundthe age of 40. Evidence of arthritis in the ribs could place ita bit older.

Evidence of epiphyseal fusion in our mastodon specimenplace the animal approximately in its 40s or 50s. Te distalradius is fused, which probably occurs during the late 40s(Haynes, 1991). Also, the vertebral centra are all fused, whichHaynes (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 the50s. Based on this information, it appears that the animal wasprobably in its late 40s or early 50s .

able 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 f n/d

Proximal 25-32 > 29 (often unfusedinto late 30s)

f n/d

ibia

Distal ca. 19 (18-20) ca. 32? f n/d

Proximal 17-24 (probably > 20) 28-32 (possibly earlier) f n/d

Humerus

Distal < 19 (ca. 13-14) > 18 f n/d

Proximal fusing 19-26 ca. 32 (often unfusedinto late 30s)

f n/d

Radius/Ulna

Distal > 24 > 32 (probably late40s)

f/f n/d

Proximal fusing at 19 < 32 (probably by 20s) f/f n/d

Vertebral centra and plates some centra fused toarches, plates attached

middle vertebrae only (<19)

lengthy fusion all arches and centrafused

all centra fused toarches, 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


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mammoths and mastodons: reconstruction of migratory behaviorusing strontium isotope ratios. Geology , 27(5): 439-442.

King, J. E., & J. J. Saunders. 1984. Environmental insularity andthe extinction of the American mastodont. Pp 315-339, in:Quaternary Extinctions: a Prehistoric Revolution, P. S. Martin &

R. G. Klein (eds), University of Arizona Press, ucson, Arizona.Kubiak, H. 1972. Te skull of Mammut praetypicum (Proboscidea,

Mammalia) from the collection of the Jagiellonian University inCracow, Poland. Acta Zoologica Cracoviensia , 17: 305-324.

Kurtén, B., & E. Anderson. 1980. Pleistocene Mammals of North America . Columbia University Press, New York, 442 pp.

Lister, A. M. 1996. Sexual dimorphism in the mammoth pelvis: anaid to gender determination. Pp 254-259, in: Te Proboscidea:Evolution and Palaeoecology of Elephants and Teir Relatives , J.Shoshani & P. assy (eds), Oxford University Press, Oxford, U.K.

Lister, A. M. 1994. Skeletal associations and bone maturation inthe Hot Springs mammoths. Pp 253-268, in: Te Hot Springs

Mammoth Site: a Decade of Field and Laboratory Research inPaleontology, Geology, and Paleoecology , L. D. Agenbroad & J. I.Mead (eds), Fenske, Rapid City, South Dakota.

Lister, A. M., & L. D. Agenbroad. 1994. Gender determination ofthe Hot Springs mammoths. Pp 208-214, in: Te Hot Springs

Mammoth Site: a Decade of Field and Laboratory Research inPaleontology, Geology, and Paleoecology , L. D. Agenbroad & J. I.Mead (eds), Fenske, Rapid City, South Dakota.

Maglio, V. J. 1973. Origin and evolution of the Elephantidae.ransactions of the American Philosophical Society , new series,63(3), 149 pp.

Miller, N. 2008. Contemporary and prior environments of theHyde Park, New York, mastodon on the basis of associated plantmacrofossils. Pp 151-181, in: Mastodon Paleobiology, aphonomy,

and Paleoenvironment in the Late Pleistocene of New York State:Studies on the Hyde Park, Chemung, and North Java Sites , W. D. Allmon & P. L. Nester (eds), Palaeontographica Americana 61.

Mol, D., & L. D. Agenbroad. 1994. Metapodials and shoulderheight of Mammuthus columbi compared with Eurasian

Mammuthus species. Pp 224-252, in: Te Hot Springs Mammoth

Site: a Decade of Field and Laboratory Research in Paleontology,Geology, and Paleoecology , L. D. Agenbroad & J. I. Mead (eds),Fenske, Rapid City, South Dakota.

Olsen, S. J. 1972. Osteology for the archaeologist: the Americanmastodon and the woolly mammoth. Papers of the Peabody

Museum of Archaeology and Ethnology, Harvard University,Cambridge , 56(3): 1-47.

Osborn, H. F. 1921. First appearance of the true mastodon in America. American Museum Novitates , 10: 4-6.

Osborn, H. F. 1936. Proboscidea: a Monograph of the Discovery,Evolution, Migration and Extinction of the Mastodonts andElephants of the World, vol. 1. American Museum Press, New

York, pp. 1-802.Osborn, H. F. 1942. Proboscidea: a Monograph of the Discovery,

Evolution, Migration and Extinction of the Mastodonts andElephants of the World, vol. 2 . American Museum Press, New

York, pp. 805-1676.Robinson, G. S., L. P. Burney, & D. A. Burney. 2005. Landscape

paleoecology and megafaunal extinction in southeastern New York State. Ecological Monographs , 75(3): 295-315.

Roth, V. L. 1984. How elephants grow: heterochrony and calibrationof developmental stages in some living and fossil species. Journalof Vertebrate Paleontology , 4(1): 126-145.

Saunders, J. J. 1996. North American Mammutidae. Pp 271-279, in:in: Te Proboscidea: Evolution and Palaeoecology of Elephants andTeir Relatives , J. Shoshani & P. assy (eds), Oxford UniversityPress, Oxford, U. K.

Shoshani, J., & P. assy. 1996a. Appendices. Pp 349-390, in: TeProboscidea: Evolution and Palaeoecology of Elephants and TeirRelatives , J. Shoshani & P. assy (eds), Oxford University Press,Oxford, U. K.

Shoshani, J., & P. assy. 1996b. Summary, conclusions, and a glimpse

into the future. Pp 335-348, in: Te Proboscidea: Evolution andPalaeoecology of Elephants and Teir Relatives , J. Shoshani & P.assy (eds), Oxford University Press, Oxford, U. K.

Simpson, G. G. 1945. Te principles of classification and aclassification of mammals. Bulletin of the American Museum of

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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 Unidentified

Skull 1 - - 37 fragments

Mandible 1 1 (R 207, L 208) - -

Basihyoid 1 - - -

Stylohyoid 2 2 (259, 262) 1 (263) -

Tyrohyoid 2 2 (260, 261) 2 (124, 289, 290) -

Cervical vertebrae 7 5 (173, 230, 231, 233, 234, 236) - -

Toracic 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 4 - - 2 (107, 175)

Scapula (L/R) 2 2 (L 200, R 222) - -Humerus (L/R) 2 2 (R 185, L 204) - -

Radius (L/R) 2 1 (L 76) - -

Ulna (L/R) 2 1 (L 78) - -

Scaphoid (L/R) 2 1 (L 97) 1 (R 205) -

Lunar (L/R) 2 2 (L 218, R 224) 1 (R 174) -

Cuneiform (L/R) 2 1 (L 128) 1 (R 70) -

Pisiform (L/R) 2 - 2 (L 125, R 104) -

rapezium (L/R) 2 1 (L 130) 1 (R 123) -

rapezoid (L/R) 2 1 (L 129) 1 (R 176) -

Magnum (L/R) 2 - 1 (R 183) -

Unciform (L/R) 2 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 2 (L 206, R 297) - -

Femur 2 2 (L 296, R 203) - -

Patella 2 1 (R 98) 1 (R 164) -

ibia 2 2 (L 79, R 72) - -

Fibula 2 1 (L 81) 1 (R 96) -

Astragalus (L/R) 2 2 (L 177, R 122) - -

Calcaneum (L/R) 2 2 (L 227, R 37) - -

Navicular (L/R) 2 2 (L 33, R 132) 1 (R 105) -

Cuboid (L/R) 2 - - -

Internal cuneiform (L/R) 2 1 (R 165) - -

External cuneiform (L/R) 2 - 1 (R 134) -

Metatarsals 5 pairs 3 (L 223) 2 (115, 142) -

Phalanges (posterior) 28 - - -

usks 2 1 (303) - -

A 47, 102, 201, 212, 232, 238-240, 242-243, 246, 249-250, 252-253, 257, 293.B 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.



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Complete skeleton of the Watkins Glen mastodon (PRI 8829).


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Complete skeleton of the Watkins Glen mammoth (PRI 8830).



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P

Figure1-2 Mastodon mandible (PRI 8829, bones 207, 208).

1. Occlusal view.2. Lateral view of left side.


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320 P A, N.


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Figure1-2 Mastodon atlas (PRI 8829, bone 173).

1. Anterior view .2. Posterior view.



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P

Figure1-2 Mastodon axis (PRI 8829, bone 234).

1. Anterior view.2. Lateral view.


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Figure1-2 Mastodon third cervical vertebra (PRI 8829, bone 231).




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P

Figure1-2 Mastodon fourth thoracic vertebra (PRI 8829, bone 246).

1. Anterior view.2. Posterior view.

3-4 Mammoth fourth thoracic vertebra (PRI 8830, bones 235).3. Anterior view.4. Posterior view.


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329

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Figure1-3 Mastodon fifth thoracic vertebra (PRI 8829, bone 293).

1. Anterior view.2. Posterior view.3. Lateral view.

4-6 Mammoth fifth thoracic vertebra (PRI 8830, bone 202).4. Anterior view.5. Posterior view.6. Lateral view.



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Figure1-2 Mastodon ninth thoracic vertebra (PRI 8829, bone 249).


3-4 Mammoth ninth thoracic vertebra (PRI 8830, bones 30 and 48).1. Anterior view.2. Lateral view.


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332 P A, N.


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Figure1-2 Mastodon thirteenth thoracic vertebra (PRI 8829, bone 252).


3-4 Mammoth thirteenth thoracic vertebra (PRI 8830, bone 294).3. Anterior view.4. Lateral view.



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Figure1-2 Mastodon first lumbar vertebra (PRI 8829, bone 251).


3 Mastodon nineteenth right rib (PRI 8829, bone 38).4 Mammoth nineteenth right rib (PRI 8830, bone 60).


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Figure1-2 Mastodon right scapula (PRI 8829, bone 222).

1. Lateral view.2. Glenoid cavity.



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Mastodon pelvis (PRI 8829, bones 206, 297).


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340 P A, N.


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Figure1-3 Mastodon left humerus (PRI 8829, bone 204).

1. Anterior view.2. Posterior view.3. Proximal view.



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Figure1-2 Mastodon left radius (PRI 8829, bone 76).

1. Lateral view.2. Medial view.

3-4 Mastodon left ulna (PRI 8829, bone 78).3. Lateral view.4. Medial view.


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344 P A, N.


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Figure1-3 Mastodon left femur (PRI 8829, bone 203).

1. Anterior view.2. Posterior view.3. Distal view.

4-6 North Java mastodon right femur (PRI 49618, bone 141).4. Anterior view.5. Posterior view.6. Distal view.



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Figure1-2 Mastodon riht tibia (PRI 8829, bone 72).

1. Anterior view.2. Posterior view.

3-4 Mastodon left fibula (PRI 8829, bone 81).3. Medial view.4. Lateral view.

5-6 Mastodon right patella (PRI 8829, bone 98).5. Ventral view.6. Lateral view.

7-8 Mammoth right patella (PRI 8830, bone 164).7. Ventral view (incomplete due to carnivore gnawing).

8. Lateral view.


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348 P A, N.


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Figure1 Mastodon left fore foot (PRI 8829).2 Mastodon left metacarpals, proximal view (PRI 8829, bones 213, 226, 39, 68).3a-h 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. rapezoid (176). h. rapezium (123).

4 Mammoth right metacarpals, proximal view (PRI 8830, bones 84, 67, 80, 86).



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Figure1-3 Mammoth right pisiform (PRI 8830, bone 104).

1. Anterior view. 2. Medial view. 3. Lateral view.4-5 North Java mastodon left pisiform (PRI 49618, bone 39). 4. Medial view. 5. Lateral view.


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Figure1-4 Mastodon left cuneiform (PRI 8829, bone 128).

1. Anterior view. 2. Posterior view. 3. Dorsal view. 4. Ventral view.5-8 Mammoth right cuneiform (PRI 8830, bone 7). 5. Anterior view. 6. Posterior view. 7. Dorsal view. 8. Ventral view.



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Figure1-2 Mastodon left lunar (PRI 8829, bone 218).

1. Ventral view. 2. Medial view.3-4 Mammoth right lunar (PRI 8830, bone 174). 3. Ventral view. 4. Medial view.


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Figure1-3 Mastodon left unciform (PRI 8829, bone 106).

1. Dorsal view. 2. Ventral view. 3. Medial view.4-6 Mammoth right unciform (PRI 8830, bone 187). 4. Dorsal view. 5. Ventral view. 6. Medial view.



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Figure1-2 Mastodon left trapezoid (PRI 8829, bone 129).

1. Dorsal view. 2. Ventral view.3-4 Mammoth right trapezoid (PRI 8830, bone 176). 3. Dorsal view. 4. Ventral view.


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Figure1-6 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.7-12 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.



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Figure1-2 Mastodon right navicular (PRI 8829, bone 123).

1. Anterior view. 2. Dorsal view.3-4 Mammoth right navicular (PRI 8830, bone 105). 3. Anterior view. 4. Dorsal view.


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Figure1-4 Mastodon right calcaneum (PRI 8829, bone 37).

1. Anterior view. 2. Posterior view. 3. Lateral view. 4. Medial view.



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Figure1-4 Mastodon right astragalus (PRI 8829, bone 122).

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


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