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Increasing and decreasing functional area of

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Increasing and decreasing functional area
of the dentition (FAD) of Mammuthus
primigenius

Ulrike Anders & Wighart von

Koenigswald

Palontologische Zeitschrift
Scientific Contributions to
Palaeontology

ISSN 0031-0220

Palontol Z
DOI 10.1007/s12542-013-0168-2

1 23
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 Author's personal copy
Palaontol Z
DOI 10.1007/s12542-013-0168-2
RESEARCH PAPER
Increasing and decreasing functional area of the dentition (FAD)
of Mammuthus primigenius
Ulrike Anders Wighart von Koenigswald
Received: 18 September 2012 / Accepted: 15 January 2013
Springer-Verlag Berlin Heidelberg 2013
Abstract The occlusal morphology and continuous molar
replacement in elephants provide a very effective func-
tional area for grinding the biomass that is more or less
abrasive. Parts of two subsequent molars contribute to the
functional area of the dentition (FAD). The FAD size,
measured in cm2, is associated with age and body size. The
FAD stage is indicated by the specific teeth contributing to
the FAD and represents the individual age. This study
concentrates on Mammuthus primigenius and compares the
FAD stages, as derived from growth series, with the fossil
Elephas antiquus, as well as the extant Elephas maximus
and Loxodonta africana. During the life history of the taxa
studied, the functional area increases simultaneously with
an increase in body size, but decreases severely in senile
age stages. In some senile individuals, the FAD is only
about 2050 % of the mean area of an adult animal. The
reduction of the FAD beyond a specific size does not mean
an immediate starvation of the animal. The general con-
stitution of the individual and the resources of fat accu-
mulation earlier may support the animal for some time but
certainly not over a longer period. Nevertheless, the highly
reduced functional area was sufficient to keep the animal
alive despite its full adult body mass. A much larger FAD
Electronic supplementary material The online version of this
article (doi:10.1007/s12542-013-0168-2) contains supplementary
material, which is available to authorized users.
U. Anders (&)
W. von Koenigswald
Steinmann-Institut fur Geologie, Mineralogie und Palaontologie,
Rheinische Friedrich-Wilhelms-Universitat Bonn, Nussallee 8,
53115 Bonn, Germany
e-mail: uanders79@gmx.de
W. von Koenigswald
e-mail: koenigswald@uni-bonn.de
in all adult stages provides the energy requirements needed
for all additional life functions including competition and
reproduction.
Keywords Elephantidae
Dentition
Growth trajectory

Tooth wear
Life history
Kurzfassung Bei den Elefanten bilden die Kauflachen
der Molaren bei einem horizontalen Zahnwechsel eine
hochst effektive Reibflache zur Zerteilung der Nahrung. An
dieser Reibflache, die hier als funktionales Areal der
Dentition (FAD) bezeichnet wird, sind meist zwei aufein-
ander folgende Backenzahne beteiligt. Die Zahnpositionen,
die zur FAD beitragen, bestimmen das FAD-Stadium. Die
FAD-Groe (in cm2 gemessen) kann mit dem Alter und der
Korpergroe korreliert werden. Die Studie konzentriert
sich auf Mammuthus primigenius und vergleicht die
Ergebnisse mit dem fossilen Waldelefanten (Elephas
antiquus) sowie den rezenten Elefanten Asiens und Afrikas
(Elephas maximus und Loxodonta africana). Wahrend des
Lebens wachst die FAD zunachst entsprechend der
zunehmenden Korpergroe an, schrumpft aber im senilen
Alterstadium wieder. Bei einigen senilen Tieren erreicht
die FAD-Groe lediglich 2050 % der durchschnittlichen
adulten FAD-Groe ausgewachsener Tiere. Aber mit dieser
stark reduzierten FAD konnten sich die voll ausgew-
achsenen Tiere noch hinreichend ernahren, wobei auch
vorher angelegte Fettreserven und eine gute allgemeine
Kondition das Uberleben uber einen kurzen Zeitraum
hinweg sicherten. Die sehr viel groere FAD in den vor-
angegangenen, adulten Alterstufen lieferte die Energie fur
alle zusatzlichen Lebensfunktionen.
Schlusselworter Elephantidae
Bezahnung

Wachstumskurve
Zahnabrieb
Lebensgeschichte
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U. Anders, W. von Koenigswald

Introduction FAD is large enough to sustain the functioning of a fully

grown individual.
During the past 2 decades, numerous frozen mammoth Our study is based on a large number of mammoth
carcasses have been found in Siberia that contribute sig- mandibles representing all age stages of M. primigenius in
nificantly to our knowledge of the life history of Mam- Germany. Specimens come from the late Pleistocene and
muthus primigenius, BLUMENBACH, 1799 (Boeskorov most originated from fluviatile deposits. Nearly complete
and Mol 2004). The early stages of the animals life his- skeletal finds allow a correlation of the FAD size with the
tory have been illuminated by the discovery of various reconstructed body size. Additional data were compiled
baby mammoths (Vereshchagin and Mikhelson 1981; from the literature, especially focused on the Siberian
Maschenko et al. 2005; Rountrey et al. 2012). The more mammoth babies to cover the early stages of the animals
functional aspect of the life history of M. primigenius can life history. These data were compared with other fossil
be documented in its dentition, which is the focus of this and extant elephants, and all elephants show a similar
article. trend.
In Mammuthus, as in other elephants, six molars occur This preliminary study will draw attention to a func-
successively because of horizontal tooth replacement. The tional aspect, especially in the senile stage of the elephant
first three molariform teeth are milk teeth (dp2/DP2, dp3/ dentition, which has so far been neglected.
DP3, and dp4/DP4), and they are followed by three molars
(m1/M1, m2/M2, and m3/M3) (Osborn 1942; Saunders
1970; Kozawa et al. 2001). The milk teeth are not replaced Methodological approach and materials
by permanent premolars. In each ramus of the mandible,
for most of the time, parts of two successive teeth function The functional area of the dentition (FAD)
simultaneously (Laws 1966). But, for a short period of
time, a single tooth is functional when the posterior part of The functional area of the dentition (FAD) is defined by the
the anterior tooth is worn out or the anterior part of the abrasive facets on one or two subsequent molars (Fig. 1).
posterior one is not fully developed. The capacity of the Only in very juvenile stages are three teeth (dm2, dm3, and
chewing apparatus of M. primigenius can be assessed using the first lophs of the dm4) functional. Normally, only two
these parts of the successive functional teeth. This subsequent teeth contribute to the functional area. But a
approach is more significant than counting the lamellae in single tooth may be active only for a short time after the
an isolated tooth. Therefore, we observed, measured, and anterior tooth is expelled. The FAD does not include those
evaluated the variability of the functional area of the
dentition (FAD) in the lower jaw during the various stages
of the life history.
The tooth wear and successive appearance of molars are
used as a criterion to determine individuals age stages in
mammoths based on correlative age stages inferred from
living elephants (Laws 1966; Sikes 1966, 1967; Krumrey
and Buss 1968; Jachmann 1985; Roth and Shoshani 1988;
Louguet-Lefebre 2005; Louguet 2006). The preferred diet
of the mammoth included grasses, sedges, and thin tufts of
gramnoids (Guthrie 2001; van Geel et al. 2008, 2010), and
this abrasive biomass had to be processed by the molars
with their transversely oriented enamel ridges. Maglio
(1972) stressed the point that molar crests are slightly
angled between the upper and lower teeth; thus, the
shearing point migrates along the crest, conserving energy
and evening out the load (von Koenigswald 1980).
Fig. 1 Functional area of the dentition (FAD) in the lower jaw of
The functional area of the dentition (FAD) increases
Mammuthus primigenius, Upper Pleistocene, Oberrheingraben near
with increasing age in mammoths because of the higher Bensheim (Meng 1011/31a). On the left side, the reflecting parts of
energy demand required by the larger body size. However, m2 and m3 represent the functional area. The FAD size is marked on
there is an extreme discrepancy in senile individuals! After the right side. The mandible is in FAD stage 10. Note the scars at the
distal end of the FAD in both m3s oriented vertically to the enamel
reaching the maximal adult FAD size, the functional area
ridges. They indicate the movement of the jaw in the mesial direction
of the last molar decreases significantly, although the body and without inclination as shown in the included mastication compass
size remains constant. Thus, the question arises about what (von Koenigswald et al. 2012) (photo: U. Anders)

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Increasing and decreasing functional area

parts of teeth that are worn down to a dentine shelf without

enamel crests and those not yet functional teeth. The FAD
size cannot be measured from isolated teeth because the
part of the FAD attributed to the preceding or following
tooth is not available. Thus, only teeth that are preserved in
the jaws provide data to calculate the FAD. The worn last
molar (m3) is an exception when definitely no other tooth
is still functional. We concentrated on data collection from
lower jaws because they are more common than upper
dentitions and often much better preserved in the fossil
record.
Pathologies in Mammuthus molars have often been
described (e.g., Guenther 1955; Kubiak 1965; e.g., Roth
1989; Adam 1994), but such abnormalities were excluded Fig. 2 Comparison of the total length of the enamel ridges on the
occlusal surface and the functional area of the dentitionFAD size
from our study. (horizontal axis) and the functional length of the enamel ridges
One way to quantify the FAD is to count the lamellae or (vertical axis) in Loxodonta sp. The regression shows the strong
the number of enamel crests. The number of lamellae in correlation between these two measurements. Thus, the more easily
relation to a defined length in isolated teeth is often used as obtainable value of the area can be used for characterizing the FAD
size [outlines for teeth are modified from Thenius (1989)]
the length-lamellar quotient or lamellar frequency for
evaluating the systematic position (e.g., Maglio 1972;
Guenther 1991, 1994; Lister 2001; e.g., Louguet 2001;
Pevzner and Vangengeim 2001; Louguet-Lefebre 2005;
McDaniel Jr. and Jefferson 2006). But these numbers are
not useful in evaluating the FAD. The number of lamellae
does not represent the functional area because the length of
the enamel ridges differs in the anterior and the posterior
part of each tooth.
Therefore, from a functional point of view, the total
length of all crests in the FAD would be more informative.
Measuring all crests, however, is more labor and time
intensive than to measure the area surrounding all crests
that are functional. A pilot study of Loxodonta showed that
the area surrounding all crests correlates well with the total
length of all crests (Fig. 2). Therefore, this value is selected Fig. 3 Comparison of the FAD size in the right and left side of
for characterizing the functional area of the dentition and complete mandibles of Mammuthus primigenius (n = 133). Most
values in the FAD stages show only a slight asymmetry (\20 %). The
used for comparative purposes. asymmetry shows some higher values in stages 10 and 11 (between 20
and 40 %). There is a strong asymmetry in single individuals in the
Applied technical methods senile stage 12. One tooth may be missing totally as in a mandible
from Sevsk (arrow) (Maschenko et al. 2006, fig. 10)
FAD measurements were acquired from photos of the
cheek teeth in the mandible, taken in occlusal view. The (Fig. 4). Maschenko et al. (2006) described one mandible
functional area was measured using ImageJ, an image from Sevsk, Russia (PIN 4353-442), in which the right m3
processing and analysis program. The measured area bor- is missing. We used the mean of values for each side in
ders all functional lamellae that show wear (Fig. 1). Where cases of moderate to strong asymmetries ([20 %).
two subsequent teeth are in use, the area encloses the entire
functional range of both teeth. If present, the functional Quantification of individual age and body size
area of each side of the mandible was measured separately.
Thus, broken jaws were included in our sample. Comparing To determine the wear pattern in mammoth, the functional
the values of both mandible sides indicates some degree of area has to be associated with an individuals age. For the
wear asymmetry (Fig. 3). Young individuals are charac- fossil M. primigenius, no data are available about at which
terized by slight asymmetries (\20 %). In adult individu- age the various teeth were replaced. Therefore, several
als, tooth asymmetry may increase to about 40 %. In some authors have used estimated ages measured in years
senile animals, the asymmetry may be distinctly higher inferred from extant Loxodonta africana (Guenther 1955;

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Haynes 1991; Adam 1994; Maschenko et al. 2005, 2006).
We use an age measurement that is independent from years
and solely based on the tooth positions that are functional.
A method of representing individual dental age was
developed for animals with vertical tooth replacement
(Anders et al. 2011), but this method is not applicable to
dentitions with a horizontal tooth replacement. Therefore,
we developed a system of 12 stages of functional area
(FAD stages) based on the six teeth contributing to the
FAD (Table 1). In uneven numbered growth stages, only
one of the six teeth is in function. In the other growth
stages, two teeth are contributing to the functional area.
FAD stage 12 includes senile teeth where only the last one
third of the potential lamellae in the m3 is functional.
Fig. 4 Senile mandible of Mammuthus primigenius (SMNS
6717.6.8.74.2) showing significant asymmetrical wear (photo: U.
Anders)
Table 1 Definition of the functional stages (FAD stages) in Elephantidae according to the molars partially in wear
FAD stages Definition
1 Only dp2 functional
2 dp2 and dp3; both functional
3 Only dp3 functional
4 dp3 and dp4; both functional
5 Only dp4 functional
6 dp4 and m1; both functional
7 Only m1 functional
8 m1 and m2; both functional
9 Only m2 functional
10 m2 and m3; both functional
11 Only m3 functional
12 m3 with only a third of possible laminae left over
The FAD stages are compared to the life history of the extant taxa Loxodonta and Elephas
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U. Anders, W. von Koenigswald
These dental stages do not represent equal time periods
of the life history but are useful for comparative purposes.
The utmost stage of worn dentitions is mentioned for the
dwarfed Island species Mammuthus exilis from the Channel
Islands. Agenbroad (2001: 474) mentions two individuals
whose mandibular teeth were missing due to advanced
age and wear, yet the animals continued to masticate
against the jaw bone.
The FAD stages can be associated, to some degree, with
important phases of the animals life history. When com-
pared to Loxodonta africana, an animal of FAD stages 15
(respectively, an individual age of 0 to about 6 years (Laws
1966; Sukumar 2003; Metcalfe et al. 2010) (Table 2) is
nursed; thus, these individuals get nutrition from mothers
milk. Subsequently, FAD stages 6 and 7 (between 8 and
15 years as seen in Table 2) cover the time from weaning
until the onset of reproductive age (Perry 1953; Laws
1966). The animals reach FAD stage 8 (m1 and m2 func-
tional) at the adult age of about 20 years (Laws 1966) and
therefore at the reproductive phase. The adult FAD stages
last until FAD stage 11 and cover the longest life period,
ranging between 30 and 40 years (Table 2). In FAD stage
12, when the m3 is heavily worn to about one third of the
entire tooth, a senile age is reached. This age system can be
used for all members of the Elephantidae, as well as
Mammuthus.
The association of FAD size with body size can be made
using either the body weight or shoulder height of an
individual. The body weight of the various age classes is
well documented for the extant species (Laws 1966; Ange
et al. 2001) and shows a fairly large variability and the
influence of sexual dimorphism (Lee and Moss 1986;
Damuth and MacFadden 1990; Haynes 1990). The shoul-
der height is also reported for extant species at each growth
stage (Laws 1966; Hanks 1972; Lee and Moss 1995). The
Biological events
Birth
Weaning
Sexual maturity
Senile age
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Increasing and decreasing functional area

Table 2 Comparison of the FAD stages to various classical age stages of mammoths and living elephants defined by other authors
FAD stages Estimated ages in Age stages Age stages Estimated ages in Age groups
years years

Elephantidae Loxodonta africana Loxodonta africana Loxodonta Mammuthus Mammuthus

africana primigenius primigenius
This study Laws (1966) Laws (1966) Sikes (1967) Adam (1994) Maschenko et al.
(2006)
1 01/2 I, II I/1I/2 Group I
2 1 III I/3I/4 Group I
3 Group I
4 23 IV, V I/5II/10 Group I/II
5 46 VI, VII III/1III/3 Group II/III
6 813 VIII, IX, X III/4IV/1 Group III/IV
7 15 XI IV/2 Group IV
8 1826 XII, XIII, XIV,XV, XVI IV/3V/1 1416 Group IV
9 2830 XVII, XVIII V/2V/3 2530 Group IV
10 3245 XIX, XX, XXI, XXII, XXIII, V/4VI/3 3035 Group IV/V
XXIV
11 4755 XXV, XXVI, XXVII, XXVIII VI/4VI/10 3550 Group V
12 5760 XXIX, XXX VI/11VI/13 5055 Group V

body weight is not accessible for fossil individuals of
Mammuthus, but the shoulder height can be calculated
from the long bones (Ziegler 1994). Therefore, skeletons of
M. primigenius are of special interest where dentition data
are linked with measurements of the long bones. Although
the number of such occurrences is very limited in
Germany, the six skeletons listed by Ziegler (1994) are
sufficient for a general comparison of features.
The material
We investigated 217 mandibles or jaws of Mammuthus
primigenius from various localities in Germany, mostly of
late Pleistocene age and a few from the late Middle
Pleistocene (see Appendix). Most specimens were dragged
from sand and gravel fluvial deposits. A rich collection
from the Oberrheingraben between Karlsruhe and Darms-
tadt is housed in the Staatliche Museum fur Naturkunde
Stuttgart (SMNS), the Hessische Landesmuseum Darms-
tadt (HLMD), and the private collection of Frank Menger,
Gro-Rohrheim near Darmstadt (Meng). Other M. primi-
genius were studied in the Naturhistorische Museum Berlin
(MFN) and the Steinmann Institute of the University in
Bonn (STIPB). Data on the more complete Mammuthus
skeletons were taken from the literature (see Appendix
ESM), especially where the shoulder height was recon-
structed (Ziegler 1994). For comparison, material from the
fossil straight-tusked elephant Elephas antiquus was stud-
ied in the same collections. This species is attributed to
Palaeoloxodon, regarded as a separate genus or subgenus
of Elephas by some authors. Since taxonomy is not the
purpose of this article, we use the term Elephas antiquus.
Extant material of Elephas maximus and Loxodonta
africana was made available in the collections of the
Naturhistorisches Museum Berlin (MFN), the Zoological
Museum in Hamburg (ZMH), Forschungsinstitut und
Naturmuseum Senckenberg (SMF), and the Alexander
Koenig Museum in Bonn (ZFMK). Thus, sampling of
representative animal sizes was accomplished. All speci-
mens and measurements are given in the AppendixESM.
The functional area of the mammoth dentition (FAD
size) during the life history
Measurements of 217 mandibles of M. primigenius show
that the functional area of the dentition increases continu-
ously from stage 1 to stage 11 (Fig. 5). The highest FAD
size values were observed when the m2 and part of the m3
(FAD stage 10) were included or when the m3 was fully
exposed (FAD stage 11). The values of the FAD size drop
distinctly in FAD stage 12, when due to wear only the last
one third of the m3 lamellae is functional.
Based on the life history and dental development of
Loxodonta africana and Elephas maximus (Perry 1953;
Laws 1966; Sikes 1966, 1967; Laws 1969; Hanks 1972;
Jachmann 1985; Haynes 1991; Sukumar 2003; Shrader
et al. 2006; Metcalfe et al. 2010), four comparable age
groups are identified for M. primigenius. These are: (1)
baby mammoth until weaning (FAD stages 15), (2) indi-
viduals between weaning and reaching reproductive age
(FAD stages 67), (3) fully grown individuals (FAD stages
811), and (4) senile individuals (FAD stage 12).

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Fig. 5 The increasing and decreasing functional area of the dentition
(FAD size) of Mammuthus primigenius (vertical axis) correlated to
FAD stages 112 (horizontal axis); the whiskers show the high
variability in the late FAD stages. The boxes cover 95 % of the
values. The red bar in FAD stage 11 indicates the mean value that is
used as 100 % of adult FAD size
Fig. 6 Comparison of FAD size and shoulder height in Mammuthus
primigenius; the FAD sizes and shoulder heights are compared to the
100 % values that are reached in FAD stage 11 (see text); data for
shoulder height of M. primigenius were taken from the literature
(Vereshchagin and Mikhelson 1981; von Koenigswald 1989; Ziegler
1994; Mol et al. 2006; Kirillova et al. 2011). The widths of the
colored zones (gray shaded for FAD size, red shaded for shoulder
height) indicate the range of variation in the single FAD stages. For
the shoulder heights only a small number of individuals could be
included; therefore, variation for that value has to be regarded with
suspicion
The mean values of the FAD size and the adult shoulder
height attained during stage 11 are set as 100 % for ease of
visualization and further comparison (Figs. 5, 6). The results
show a continuous and parallel growth in both the FAD size
and shoulder height of Mammuthus (Fig. 6). The functional
area increases up to about 30 % of the mean maximum FAD
size until the end of nursing (about FAD stage 5). Once the
reproductive age (FAD stage 8) has been reached, the FAD is
about 60 % of the mean maximum FAD size (Fig. 6). At
that time, the body size is still increasing. Only when the
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U. Anders, W. von Koenigswald
individuals attain their full body size is the maximum FAD
size available (FAD stages 10 and 11).
For the comparison, the reports of more complete
skeletal material are more informative than those of iso-
lated jaws.
Individuals of Mammuthus primigenius until weaning
time (FAD stages 15) (Fig. 7)
Baby mammoth jaws are rare in river deposits because of
their fragile structure. They are better known from the
carcasses from the Siberian Permafrost, where complete
individuals are preserved.
Lyuba from the Yurubei River, Yamal Peninsula,
Russia (Kosintsev et al. 2010; Rountrey et al. 2012), rep-
resents the rare FAD stage 1. In this animal, the dp2 shows
slight wear, whereas the dp3 shows no wear at all. The
individuals age of about 30-35 days was calculated from
the dentine formation after the neonatal line (Rountrey
et al. 2012). The FAD size measured from Rountrey et al.
(2012; fig. 4) is 0.5 cm2 and is only 0.4 % of the mean
maximum area in the adult stages. Because juveniles
exclusively suckle at this age, it is not clear whether the
exposed dentine is due to tooth wear, a result of spalling, or
incomplete mineralization (Rountrey et al. 2012). How-
ever, the teeth in these early stages are only of minor
importance for the animals energy requirement, and later,
with progressing age, their functionality increases.
Dima, a baby mammoth discovered in 1977 in
northeastern Siberia in the Kolyma basin near the Kirgilj-
ach River, is regarded as a fairly young (Vereshchagin and
Mikhelson 1981). The shoulder height of this individual is
100 cm, and the FAD size can be reconstructed at about
10 cm2. That is only about 5 % of the mean maximum area
in an adult individual, while the shoulder height attains
about 30 % of the maximum adult height. But, with the
input of mothers milk, the energy needs of the juvenile can
be covered.
Hunermann (1985) figured the upper and lower dentition
of a neonate mammoth from Niederweningen near Zurich.
According to an old figure from A. Lang (in Hunermann
1985), the right lower jaw carries two teeth, the rooted dm2
and dm3. The dm2 may have wear facets. The dm3 shows
initial facets on the first three lophs, and these features
indicate an early FAD stage 2. The Dm2 and Dm3 are
present in the upper dentition, and slight facets are visible.
No correlated skeletal material is available to evaluate the
juveniles size. It is possible that this individual might have
been even slightly younger than Dima.
The Saalian site, Pfannerhal, in the Geiseltal near Halle
(Germany) provided two skeletons of Mammuthus
(Toepfer 1957). The younger one can be attributed to stage
5, representing a late stage of a suckling individual. Here,
 Author's personal copy
Increasing and decreasing functional area
Fig. 7 Juvenile mandibles (FAD stages 25) of Mammuthus prim-
igenius; a mandible in FAD stage 2 from Benshheim (Meng 1033);
b mandible in FAD stage 3 from Geinsheim (Meng 1038); c mandible
the FAD size has increased up to about 30 % of the
maximal mean adult value. This individual reached about
50 % of adult shoulder height.
The teeth of newborn individuals appear to have no
facets. In Loxodonta, the portion of non-milk nutrition
increases distinctly, but the suckling duration is consistent
(Sukumar 2003). The time of weaning is variable, lasting
from 4 to 9 years in extant elephants. At the end of suck-
ling, the FAD size in FAD stage 5 has increased up to 30 %
of the adult individuals. This value seems to be the minimal
FAD for an individual to become independent of mothers
milk and enough area to cover the energy demand by
ingestion of plant biomass.
FAD stages between weaning and the beginning
of reproductive age (FAD stages 6 and 7)
This age group is represented by a large number of lower
jaws (Fig. 8). The FAD size in the Late Upper Pleistocene
Mammuthus varies from 40 to 80 cm2 per side in FAD
in FAD stage 4 from Bensheim (Meng 1030); d mandible in FAD
stage 5 from Lampertheim (SMNS 6316.2.3.75.1); (scale 5 cm)
(photos: U. Anders)
stage 6 and between 35 and 125 cm2 in FAD stage 7. These
values are 4555 % of the mean maximum FAD size. We
assume that the shoulder height corresponds to about
200220 cm (6070 %) (Fig. 6). However, none of the
German skeletons represent these FAD stages.
After weaning, the animal has to gain its energy
exclusively from the intake of plant biomass, and the FAD
size has to be large enough to satisfy the nutritional
requirements. This stage occurs at the age of 1012 years
in extant elephants (Perry 1953).
Adult individuals of Mammuthus primigenius
(FAD stages 811)
The data for FAD stages 811 come from 153 mandibles of
adult individuals of the Upper Pleistocene Mammuthus
primigenius. In stage 8, the FAD size varies between 55
and 110 cm2; in stage 9, between 50 and 180 cm2; in stage
10, between 90 and 190 cm2; and in stage 11 between
60 and 230 cm2. The values for Middle Pleistocene
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Fig. 8 Mandibles of Mammuthus primigenius in puberty based on the
life history and dental development in extant elephants; a mandible in
FAD stage 6 from Otterstadt (SMNS 6616.7.7.79.1); b mandible in
individuals from Pfannerhall (n = 1) and Steinheim/Murr
(n = 1) fall in the same intervals.
The mammoth skeleton found at Ahlen, near Hamm in
Westfalen (Siegfried 1959), is an adult individual with m2
and part of a functional m3 (FAD stage 10). Unfortunately,
the m2 has been lost, but in comparison to other mandibles
with similar wear patterns on the third molar, the FAD size
is about 140 cm2 in each jaw. The shoulder height of this
individual was reconstructed at 320 cm (Ziegler 1994).
Thus, the Ahlen specimen represents an average FAD size
and body size for FAD stage 10 mammoths (Fig. 9).
The mammoth skeleton from LAa (Nord Pas de Calais,
France) (Pointer 1913) represents FAD stage 11 because
only the m3 is functional. However, the FAD size does not
cover the distal lophs (Fig. 9). It measures 240 cm2 on the
right side and about 236 cm2 on the left side; this represents
an extremely large FAD size, almost double the mean FAD
size in that growth stage (Fig. 9). Ziegler (1994) calculated
a shoulder height of 301 cm for this individual. Thus, the
tooth replacement patterns indicate a more mature indi-
vidual from Ahlen, although the shoulder height is smaller.
Another mammoth from the Saalian Pfannerhall site, in the
Geiseltal near Halle, represents a full adult (Toepfer 1957).
The m3 is fully exposed (FAD stage 11), and the FAD size
measures about 185 cm2 per side. This value lies above the
mean (130 %) for Weichselian mammoths (Fig. 9). However,
at 300 cm (Ziegler 1994), the shoulder height of this individual
is somewhat below the mean of Weichselian mammoths.
Although the FAD size is slightly smaller, the Pfannerhall
specimen is comparable to the mammoth from LAa.
The skeleton found in Siegsdorf near Rosenheim,
Bavaria, Germany, has both m3 teeth fully exposed (Heiig
and Bredow 1987; Ziegler 1994). The most anterior lophs
are worn out completely (Fig. 9). The FAD size is about
130 cm2 per side, just below the average maximum in FAD
123
U. Anders, W. von Koenigswald
FAD stage 7 from Altrip (SMNS 6516.4.10.65.1) (scale 5 cm) (photo:
U. Anders)
stage 11. This animal was very large with a shoulder height
of 360 cm (Ziegler 1994). The FAD dimensions reflect the
beginning of the FAD size decrease in late stage 11, despite
a consistency in shoulder height.
In the Borna mammoth, Sachsen (Felix 1912), this trend
is even more pronounced. In contrast to the original
description, we regard the teeth of the mandible as third
molars. They are heavily worn in the anterior part where a
dentine shelf has developed. There is no indication for a
following tooth at the distal end. Thus, the individual is a
full FAD stage 11, but not yet senile. The FAD size taken
from the published figures is about 70 cm2, which is only
about 50 % of the mean maximum FAD (Fig. 9). With a
shoulder height of 320 cm (Ziegler 1994), the Borna
individual had an average maximum body size as defined
in the current paper.
At the time full adult individuals of FAD stages 811
reach their maximum body size, the FAD size reaches its
maximum as well. However, the feature becomes slightly
reduced when the anterior part of the m3 is worn (Fig. 9).
Tooth asymmetry is moderate between the two sides of the
jaw when 49 mandibles were evaluated and attributed to
this age group (Fig. 3). Hence, there is high individual
variability that is partly related to the differences in sexes.
Sexual dimorphism in body size becomes conspicuous
when an individual reaches the reproductive age at about
20 years (Loxodonta africanaFAD stage 8) (Lee and
Moss 1986). Thus, males can easily obtain double the
weight of females (Hanks 1972).
Senile individuals of Mammuthus primigenius
(FAD stage 12)
While mandibles of full adult individuals are frequently
preserved, those of senile mammoths are very rare. The
 Author's personal copy
Increasing and decreasing functional area
Fig. 9 Comparison of shoulder
height and FAD size in adult
(FAD stages 10 and 11) and
senile (FAD stage 12)
Mammuthus primigenius
skeletons; data for shoulder
height were taken from the
literature (Felix 1912; Pointer
1913; Toepfer 1957; Siegfried
1959; von Koenigswald 1989;
Ziegler 1994; Kirillova et al.
2011)
FAD size in a mandible of Mammuthus primigenius (SMNS
6717.6.8.74.2) from the Oberrheingraben, St. Leon near
Wiesloch, measures 10 cm2 on the left side and 38 cm2 on
the right side, and it exhibits a significant asymmetrical
shape (Fig. 4). Thus, the remaining FAD size of about 20 %
of the mean maximum area appears to have been enough for
processing the food required to maintain life.
Another left mandible of a senile Mammuthus primige-
nius was found in Nierstein in the northern Oberheingraben
(HLMD-RS 146) (von Koenigswald 1989). The m3 in this
individual is heavily worn and mostly represented by a
dentin shelf (Fig. 10a). Only the most basal parts of three
or four lamellae were functional, and these are strongly
reduced in their width. Thus, the remaining FAD size
measures only 15 cm2. However, asymmetries are frequent
in senile ages, and this extremely small value may have
been balanced by a more complete tooth on the other side.
The only skeleton attributed to this age group is the almost
complete mammoth from Polch, near Koblenz (von
Koenigswald 1989). The lower jaw holds both m3 teeth,
which are heavily worn (Fig. 10b). More than half of each
tooth is a dentine shelf. Thus, the FAD size is strongly
reduced and measures 64 cm2 on the right side and 75 cm2on
the left side. Both teeth are worn symmetrically (Fig. 10b).
These values approximate 50 % of the mean maximal FAD
size (Fig. 9). The individuals shoulder height was calcu-
lated at 320 cm (Ziegler 1994) and represents an average
body size. The Polch skeleton demonstrates that the upper
M3s are not worn to the same degree as the lower ones
according to their later appearance and different structure.
Senile m3s are recognisable even as isolated teeth.
Becker et al. (2009) depict a right molar from the Va Tche
Tcha, Ajoie, Switzerland. This m3 is slightly less worn
than both m3s from the Polch specimen. The FAD size is
measured at about 75 cm2 (Becker et al. 2009; fig. 1).
The FAD size in senile mammoths is reduced to 50 % or
even more, and asymmetrical tooth wear can be expected
(Figs. 3, 4). Therefore, one-sided mandibles or isolated
teeth are more difficult to evaluate.
Nevertheless, these teeth with the strongly reduced FAD
in stage 12 were still able to process enough plant material
to keep the fully grown mammoth alive. The minimal
FAD, however, is close to the final limit, starvation. The
animal might have selected softer plants, if available, and
used up resources of fat and other tissues accumulated in
earlier times. But such resources certainly are not able to
expand the life span for a longer period. Therefore, the
minimal FAD roughly marks the lower limit of the dental
area that is required to sustain the basic needs of a fully
grown mammoth. Consequently, the much lager teeth
during FAD 911 are able to provide a major surplus of
energy. This additional energy is essential for all activities
beyond the basic requirements, e.g., larger migrations,
intraspecific competition, and reproduction.
123
 Author's personal copy
Fig. 10 Mammuthus primigenius mandibles of FAD stage 12. a Left
mandible with the heavily worn m3 of the mammoth from Nierstein
in the Oberrheingraben (HLMD-RS 146), Germany, with only 34
Fig. 11 Comparison of the functional dental area (FAD size) of all
taxaMammuthus primigenius, Elephas antiquus, Elephas maximus,
and Loxodonta sp
Comparison with other elephants
Elephas antiquus
In this preliminary study, our sample of Elephas antiquus
comprises 50 individuals. However, the very early stages
14 and senile stage 12 are not represented in this data set.
The available data indicate that FAD size increases steeply
during the early growth stages until FAD stage 8, and the
trajectory decreases up to stage 11 (Fig. 11). The expected
decrease in stage 12 is not represented in our limited
material. The FAD size is slightly larger in E. antiquus than
in the other taxa studied because of the animals larger
body size (Fig. 11). During the stages of reproduction
(FAD stage 8)as deduced from Loxodontathe FAD
123
U. Anders, W. von Koenigswald
lamellae left; b mandible of the senile mammoth from Polch near
Koblenz (STIPB M4669), Germany, with heavily worn m3 on both
sides (von Koenigswald 1989) (i10 cm) (photos: G. Oleschinski)
Fig. 12 Comparison of the percentages of the FAD sizes to the mean
maximum FAD (100 %) in Mammuthus primigenius, Elephas
antiquus, Elephas maximus, and Loxodonta sp.; the times of weaning
and sexual maturity are inferred based on the life history of
Loxodonta africana
size reaches almost 8090 % of the mean maximum area
(Fig. 12). The FAD size in this FAD stage 8 is distinctly
higher than in Mammuthus primigenius and more similar to
Loxodonta and Elephas maximus. More information on
E. antiquus will be obtained from further studies of the
skeletal remains of the locality Neumark-Nord, near Halle/
S (Palombo et al. 2010).
Elephas maximus
The data set for Elephas maximus is limited to 15 indi-
viduals per FAD stage, and FAD stage 3 is not represented.
The measurements show the same increase of FAD up to
50 % at the time of weaning in age group 5 (Fig. 12). The
 Author's personal copy
Increasing and decreasing functional area
mean maximum FAD size (100 %) is reached in stage 8
where the reproductive phase starts (Fig. 12). In contrast
with the other taxa, the FAD values do not increase during
stages 811. The functional dental area in the adult stages
is distinctly smaller than in other elephants (Fig. 11). The
sample demonstrates the extreme decrease in area during
the senile stage 12, reaching only 5060 % of the mean
maximum area (Fig. 12).
Loxodonta sp.
Loxodonta is a well-represented taxon with more than 130
jaws, mostly belonging to L. africana. Here, FAD size
increases from stage 1 to 8 (Fig. 11). Stage 11 is repre-
sented by only one individual, and, thus, the unusual
decrease in the trend should not be considered as the norm
because there is a high variability in Mammuthus as well as
in both Elephas species in this stage (Fig. 5). The decrease
in FAD associated with stage 12 is well documented.
Conclusions
All elephant species investigated for this study show a
simultaneous increase of body and FAD size during the
early stages of their life history that continues until sexual
maturity. In stage 11, the maximal FAD size is reached
when the full m3 is functional. All species show a drastic
decrease in FAD size during senile FAD stage 12. This
senile minimum lies between 40 and 60 % of the mean
maximum value found in the FAD size. A severe asym-
metry between both sides of the mandible may occur in this
stage. Thus, the m3 on one side may decrease to only 20 %
of its former size or even may be lost completely.
Slight differences can be observed when the four studied
species are compared. Mammuthus differs by exhibiting
comparatively low FAD sizes during stage 8 when the other
species reach their reproductive age. Guthrie (2001) stressed
that the life history of Mammuthus may have been slightly
different from extant elephants of the tropical zones. Mam-
muthus reaches the highest values for FAD sizes during stage
10 and 11, which is similar to the other elephants.
The reduced FAD size during the senile age raises the
question about how large the functional area has to be to
support a fully grown animal. Our investigation shows that
the minimal area of a senile animal is similar to that prior
to sexual maturity. At this stage, about 60 % of the max-
imal shoulder height is gained in Mammuthus primigenius,
like in Loxodonta. Hence, in FAD stages 911, the FAD
provides a surplus to the immediate needs of the individual
to sustain body function. This surplus of at least one third is
required to satisfy the various additional functions,
including intraspecific competition and reproduction. Such
a surplus allows for the accumulation of reserves to outlive
critical environmental periods.
The rarity of preserved senile individuals of Mammuthus
primigenius in the fossil record may be caused by two
factors. One may be the result of a sampling bias, whereas
the second may be the rarity of old individuals in any
population. Heavily worn teeth may be less attractive to
collectors and thus be underrepresented in museum col-
lections. On the other hand, senile individuals are also rare
in stable populations of extant elephants. The peak mor-
tality is in young individuals (Haynes 1985, 1991) as fully
grown elephants are not endangered by predators. How-
ever, old elephants are more affected by harsh environ-
mental factors. Thus, only a few individuals survive until
senile age, and their preservation potential is low (Haynes
1991).
Acknowledgments This study was supported by the Deutsche
Forschungsgemeinschaft (DFG, German Research Foundation). It is
part of the DFG Research Unit 771 Function and performance
enhancement in the mammalian dentitionphylogenetic and onto-
genetic impact on the masticatory apparatus and publication no. 49.
We are much obliged for the help of the curators who allowed us to
work on the fossil and extant elephant material in their collections.
We want to name especially Dr. F. Mayer and Dr. O. Hampe, MFN
Berlin; Dr. R. Hutterer, ZMFK Bonn; K. Krohmann, SMF Frankfurt
a. M.; Dr. E. Schulz, ZMH Hamburg; F. Menger, Gro-Rohrheim; Dr.
R. Ziegler, SMNS Stuttgart. G. Oleschinki, STIPB Bonn, supported
us with photos. We are deeply indebted to Prof. R. A. Gastaldo, Colby
College, who revised the English manuscript draft. We are grateful to
Daniel Fischer and Maria Palombo for their constructive review.
References
Adam, K.D. 1994. Anomalien des Zahnwechsels bei Elephas
primigenius aus dem Quartar des Oberrheins. Stuttgarter Beit-
rage zur Naturkunde Serie B 211: 136.
Agenbroad, L.D. 2001. Channel Islands (USA) pygmy mammoths
(Mammuthus exilis) compared and contrasted with M. columbi,
their continental ancestral stock. In The world of elephants
international congress. Rome.
Anders, U., W. von Koenigswald, I. Ruf, and B.H. Smith. 2011.
Generalized individual dental age stages for fossil and extant
placental mammals. Palaontologische Zeitschrift 85(3): 321339.
Ange, K., S.D. Crissey, C. Doyle, K. Lance, and H. Hintz. 2001. A
survey of African (Loxodonta africana) and Asian (Elephas
maximus) elephant diets and measured body dimensions com-
pared to their estimated nutrient requirements. In Proceedings of
the American Zoo and Aquarium Association (AZA) Nutrition
Advisory Group. Fourth conference on zoo and wildlife nutri-
tion, eds. M.S. Edwards, K.J. Lisi, M.L. Schlegel, and R.E. Bray.
Lake Buena Vista, Florida: The AZA Nutrition Advisory Group.
Becker, D., D. Aubry, and J. Detray. 2009. Les dolines du Pleistocene
superieur de la combe de Va Tche Tcha (Ajoie, Suisse): Un
piege a restes de mammiferes et artefacts lithiques. Quaternaire
20(2): 135148.
Boeskorov, G.G., and D. Mol. 2004. Quaternary mammal collections
in the Museums of Yakutsk (Eastern Siberia, Yakutia, Russia).
Cranium 21(12): 1932.
123
 Author's personal copy
Damuth, J., and B.J. MacFadden. 1990. Body size in mammalian
paleobiology. Cambridge: Cambridge University Press.
Felix, J. 1912. Das Mammuth von Borna. Veroffentlichungen des
Stadtischen Museums fur Volkerkunde 4: 152.
Guenther, E.W. 1955. Mibildungen an den Backenzahnen diluvialer
Elefanten. Meyniana 4: 1236.
Guenther, E.W. 1991. Backenzahne eiszeitlicher Elefanten aus
Schottern des Oberrheintals, der weiteren Umgebung von
Offenburg (Baden). Quartar 41(42): 6385.
Guenther, E.W. 1994. Die Mammutfunde von Stuckenbruch bei
Herten. Geologie und Palaontologie in Westfalen 28: 128.
Guthrie, R.D. 2001. Reconstructions of Woolly Mammoth life history.
In The world of elephantsinternational congress. Rome.
Hanks, J. 1972. Growth of the African elephant (Loxodonta africana).
East African Wildlife Journal 10: 251272.
Haynes, G. 1985. Age profiles in elephant and mammoth bone
assemblages. Quaternary Research 24: 333345.
Haynes, G. 1990. The mountains that fell down: Life and death of
Heartland Mammoths. In Megafauna and man: Discovery of
Americas Heartland, eds. L.D. Agenbroad, J.I. Mead, and L.W.
Nelson, 2531. Hot Springs, South Dakota: The Mammoth Site
of Hot Springs, South Dakota, Inc., Scientific Papers.
Haynes, G. 1991. Mammoths, mastodonts, and elephants: Biology,
behavior, and the fossil record. Cambridge: Cambridge Univer-
sity Press.
Heiig, K., and B.R. Bredow. 1987. Das Mammut von Siegsdorf. Die
Fundgeschichte. In Der Eiszeit auf der SpurKatalog der
Mineralientage Munchen 1987 Achat. Eiszeit. Siegerland.,
134141. Freunde der Bayerischen Staatssammlung fur Palaon-
tologie und Historische Geologie.
Hunermann, A. 1985. Eiszeit-Saugetiere aus dem Kanton Zurich.
Vierteljahrschrift der naturforschenden Gesellschaft in Zurich
130: 229250.
Jachmann, H. 1985. Estimating age in African elephants. African
Journal of Ecology 23: 199202.
Kirillova, I.V., F.K. Shidlovskiy, and V.V. Titov. 2011. Kastykhtakh
mammoth from Taimyr (Russia). Quaternary International 30:
19.
Kosintsev, P.A., E.G. Lapteva, S.S. Trofimova, O.G. Zanina, A.N.
Tikhonov, and J. van der Plicht. 2010. The intestinal contents of
a Baby Woolly Mammoth (Mammuthus primigenius Blumen-
bach, 1799) from the Yuribey River (Yamal Peninsula). Doklady
Biological Sciences 432: 209211.
Kozawa, Y., H. Mishima, K. Suzuki, and M.W.J. Ferguson. 2001.
Dental formula of elephant by the development of teeth germ. In
The world of elephants, ed. G. Cavaretta, P. Gioia, M. Mussi, and
M.R. Palombo, 639642. Rome: Proceedings of the First
International Congress.
Kroll, W. 1991. Der Waldelefant von Crumstadt. Ein Beitrag zur
Osteologie des Waldelefanten, Elephas (Palaeoloxodon) antiqu-
us Falconer & Cautley (1847). Dissertation, Ludwig-Maximi-
lians-Universitat Munchen, Munchen.
Krumrey, W.A., and I.O. Buss. 1968. Age estimation, growth, and
relationships between body dimensions of the female African
elephant. Journal of Mammalogy 49(1): 2231.
Kubiak, H. 1965. The fossil elephants of South Poland. Folia
Quaternaria 19: 161.
Laws, R.M. 1966. Age criteria for the African elephant, Loxodonta a.
africana. East African Wildlife Journal 4: 137.
Laws, R.M. 1969. Aspects of reproduction in the African elephant,
Loxodonta africana. Journal of Reproduction and Fertility.
Supplement 6: 193217.
Lee, P.C., and C.J. Moss. 1986. Early maternal investment in male
and female African elephant calves. Behavioral Ecology and
Sociobiology 18: 353361.
123
U. Anders, W. von Koenigswald
Lee, P.C., and C.J. Moss. 1995. Statural growth in known-age African
elephants (Loxodonta africana). Journal of Zoology London 236:
2941.
Lister, A.M. 2001. Gradual evolution and molar scaling in the
evolution of the mammoth. In The world of elephants
international congress. Rome.
Louguet-Lefebre, S. 2005. Les megaherbivores (Elephantides et
Rhinocerotides) au Paleolithique moyen en Europe du Nord-
OuestPaleoecologie, taphonomie et aspects palethnographi-
ques. BAR S1451. Oxford: Archaeopress, Publishers of British
Archaeological Reports.
Louguet, S. 2001. The Middle and Late Pleistocene Mammoth
remains from Hanhoffen (Bas-Rhin, France). In The world of
elephantsinternational congress. Rome.
Louguet, S. 2006. Determining the age of death of proboscids and
rhinocerotids from dental attrition. In Recent advances in ageing
and sexing animal bones, ed. D. Ruscillo, 179188. Oxbow:
Oxbow Books.
Maglio, V.J. 1972. Evolution of mastication in the Elephantidae.
Evolution 26: 638658.
Maschenko, E.N., S.S. Gablina, A.S. Tesakov, and A.N. Simakova.
2006. The Sevsk woolly mammoth (Mammuthus primigenius)
site in Russia: Taphonomic, biological and behavioral interpre-
tations. Quaternary International 142143: 147165.
Maschenko, E.N., A.N. Tikhonov, and R.D.E. MacPhee. 2005.
Mammoth calf from Bolshoi Lyakhovskii Island (New Siberian
Islands, Arctic Siberia). Russian Journal of Theriology 4(1):
7988.
McDaniel Jr, G.E., and G.T. Jefferson. 2006. Dental variation in the
molars of Mammuthus columbi var. M. Imperator (Proboscidea,
Elephantidae) from a Mathis graval quarry, southern Texas.
Quaternarty International 142143: 166177.
Meller, H. 2010. Elefantenreich. Halle (Saale): Landesamt fur
Denkmalpflege und Archaologie Sachsen-AnhaltLandesmuse-
um fur Vorgeschichte.
Metcalfe, J.Z., F.J. Longstaffe, and G.D. Zazula. 2010. Nursing,
weaning, and tooth development in woolly mammoths from Old
Crow, Yukon, Canada: Implications for Pleistocene extinctions.
Palaeogeography, Palaeoclimatology, Palaeoecology 298:
257270.
Mol, D., J. Shoshani, A.N. Tikhonov, B. van Geel, S. Sano, P.
Lazarev, G.G. Boeskorov, and L.D. Agenbroad. 2006. The
Yukagir Mammoth: Brief history, 14C dates, individual age,
gender, size, physical and environmental conditions and storage.
Scientific Annals, School of Geology Aristotle University of
Thessaloniki 98: 299314.
Osborn, H.F. 1942. Proboscidea. New York: American Museum
Press.
Palombo, M.R., E. Albayrak, and F. Marano. 2010. The straight-
tusked elephants from Neumark-Nord. A glance into a lost
world. In ElefantenreichEine Fossilwelt in Europa, ed.
H. Meller, 219252. Halle: Landesamt fur Denkmalpflege und
Archaologie Sachsen-AnhaltLandesmuseum fur Vor-
geschichte Halle (Saale).
Perry, J.S. 1953. The reproduction of the African Elephant,
Loxodonta africana. Philosophical Transactions of the Royal
Society of London 237(643): 93149.
Pevzner, M.A., and E.A. Vangengeim. 2001. Age of some European
localities with elephant remains determined by the biometric
method. In The world of elephantsinternational congress.
Rome.
Pointer, G. 1913. Etude sur le Mammouth de lAa. Bulletin de la
Societe Prehistorique Francaise 10: 621646.
Roth, V.L. 1989. Fabricational noise in elephant dentitions. Paleo-
biology 15(2): 165179.
 Author's personal copy
Increasing and decreasing functional area
Roth, V.L., and J. Shoshani. 1988. Dental identification and age
determination in Elephas maximus. Journal of Zoology London
214: 567588.
Rountrey, A.N., D.C. Fisher, A.N. Tikhonov, P.A. Kosintsev, P.A.
Lazarev, G.G. Boeskorov, and B. Buigues. 2012. Early tooth
development, gestation, and season of birth in mammoths.
Quaternary International 255: 196205.
Saunders, J.J. 1970. The distribution and taxonomy of Mammuthus in
Arizona. Master Thesis, University of Arizona.
Shrader, A.M., S.M. Ferreira, M.E. McElveen, P.C. Lee, C.J. Moss,
and R.J. van Aarde. 2006. Growth and age determination of
African savanna elephants. Journal of Zoology 270(1): 4048.
Siegfried, P. 1959. Das Mammut von Ahlen Mammonteus primige-
nius (Blumenb.). Palaontologische Zeitschrift 33(3): 172184.
Sikes, S.K. 1966. The African elephant, Loxodonta africana: A field
method for the estimation of age. Journal of Zoology London
150: 279295.
Sikes, S.K. 1967. The African elephant, Loxodonta africana: A field
method for the estimation of age. Journal of Zoology London
154: 235248.
Sondaar, P.Y., and G.J. Boekschoten. 1967. Quaternary mammals in
the South Aegean Island arc; with notes on other fossil mammals
from the coastal regions of the Mediterranean. Koninkl. Nederl.
Akademie van WetenschappenAmsterdam; reprinting from
Proceedings, Series B 70(5): 556567.
Sukumar, R. 2003. The living elephants: Evolutionary ecology,
behaviour, and conservation. Oxford: Oxford University Press.
Thenius, E. 1989. Zahne und Gebi der Saugetiere. New York:
Walter de Gruyter.
Toepfer, V. 1957. Die Mammutfunde von Pfannerhall im Geiseltal.
Halle (Saale): Veroffentlichungen des Landesmuseums fur
Vorgeschichte in HalleVEB Max Niemeyer Verlag.
van Geel, B., A. Aptroot, C. Baittinger, H.H. Birks, I.D. Bull, H.B.
Cross, R.P. Evershed, et al. 2008. The ecological implications of
a Yakutian mammoths last meal. Quaternary Research 69:
361376.
van Geel, B., R.D. Guthrie, D.C. Fisher, J.G. Altmann, P. Broekens,
I.D. Bull, F.L. Gill, et al. 2010. Paleoecological studies of
mammoth and mastodon feces and the evidence for coprophagy.
Quaternaire, Hors serie 3: 9899.
Vereshchagin, N.K., and V.M. Mikhelson. 1981. Magadan baby
mammoth, Mammuthus primigenius (Blumenbach). Leningrad:
Nauka [in Russian].
von Koenigswald, W. 1980. Schmelzstruktur und Morphologie in den
Molaren der Arvicolidae (Rodentia). Abhandlungen der Senc-
kenberg Gesellschaft fur Naturforschung 539: 1129.
von Koenigswald, W. 1986. Ein Skelett vom Waldelefanten aus dem
jungpleistozanen Rheinschottern von Crumstadt bei Darmstadt.
Cranium 3(1): 813.
von Koenigswald, W. 1989. Das Mammut von Polch bei Mayen
(Eifel). Eiszeitalter und Gegenwart 39: 8797.
von Koenigswald, W., U. Anders, S. Engels, J.A. Schultz, and O.
Kullmer. 2012. Jaw movement in fossil mammals: analysis,
description and visualization. Palaontologische Zeitschrift 87:
141159.
Ziegler, R. 1994. Das Mammut (Mammuthus primigenius Blumenbach)
von Siegsdorf bei Traunstein (Bayern) und seine Begleitfauna.
Munchner Geowissenschaftliche Abhandlungen 26: 4980.
123