Proboscidea

Incertae sedis: Moeritherium

Plesielephantiformes

Family Numidotheriidae: Phosphatotherium, Daouitherium, Numidotherium

Family Barytheriidae: Barytherium
Family Deinotheriidae: Chilgatherium, Deinotherium (incl. Prodeinotherium)

Elephantiformes

Incertae sedis: Hemimastodon, Erythreum

Family Palaeomastodontidae: Palaeomastodon
Family Phiomiidae: Phiomia

Superfamily Mammutoidea

Family Mammutidae: Eozygodon, Zygolophodon, Mammut

Superfamily Gomphotherioidea

Family Gomphotheriidae

Choerlophodontinae: Choerolophodon
Gomphotheriinae: Gomphotherium
Amebelodontinae: Archaeobelodon, Protanancus, Serbelodon, Amebelodon, Platybelodon
Rhynchotheriinae: Rhynchotherium, Cuvieronius, Stegomastodon, Gnathabelodon,
Eubelodon, Sinomastodon

Superfamily Elephantoidea

Incertae sedis: Tetralophodon, Anancus, Stegotetrabelodon

Family Stegodontidae: Stegolophodon, Stegodon, Selenotherium
Family Elephantidae: Primelephas, Loxodonta, Elephas (incl. subgenus Palaeoloxodon),
Mammuthus

Tab. 1
Classification of the Proboscidea to genus level. Modified, mostly
simplified, after Shoshani/Tassy 2005, and taking an analysis by
Prado/Alberdi 2008 into account. Order, superfamily (upper case,
and with suffix -oidea), family (suffix -idae), subfamily (-inae), tribe
(-ini), genus (written in italics and upper case), subgenus (italics,
uppercase, and usually between brackets after genus), species
(italics) and subspecies (italics, after species name) are categories
at different levels.
Systematik der Proboscidea bis zur Gattungsebene. Abgeändert und
mehrheitlich vereinfacht nach Shoshani/Tassy 2005, unter Berücksich-
tigung der Analyse von Prado/Alberti 2008. Ordnung, übergeordnete
Familie (groß und mit Nachsilbe -oidea), Familie (Nachsilbe -idea),
Unterfamilie (Nachsilbe -inea), Stamm (-ini), Gattung (Kursivschrift und
groß), Untergattung (Kursivschrift, groß und gewöhnlich in Klammern
nach der Gattung), Art (Kursivschrift) und Unterart (in Kursivschrift,
nach dem Artnamen) sind Kategorien auf unter­schiedlichen Ebenen.

340 Jan van der Made // The evolution of the elephants and their relatives
One of the classic examples of evolution is that of the Pro-
boscidea (elephants and their relatives). It has been inten-
sively studied for a long time and is relatively well known.
These animals show a wide range of adaptations to the envi-
ronments in which they lived. They originated in a world
with a geography and climate that were very different from
the present, and they evolved while this world was being
transformed into the one we know today. This paper intends
to describe their evolution and distribution in relation to
these climatic and geographic changes.
Ein klassisches Beispiel der Evolution sind die Probosci-
dea (Elefanten und ihre Verwandten). Sie sind seit langem
erforscht worden und relativ gut bekannt. Diese Tiere zeigen
ein breites Spektrum der Anpassung an die Umwelt, in der
sie lebten. Ihren Usprung haben die Tiere in einer Umwelt
mit einer Geografie und einem Klima, welche sich deutlich
von der heutigen unterscheidet, und sie haben sich weiter-
entwickelt, während sich jene Umwelt in die wandelte, die
wir heute kennen. Dieser Beitrag möchte die Evolution und
Verteilung der Proboscidea auf der Erde im Kontext mit den
klimatischen und geografischen Veränderungen beschreiben.

The evolution of the elephants and their relatives

in the context of a changing climate and geography
Die Evolution der Elefanten und ihrer Verwandten im Kontext des Wandels von Klima und Geografie

Jan van der Made

Introduction
and played an important role in the evolution and dis-
It is the aim of this paper to describe the evolution tribution of the elephants.
and biogeography of the »elephants« as a context to the Fig. 1 gives an idea of the changed geography, while
elephants from Neumark-Nord. The living elephants Fig. 2–6 describe evolution and distribution of the ele-
are the Indian elephant (genus Elephas and species phants. As said before, there are over 15o fossil species
maximus) and African elephant (Loxodonta africana), recognized and many genera, but there is discussion
which belong to the family Elephantidae and for many on which species or genera are valid or not. It is not the
specialists the vernacular word »elephant« refers to intention to treat all species or even genera here and
this family. Alternatively the vernacular name could discuss details of nomenclature (Tab. 1). This overview
be applied to the order Proboscidea or »trunkers«, refer- of »elephant« evolution is more general when the older
ring to the trunk, which is the most typical character forms are concerned and becomes more detailed for
of these animals. As a comparison, other orders are groups that are more closely related to Elephas (Palaeo­
the Artiodactyla (even-toed ungulates), which includes loxodon) antiquus of Neumark-Nord.
among others the living families Suidae (pigs), Hip-
popotamidae (hippos), Cervidae (deer), Giraffidae The world of the earliest proboscideans
(giraffes), Bovidae (cattle, goats and antelopes), the During the Palaeogene (65–24 Ma, or Megaannum or
Perissodactyla (odd-toed ungulates) with horses, rhi- millions of years ago), the Red Sea did not exist and
noceroses and tapirs, and the Carnivora. Proboscidea Africa and Arabia formed one large continent. This
originated about 6o million years ago and belong to the landmass, like the Indian Subcontinent, was an iso-
Afrotheriidae, a large group of animals that all origi- lated landmass separated from Europe and Asia by a
nated from the African continent and which includes wide ocean, called the Tethys (Fig. 1). The level of the
also sirens, aardvarks, hyraxes, etc. Geography and oceans was then much higher than today (Fig. 3). These
climate have changed much in these 6o million years continents were then situated much more to the south

// Die Evolution der Elefanten und ihrer Verwandten 341


Atlantic
Erytherium – 60 Ma
Phosphatotherium – 55 Ma
Daouitherium – 55 Ma
Numidotherium – .. Ma
Barytherium – .. Ma
Moeritherium – 34 Ma
Palaeomastodon – 34 – 27 Ma Eozygodon – 22 Ma
Early Oligocene
Deinotherium – 16.5 Ma
Zygolophodon – 18 Ma
Gomphotherium – 18 Ma
WP
Mediterranean
Late Burdigalian
African taxa, indicating that Africa was still isolated at
that time (Kappelman et al. 2oo3).
Early Proboscidea
Paratethys
The Proboscidea originated in this isolated landmass
formed by Africa and Arabia with a tropical climate
and for many millions of years, they evolved there in
isolation.
Tethys
The earliest known species, called Erythreum azzour-
zorum, is from the Palaeocene (65–55 Ma). It was a
small animal with an estimated body weight of 3 to
8 kg. The lower premolars have one large main cusp
Phiomia – 34 – 27 Ma
and the uppers have a pair of two large cusps. The
Chilaghtherium – 30 Ma molars are bunodont (that is: with cusps, not with
Erythreum – 25 Ma
Indian Ocean
Gomphotherium – 27 Ma crests) and have two lobes each with a pair of main
cusps, and the third lower molar has an incipient third
lobe (Gheerbrant 2oo9).
The next oldest species is Phosphatotherium escuil-
lei (Fig. 3). It shows a tendency towards more lophodont
molars, which means that pairs of cusps that are next
? Platybelodon ≈ 15 Ma
Choreolophodon – 16.5 Ma to each other are fused into a smooth transverse crest
Zygolophodon – 16.5 Ma
Gomphotherium – 18 Ma
or loph (Gheerbrant et al. 1996). This evolution towards
lophodont teeth continued reaching perfect lophodonty
Eastern Paratethys ?
in Daouitherium, Numidotherium, Arcanotherium and
Barytherium (Gheerbrant et al. 2oo2; Mahboubi et al.
Deinotherium – 18 Ma
Me Zygolophodon – 20 Ma 1986; Delmer 2oo 9; Andrews 19o 6). Arcanotherium
sop Choerolophodon – 20 Ma
ota
mi Gomphotherium – 20 Ma
is based on a species which formerly was placed in
an
ba
sin Hemimastodon – 22 Ma Numidotherium. In Numidotherium the upper first and
third incisors are reduced in size and the second one
is large. There are two lower incisors; a large mesial
Indian Ocean one and a small lateral one. There is an upper canine,
but no lower one and there are three upper and lower
premolars. There is a long diastema (empty space) in
front of the cheek teeth. The skull is short and high,
which is an advanced condition. The preservation of
and moreover the global climate was much warmer so many incisors and a canine are primitive features.
than today (Lear et al. 2ooo). Living elephants have two upper tusks, which were
On this continent lived a fauna that was rather dif- believed to be second incisors. Many of the Miocene
ferent from that of the other continents and had cav- proboscideans have lower tusks, which usually were
iomorph rodents, creodonts (primitive carnivorous also believed to be second incisors. However, recently
animals), hyraxes of all sizes, primates, Embritopoda it has been proposed on the basis of material of Arcan-
(large animals related to elephants) and, as we have otherium, that the lower tusks are in fact the first lower
seen, the primitive relatives of the elephants (Maglio/ incisors.
Cooke 1978). During this time, there must have been Moeritherium is a strange animal, with a very elon-
some sporadic and not well understood connections gated body and very short legs, and some have doubts
with other continents. For instance, African primates whether it should be included in the Proboscidea,
and rodents reached South America and the hippo- though in cladistic analysis it is situated within that
potamus-like anthracotheres reached Africa. The latest group. The animal did not have a nasal cavity with a
Oligocene fossil record is not very dense, but a fauna large opening, suggesting that it did not have a trunk.
with an age of 27 Ma is rich in species, but all typically It has very small first lower incisors and large second
342 Jan van der Made // The evolution of the elephants and their relatives
lower incisors (Fig. 3), which gave rise to the interpre- mandible. In the most extreme way this is in elephants 1 (opposite/linke Seite)
tation that the lower tusks of the later proboscideans with short mandibles, very long molars and very heavy Early Oligocene paleogeography
and the proboscideans that
are second lower incisors. Now Moeritherium is inter- wear. In these animals milk teeth are worn of and shed remained in isolation in Africa
preted to be a form that evolved in a different direction before they are replaced by premolars, and the molars during the Palaeogene (with
and that did not give rise to the later proboscideans. move forward (like on a conveyor belt) as they are their approximate ages) (above)
and Late Burdigalian (Early
Its molars were bunodont and bilophodont, which worn down. There are usually not more than one or Miocene) paleogeography and
does not strictly refer to having lophs, but refers to two specimens in function in the same jaw. the proboscideans that dis-
the number of pairs of cusps or lobes. If it is said that persed out of Africa during the
so called »Proboscidean Datum
a species is trilophodont (or tetralophodont), this does Changing geography and the »Proboscidean Datum Event« Event« and the approximate
generally refer to the fourth premolar and the first two During the Palaeogene and earliest Miocene Africa ages of their dispersal (below);
molars (which usually, but not always have the same was isolated from Europe, Asia and the Indian Sub- maps after Rögl 1988, modified
(grey area = lack of information).
number). Later proboscideans may be tri-, tetra- or pen- continent by the Tethys. There must have been some Paläogeografie des Früholigozäns
talophodont, etc. The third molar generally has more sporadic faunal exchange, but the fauna remained in und Angabe der Rüsseltiere, die
lobes. In the case of Moeritherium, it had a third lobe large parts endemic. By a process called plate tecton- während des Paläozäns in Afrika
in Isolation verblieben sind (mit
(Andrews 19o6; Tassy 1981; Delmer 2oo9; Fig. 2). ics, these continents moved north, but the southern ihrem ungefähren Alter) (oben)
Palaeomastodon (Fig. 2; 3) and Phiomia (Fig. 2; 3) ones a little faster than the northern ones. The result und Paläogeografie des Spät­
are later forms. They are well known from very good is that the Tethys became narrower and the Alps and burdigals (frühes Miozän) und
die Rüsseltiere, die sich während
material from the Fayum Oasis in Egypt (e. g. Andrews Himalayas started to develop. North of this mountain des sogenannnten »Proboscidean
19o6; Osborn 1936). In these animals trilophodonty was chain a large sea developed, named the Paratethys. As Datum Event« von Afrika aus
acquired; all later forms are at least trilophodont, save this slow and gradual process continued, the separa- verbreiteten und die ungefähren
Altersangaben ihrer Verbreitung
for the deinotheres, which have first molars with three tion between Africa and Eurasia became narrower (unten); Karten nach Rögl 1988,
lobes and second molars with just one lobe. They have and shallower. At the same time, global temperatures geändert (graue Flächen =
reduced nasal bones and large nasal openings, indicat- were fluctuating and overall decreasing and extensive Fehlen von Informationen).

ing that they had trunks. They have long diastemas and glaciers started to develop on Antarctica. Since the ulti-
one pair of upper and one pair of lower incisors. These mate source of that ice is the water of the oceans, this
Oligocene genera have many similarities with some of process caused fluctuations of the level of the world’s
the Miocene proboscideans. oceans (left hand side of Fig. 3–5). The interaction of
Eritreum is known from mandible fragments of one tectonics and sea level caused fluctuating terrestrial
individual from Eritrea, with an age of about 27 Ma connections between Africa, India and Eurasia (e. g.
(Shoshani et al. 2oo6). It is of the size of Palaeomas- Fig. 1) resulting in an equal number of events of fau-
todon and Phiomia and has an estimated body weight nal exchange.
of some 5oo kg. The mandible has an elongatedd sym- These first faunal exchanges were called the »Pro-
physis and two incisors with a section that is higher boscidean Datum Event« (Madden/Van Couvering
than wide. The bunodont second molar has three lobes 1976) and had been believed to be one event. Later it
and the third one four. The buccal cusps have a tre- turned out that there were three (Thomas 1985) or at
foil pattern, which is better developed than in the two least six events (Van der Made 1996; 1999). Work in
older genera. The mandible is interpreted and shows the Bugti area and Zinda Pir dome is shedding light on
evidence for »horizontal« tooth replacement and Eri- early contacts between Africa, the Indian Subcontinent
treum is thus the most primitive proboscidean known and the rest of Eurasia. An elephantid proboscidean
in which this occurs. incisor was reported from strata with an age of about
The later and more evolved proboscideans have a 24 Ma, proving that faunal exchange started earlier
peculiar type of tooth replacement. Normal is »verti- than previously believed (Antoine et al. 2oo3; Lindsay
cal« tooth replacement, in which the milk molars are et al. 2oo5).
replaced by premolars, while the molars erupt behind Around or shortly after 22 Ma ago, numerous types
them and remain at their positions, so that in an adult of animals of Eurasian origins dispersed into Africa
individual all premolars and molars are in function. and the Indian Subcontinent: hedgehogs, a variety of
This is the case in Deinotherium and the primitive pro- rodents, lagomorphs (»hares and rabbits«), true carni-
boscideans. Horizontal tooth replacement is when the vores, chalicotheres (clawed rhinoceros-like animals),
teeth move forward in the mandible, worn teeth drop rhinoceroses, pigs and ruminants. Hyraxes, creodonts
out of the mouth and new teeth erupt at the back of the and at least one species of proboscidean dispersed from

// Die Evolution der Elefanten und ihrer Verwandten 343

Africa to the Indian Subcontinent, but the anthraco- 16,5 Ma in Europe, and did not disperse as far and not
there Brachyodus was possibly the only mammal that reaching northern and eastern Asia. In all areas where
dispersed to the north. For Africa this was a very deinotheres occur there is a size increase in time and
important event, comparable to similar events called the latest forms are really large (from this the name
Grand Coupure around the Eo-Oligocene transition, Deinotherium, meaning terrible mammal) and the ear-
when Europe came into contact with Asia, and the liest and smaller species of deinotheres are often placed
Great American Interchange, when a land connection in the genus Prodeinotherium, and the later ones in
between North and South America was established. Deinotherium. Since a genus can have only one origin,
After 2o Ma, and possibly close to 18 Ma, various this classification implies that the larger forms evolved
other mammals dispersed into Africa, while a creodont in one area and dispersed to others. However, that has
and the proboscideans Gomphotherium and Zygolo- not been proven.
phodon reached Europe and Gomphotherium reached Lophodont dentitions are believed to be an adapta-
also East Asia. Around 16,5 Ma, there was another tion to folivory (leaf eating). Lophodonty is when the
faunal exchange, and this time the proboscidean Dei- molars (and in some cases also premolars) have lophs,
notherium reached Europe, while Zygolophodon and which consist of a pair of cusps connected by a smooth
Choerolophodon may have reached Eastern Asia. Note- ridge. Carbon isotopes in tooth enamel give informa-
worthy is that at this time, the first primates of Afri- tion about the diet of a species and confirm that dei-
can origin may have reached the Indian Subcontinent notheres were folivores and remained to be so, when
and China. Some more of these events occurred, but environmental changes caused other herbivores to
were of less importance to proboscidean biogeography. include more grasses in their diets (Cerling et al. 1997).
Eventually progressive tectonics and sea level lowering The adherence to folivory may have lead to progres-
lead to the establishment of a permanent land con- sive extinction of the deinotheres at higher latitudes
nection and continuous or frequent opportunities of with a more seasonal climate and winters without the
faunal exchange. preferred food of these animals.

Deinotheriidae Hemimastodon

Deinotheres are the only proboscideans which lack
upper tusks, but instead they have lower tusks, which
appear from the mandible in downward direction
and then curve backward (Fig. 3). They are perfectly
lophodont. The first molar has three lobes, each with a
loph, but the second molar and third upper molar have
only two lobes, while the third lower molar has a very
small third lobe (Fig. 2). In Palaeomastodon a third lobe
starts to develop in the first and second molar, so Deino‑
therium must either have lost the third lobe in the sec-
ond molar, or evolved from a more primitive ances-
tor than Palaeomastodon. Deinotheres lost the first
two premolars, but still have all the permanent cheek
teeth at the same moment in function. This is normal
in mammals, but in the following proboscideans the
premolars and even the anterior molars tend to disap-
pear when the third molar is in function.
The earliest possible deinothere is Chilgatherium,
with an age of 27 Ma (Sanders et al. 2oo4), but the
material is very poor and many of the characteristics
of deinotheres cannot be confirmed here. During the
»Proboscidean Datum Event«, deinotheres dispersed
later than other proboscideans, appearing some time
before 18 Ma on the Indian Subcontinent and around
Hemimastodon is known by two upper molars from
the Indian Subcontinent, which had been assigned to
a large pig, but which in fact represent the most primi-
tive elephantoid (Tassy 1988). The exact stratigraphic
provenance of these remains are not known, but an
incisor with a morphology different from Phiomia and
older Proboscidea, but similar to that of the Elephan-
toidea is interpreted as Hemimastodon and comes from
strata with an age of about 24 Ma in the same area in
Pakistan (Antoine et al. 2oo3).
Mammutidae
Mammutidae are zygodont, which means that they
have molars with transverse crests, each one formed by
a pair of principal cusps connected by additional cusps
(but not by a smooth crest; Fig. 2). Like lophodonty,
zygodonty is believed to be an adaptation to folivory.
The first and second molars and the last milk tooth
had three lobes (trilophodont), while the last molar had
four lobes. All following proboscideans had this tooth
pattern or had molars with even more lobes. Contrary
to what happened in the deinotheres, this group had
large upper tusks, and initially also large lower tusks,

344 Jan van der Made // The evolution of the elephants and their relatives
 2

Mammuthus
The evolution of the morphology

Loxodonta
of the third lower molar (wisdom

Stegodon

Elephas
Stegomastodon warringi tooth). Occlusal surfaces are
shown of left molars (or mirror

Anancus

Stegotetrabelodon
0

Zygolophodon - Mammut
Pliocene Pleisto

Stegolophodon
images of right molars). Figures

Selenotherium

Primelephas
are not to scale. In the case of
Deinotherium, Zygolophodon,
Stegotetrabelodon, Primelephas
5 Amebelodon Anancus
and Elephas also the lingual side

Platybelodon / Amebelodon
Stegotetrabelodon is shown. (»occlusal« is the side

Choerolophodon
that during mastication is in

Paratetralopho-
Tetralophodon /
American gomphotheres
contact with the upper molar
and »lingual« is the side of

Gomphotherium
10
Deinotherium

the tongue). Conventions

don
Loxodonta africana
Tetralophodon as in Fig. 3.
Die Morphologieentwicklung des
Miocene

Zygolophodon dritten unteren Backenzahns

Primelephas (Weisheitszahn). Die okklusale
15
Gomphotherium angustidens
Oberflächen der linken Backen-
zähne (oder Spiegelbilder der
Choerolophodon Stegodon
rechten Backenzähne). Abbildun-
Stegolophodon Elephas hysudricus gen sind nicht maßstabsgetreu.
20 Deinotherium Im Falle von Deinotherium,
Gomphotherium cooperi Zygolophodon, Stegotetrabelodon,
Phiomia
Primelephas und Elephas wird
Palaeomastodon auch die linguale Seite gezeigt
(»okklusal« ist die Seite, die wäh-
Oligo

25 Moeritherium
rend des Kauvorgangs mit den
oberen Backenzähnen in Berüh-
rung kommt und »lingual« ist die
Seite zur Zunge hin). Angaben
wie in Abb. 3.

but in the course of evolution these became very small Upper and upper Middle Pleistocene. The American
or even absent (Fig. 3). Mammut was abundant and survived until the end of
The earliest genus of this group is Eozygodon from the Pleistocene and as a result they are well known.
Africa (around 22 Ma). During the first major faunal The stomach contents of Mammut included spruce need­
exchange of the »Proboscidean Datum Event«, Zygolo- les, pine cones and grass (!), occasionally gourd and
phodon spread to the Indian Subcontinent and during vine leaves (Lambert/Shoshani 1998).
a later faunal exchange, it spread to Europe, northern In northern Eurasia, the zygodont proboscideans
Asia and North America. In all these areas, the later became extinct around 2,6 Ma. This was probably
forms have better developed zygodonty and reduced related to a climatic change, when a glaciation occurred
lower tusks, but the transition is poorly documented. with ice accumulation on the northern continents,
In North America, this seems to have happened around and when climate became periodically more seasonal
11 Ma and the younger forms are placed in the genus (see section »The Great American Interchange«). This
Mammut (Lambert/Shoshani 1998). In Asia, this may increase in seasonality led to colder winters, which was
have happened at roughly the same time (Tobien et al. reflected in the vegetation: plants protect themselves
1988), but in Europe this seems to have happened later from cold winters by shedding their leaves and they do
and different authors place the species »borsoni« from not produce fruit. This led to the extinction in northern
northern Eurasia either in Mammut (which implies a Eurasia of folivorous and frugivorous animals, such as
dispersal from or to North America), or in Zygolopho- tapirs and the zygodont proboscideans. In North Amer-
don (which implies local evolution in the two areas). ica Mammut survived and even became very abundant.
The zygodont molars suggest that these probosci- The species became extinct during the late Quaternary
deans were folivores or at least no grazers. Balls of extinction event (see chapter »Megafauna extinctions
plant remains at the place of a stomach in an excavated in the wake of human dispersal«).
skeleton are commonly interpreted as stomach content.
This kind of preservation is mainly known from the

// Die Evolution der Elefanten und ihrer Verwandten 345

3 Africa S Asia Europe N Asia N America
Phylogeny and distribution of eustaticsea level (m)
the early proboscideans (black), –140 –100 –60 –20 20 60 100 140

Choerolophodon

Deinotherium
Deinotheriidae (green), Mam-

Deinotherium
0

M / Z borsoni

M / Z borsoni

Mammut
Zygolophodon
Plioc Plei
mutidae (blue) and Choerlopho- Great American

Choerolophodon
Interchange

Zygolophodon
don (red). On the left side, age
5
in millions of years, and eustatic »Camel event«
sea level after Miller et al.
Deinotherium
? ? ?
2005. The grey horizontal areas 10
»Hipparion event«

Zygolophodon

Zygolophodon
Deinotherium
indicate the temporal intervals in

»ProboscideanDatumEvent«
Miocene
which the so called »Probosci- 15 Choerolophodon
dean Datum Event« occurred, as

Palaeomastodon
well as several other interconti-

Chilgatherium

Eozygodon
nental dispersal events that were 20

Gomphotherium
Eritreum
Phiomia
allowed for by low sea level.
On the right the different pro- 25

Hemimastodon
boscideans per continent or part
Oligocene

Palaeomastodon

Moeritherium
of continent. Southern Asia cor-
30
responds to Asia south of the Mammut

Alpine-Himalayan mountain Moeritherium

chain and includes Southeast 35
Asia, while northern Asia corre-

Barytherium
sponds to the part north of that

Numidotherium
40
chain and includes southern Phiomia

China. Species or genera that Phosphatotherium

Eocene

occur only at the border of one 45 Daouitherium

of the areas are not included

Erytherium

in the graphs.
Thick lines indicate the approxi-
mate chronologic distribution of
the different Proboscidea. Thin
lines indicate simplified pre-
sumed phylogenetic relation-
ships mostly after Gheerbrant/
Tassy 2009 and Tassy 1990.
Dashed lines indicate a lesser
level of certainty. Arrows in the
thick lines indicate that full
stratigraphic range is not given.
Skulls of selected genera are
shown in right lateral view and
are not to scale; occasionally
mandibles are shown in
occlusal view.
50
Phosphatotherium
55
Faunal exchange between Asia and North America
the lower third molars, but may have five in the more
The opening of the Atlantic Ocean separated first advanced forms (Fig. 2). The milk teeth are changed,
South America from Africa and during the Eocene but the premolars are shed after a short use and in
North America from Europe. North America was rela- an adult gomphothere, only the molars are in use and
tively isolated, but it had sporadic contacts with Asia in an old one, only the posterior molars (»horizontal
across in Beringia (the area of the Bering Strait). These replacement«).
connections were in part controlled by sea level. Dur- Material as old as 27 Ma from Africa is assigned to
ing the Late Eocene and Oligocene (34–24 Ma), increas- Gomphotherium (Kappelman et al. 2oo3). The genus dis-
ingly colder conditions and the formation of extensive persed probably around 2o Ma ago into the Indian Sub-
glaciers on Antarctica caused decreasing sea levels and continent, around 18 Ma ago into Europe and 16.5 Ma
major faunal exchange event occurred between Europe, ago into northern and eastern Asia and North America.
Asia and North America, known as the Grande Cou- While in North America it seems to have lived on until
pure. Later more, but less dramatic faunal exchanges about 2 Ma ago, in the Old World it was replaced by
between North America and Asia occurred, some of Tetralophodon around 11–12 Ma ago. The latter genus
them being coincident with the »Proboscidean Datum generally is believed to be its descendant, but it is not
Event« and even bringing the first proboscideans to clear where this evolution occurred, though apparently
North America. intermediate forms are known from Europe.

Gomphotheriidae – Gomphotherium Gomphotheriidae – Choerolophodon

Gomphotherium has downward directed upper tusks The genus name Choerolophodon derives from the »pig-
and flattened lower tusks inserted in a very elongated like« teeth, which have additional cusps and crenulated
symphysis (Fig. 4). This morphology is already fore- enamel. They are trilophodont, while the third lower
shadowed by Phiomia (Fig. 3). It is trilophodont (three molar has four lobes (Fig. 2). The upper tusks are large,
lobes in the last milk tooth and the first two molars) while the lower incisors are reduced (Fig. 3). The man-
and the third molar has more lobes, initially four in dibular symphysis is elongated, which suggests that

346 Jan van der Made // The evolution of the elephants and their relatives
they evolved from a form with an elongated symphysis and unlike Stegotetrabelodon. Material from Sahabi, Phylogenese und Verteilung der
and incisors. described as Amebelodon cyrenaicus is probably con- frühen Rüsseltiere (schwarz),
Deinotherien (grün), Mammutidae
The oldest Choerolophodon known is from the specific with »M.« grandincisivus. (blau) und Choerlophodon (rot).
Indian Subcontinent and there is no record from Africa In the Old World, the shovel tuskers are mainly Auf der linken Seite, Altersanga-
of a similar or slightly younger age (Tassy 199o), sug- restricted to areas which probably had dry or open ben in Millionen von Jahren und
eustatischer Meeresspiegel nach
gesting the possibility that Choerolophodon may have landscapes. Platybelodon at Tongxin occurs in sedi- Miller et al. 2005. Die grauen
originated in the Indian Subcontinent from Gompho­ ments with gypsum crystals (Guan/Rice 199o), which horizontalen Felder markieren die
therium and subsequently dispersed into Africa. may have been formed by evaporation typical of an zeitlichen Intervalle, in denen das
sogenannte »Proboscidean Datum
The genus is known from Pikermi and Chios in arid climate. This contrasts with traditional views in Event« stattfand, sowie weitere
Greece, but did not disperse far into Europe (Tobien which these animals waded in ponds and marshes interkontinentale Verteilungsereig-
198o). Its geographic distribution suggests that it was and used their shovel-like incisors to collect food in nisse, die im Zuge der Senkung
des Meeresspiegels möglich
adapted to more or less open or dry environments. the mud. More recently, it has been suggested that the waren.
large incisors are used in conjunction with the trunk Auf der rechten Seite sind die
Gomphotheriidae – Shovel tuskers verschiedenen Rüsseltiere auf
to cut tough vegetation (Lambert/Shoshani 1998).
jedem Kontinent oder Teil eines
The Amebelodontinae or shovel tuskers are a subfamily Kontinents angegeben. Südasien
of the Gomphotheriidae. Their most typical character- Proboscidean dispersal and Mid Miocene bezieht sich auf Asien südlich
istic is the enlarged and widened lower incisors with Climatic Optimum der Alpen-Himalaja Bergkette
und schließt Südostasien mit ein,
tubular dentine, which have been compared to a shovel. As part of the »Proboscidean Datum Event« at least six während sich Nordasien auf den
They maintain small upper tusks, are trilophodont proboscideans dispersed out of Africa. But for a long Teil nördlich dieser Bergkette
and have third molars that are moderately elongated period after about 15 Ma no dispersals out of Africa bezieht und Südchina mit ein-
schließt. Arten und Gattungen,
(lower third molar with five or more lobes). occurred, though there was a land connection and die nur an der Grenze einer
Material that initially was considered to belong to at least four to five dispersals of proboscideans into Gegend auftreten, sind in dem
Gomphotherium, and which is very similar, was named Africa occurred. Why? Diagramm nicht angegeben.
Dicke Linien zeigen die ungefähre
Archaeobelodon. On the one hand, this genus, with Dispersals out of Africa are dispersals to higher chronologische Verteilung der ver-
a distribution in Africa and Europe, is however not latitudes (even if later the direction of dispersal is schiedenen Rüsseltiere an. Dünne
considered to have given rise to other shovel tuskers deflected towards the southeast). Higher latitudes tend Linien geben die vereinfachten
und angenommenen phyloge-
(Tassy 199o). On the other hand, the validity of the to have more seasonal climates. This may restrict the netischen Verwandtschaftsbe-
genus and the relationship of these specimens with availability of food for herbivores. Herbivores tend to ziehungen in der Mehrheit nach
Gheerbrandt/Tassy 2009 und
the shovel tuskers was doubted (Mazo 1996). The next adapt to this to such an extent that procreation occurs
Tassy 1990 wieder. Gestrichelte
most primitive genus is Protanancus, which dispersed in conjunction with the seasons. There are thus prob- Linien zeigen einen geringeren
into the Indian Subcontinent, but at the same time it lems to overcome for a species dispersing northward. Grad an Bestimmtheit an. Pfeile
in den dicken Linien bedeuten,
may have dispersed into northern Asia, where it gave If global temperatures are higher, seasonality has less
dass die gesamte stratigraphische
rise to Platybelodon and shortly later into North Amer- effect on flora and fauna and therefore these problems Bandbreite nicht angegeben ist.
ica where it gave rise to Amebelodon and Serbelodon. for the northward disperser are minimized. Schädel von bestimmten Gattun-
gen sind in rechter Seitenansicht
The latter two genera are closely related and follow Most of the »Proboscidean Datum Event« coin-
abgebildet und nicht maßstabs-
each other in time. They are tetralophodont and have cided with the Mid-Miocene Climatic Optimum. As getreu. Zuweilen werden die
third lower molars with around five cusp pairs (Fig. 2). mentioned before, global temperatures were higher in Unterkiefer in okklusaler Ansicht
gezeigt.
Compared to Gomphotherium (Fig. 4), the mandibular the Eocene but started to decrease (Lear et al. 2ooo).
symphysis is also elongated, but wider to accommo- During the Middle Miocene, broadly between 16–17
date the much larger incisors (Fig. 5). Platybelodon is Ma and 12–14 Ma, there was a reversal of this trend,
similar, but has a shorter and more robust symphysis which ended with the so-called Mid-Miocene Crisis,
with extremely wide incisors. In America it evolved when temperatures dropped again. These generally
into Torynobelodon, which is similar, but with a still warm conditions did not impede that in certain areas
more robust symphysis. (such as Antarctica) temperatures fluctuated, reached
There has been some discussion on the affinities of low values and caused global sea level changes.
the species »Mastodon« grandincisivus (see discussion This climatic optimum is noted in the northward
by Tassy 1999). Some authors include a wide range of expansion of many marine and continental organisms.
material and assign the material to Stegotetrabelodon. Termophile reptiles and amphibians, such as monitor
However »M.« grandincisivus has large lower tusks lizards, chameleons and crocodiles dispersed to Europe,
(Fig. 5) with tubular dentine, as in the shovel tuskers but became extinct again after the climatic optimum

// Die Evolution der Elefanten und ihrer Verwandten 347

(Böhme 2oo3). The same pattern is observed in mam- they became extinct as part of the late Quaternary
mals. For instance, tragulids (mouse deer), which are extinction event.
frugivores, dispersed to Europe and for some time
Sinomastodon and the dispersal of Camels
were abundant and diverse, but after the climatic opti-
to the Old World
mum they became rarer and finally disappeared.
Proboscideans that reached northern Eurasia dur- Sinomastodon is a late gomphothere from China, with
ing the Mid-Miocene Climatic Optimum were prob- a mixture of primitive (bunodont, trilophodont) and
ably all browsers, but they evolved and adapted them- advanced characters: lack of lower incisors and a
selves to the environment and flourished during some short mandibular symphysis. An endemic species of
time. However, most became finally extinct in relation this genus is reported from the Early Pleistocene Satir
to climatic change. African proboscideans were not fauna from Java (Van den Bergh 1999).
capable of dispersing into higher latitudes, until they The primitive characters of Sinomastodon preclude
evolved different feeding strategies (Van der Made/ a close relationship with Tetralophodon or Anancus,
Mazo 2oo3). but this combination of characters is found in some
American gomphotheres (Tobien et al. 1988; Prado/
American gomphotheres
Alberdi 2oo8). This would imply a latest Miocene or
Gomphotherium dispersed to America, where it sur- Early Pliocene dispersal from America across the Ber-
vived much longer than in the Old World and gave ing Strait to East Asia.
rise to a number of genera (Lambert/Shoshani 1998; Camels originated during the Eocene in North
Prado/Alberdi 2oo8; Fig. 4). These are all forms with America and remained isolated in this continent until
low crowned trilophodont anterior molars and simple they dispersed across Beringia into the dry areas of
third molars that are not much elongatedd. Eubelodon Asia and Africa and even reached southern Spain (Van
and Gnathobelodon lost their lower tusks, but the sym- der Made et al. 2oo2). The dispersal of these camels
physis acquired a peculiar shape, suggesting that it was and of Sinomastodon may have occurred at the same
functional, even without incisors. Rhynchotherium is time.
rather similar to Gomphotherium, but differs in hav-
ing lower tusks that are laterally compressed and that Great American Interchange
may have enamel bands (Fig. 4). Stegomastodon lost its The opening of the Atlantic Ocean separated South
lower incisors, acquired a short symphysis, its skull or America from Africa during the Mesozoic (before 65
face became shortened (Fig. 4) and its molars became Ma), but it remained connected to Antarctica and Aus-
heavily wrinkled enamel with small additional cusps tralia. Marsupials flourished in this large continent. By
(Fig. 2). Cuvieronius shares with the previous genus the the Oligocene South America was disconnected from
reduction of the lower incisors and the shortening of Antarctica and became isolated save for some connec-
the symphysis and face, but differs in having very large tion with Africa, which must have occurred during
upper tusks with twisted enamel bands (Fig. 4). None the Eocene, allowing the placental caviomorph rodents
of these genera have molars with high crowns, but the and primates access to South America. Marsupials and
molars with wrinkled enamel and additional cusps of endemic placentals evolved during a long period of
Stegomastodon suggest that it became a grazer. Car- isolation and the South American fauna was totally
bon isotope analysis suggests that this genus was an different from that of any other continent, until a firm
opportunistic grazer, browser or a mixed feeder, while contact with North America occurred. There may have
Cuvieronius was a browser or mixed feeder (Sánchez been some sporadic contacts allowing few animals to
et al. 2oo4). disperse into South America, but around 2,6 Ma ago,
When the Great American Interchange occurred the connection in the area of the Panama Isthmus was
(see corresponding section below), Cuvieronius dis- established.
persed into South America. Stegomastodon did the The land connection between North and South
same in the Middle Pleistocene. Several of the gom- America was produced by tectonic processes, but prob-
photheres in North America became extinct around ably climate interfered as well. Around 2,6 Ma global
the time of the dispersal of Mammuthus in this conti- climate changed reaching still lower temperatures and
nent. Stegomastodon survived in South America and glaciations occurred on the northern hemisphere, caus-
Cuvieronius in both North and South America until ing ocean levels to drop (see left hand side of figs. 3–4).

348 Jan van der Made // The evolution of the elephants and their relatives
 Africa S Asia Europe N Asia N America S America
eustaticsea level (m)
–130–110–90 –70 –50 –30 –10 10 30 50

Anancus

Cuvieronius
Stegotetrabelodon

Anancus
0

Gomphotherium
Pliocene Pleisto

Rhynchotherium
Anancus

Stegomastodon

Stegomastodon
Sinomastodon

Cuvieronius
Gnathbeldon
Paratetra-

Paratetralophodon

Tetralophodon
lophodon

Tetralophodon
Tetralophodon

Anancus
5 4

Tetralophodon
Phylogeny and distribution of

Eubelodon
Gomphotherium, Tetralophodon,
?
Anancus and the American Gom-
10
photheriidae. Simplified pre-
sumed phylogenetic relation-
Stegomastodon
ships mostly after Gheerbrant/
Stegotetrabelodon
Tassy 2009; Tassy 1990; Tassy/

Gomphotherium

Gomphotherium
Miocene

Darlu 1987; Lambert/Shoshani

15 Rhynbchotherium
1998. Conventions as in Fig. 3.
Phylogenese und Verteilung von

Stegolophodon

Gomphotherium
Gomphotherium Gomphotherium, Tetralophodon,
Anancus und amerikanischen
Gomphotherium

20 Cuvieronius Gomphotheriidae. Vereinfachte

angenommene phyogenetische
Verwandtschaftsbeziehungen
mehrheitlich nach Gheerbrant/
Anancus
Tassy 2009; Tassy 1990; Tassy/
Oligo

25
Darlu 1987; Lambert/Shoshani
1998. Angaben wie in Abb. 3.

The coincidence of tectonics and this process may have the lower incisors and the size of the symphysis, but
caused a »land bridge«, allowing for a major faunal this is not documented in the record. There seems to be
exchange, which is known as the Great American Inter- material from Spain that is intermediate, for instance
change (Webb 1991). Initially, a large number of North in being nearly tetralophodont, but maintaining an
American mammals dispersed into South America, enamel band on the tusks (Mazo 1996). Gheerbrant/
including the relatives of camels (like the living llama Tassy (2oo 9) indicated a Gomphotherium from Ger-
and vicuña), while most of the dispersals to the north many to be particularly closely related to Tetralopho-
occurred a little later. The proboscidean Cuvieronius don longirostris. These Spanish and German fossils are
was one of the animals dispersing to the south. Later about 12–11 Ma old.
also Stegomastodon followed. Anancus replaced Tetralophodon in Europe, Asia and
Africa. The Indian Paratetralophodon (Fig. 4) seems to
Tetralophodon – Anancus
be close to Anancus and the first record of that genus
Tetralophodon is advanced over Gomphotherium in seems to be also in the same area. Anancus was abun-
being tetralophodont (four lophs or pairs of cusps dant in East Africa, but disappeared there, possibly
in the fourth milk tooth and first and second molar). in relation to the spread of C4 grass lands (see sec-
Similarly its third molar is more complex (Fig. 2). The tion on Atmospheric carbon dioxide). In North Africa,
lower tusk is reduced, but the symphysis is still long. which maintained different vegetation, it survived
Anancus is still more advanced in this way and may until around 1,8 Ma when isotopes suggest increas-
be even pentalophodont. Its upper tusks are immense ingly dry conditions (Sahnouni et al. in press). Anan-
and the mandibular symphysis is shortened as much cus became extinct worldwide after about 1,8 Ma ago,
as in the Elephantidae (Fig. 4). around or after a time, when the climate is believed to
Tetralophodon is generally believed to have evolved have become colder and in a wide area and also more
from Gomphotherium. Where this happened is not arid (deMenocal 1985).
quite clear, but the genus appeared in more or less the These genera were mostly classified in the Gomphoth-
same time in Africa southern and northern Asia and eriidae (e. g. Tassy 1986; Tobien et al. 1988; Gheerbrant/
in North America. This transition should be visible in Tassy 2oo9), more recently they are being classified
a spectacular reduction in the size and eventual loss of outside the Gomphotheriidae and in the Elephantoidea

// Die Evolution der Elefanten und ihrer Verwandten 349

 Africa S Asia Europe N Asia N America
eustaticsea level (m)
–130–110–90 –70 –50 –30 –10 10 30 50 Platybelodon

»M.« grandincisivus?

Stegodon

Stegodon
0

Stegolophodon
Stegolophodon
Pliocene Pleisto

Selenotherium

Platybelodon-Torynobelodon
Stegodon
Primelephas
5

Amebelodon-Serbelodon
Phylogeny and distribution of Stegodon
»M.« grandincisivus

the Amebelodontinae (red), 5

»M.« grandincisivus
Stegodontidae (blue) (insular
species omitted). Simplified

Platybelodon?

Platybelodon
»M.« gran-
presumed phylogenetic relation-

dincisivus
Protanancus
ships mostly after Gheerbrant/
10
Tassy 2009; Tassy 1990; Kalb
et  al. 1996a; Mackaye et  al.
2005; Lambert/Shoshani 1998. Stegolophodon
Miocene

Conventions as in Fig. 3.

Protanancus
Phylogenese und Verteilung 15
der Amebelodontinae (rot),

Gomphotherium
Stegodontidae (blau) (Inselarten
sind weggelassen). Vereinfachte

Gomphotherium
Gomphotherium
angenommene phyogenetische 20

Gomphotherium
Verwandtschaftsbeziehungen
mehrheitlich nach Gheerbrant/
Tassy 2009; Tassy 1990; Kalb Amebelodon Platybelodon
et  al. 1996a; Mackaye et  al.
Oligo

2005; Lambert/Shoshani 1998. 25

Angaben wie in Abb. 3.

(Shoshani/Tassy 2oo5). However, in cladistic analyses of Europe and America in the Eocene, they became
Tetralophodon and Anancus, or »tetralophodont gom- extinct in Europe, but continued evolving in North
photheres« are placed in a basal position relative to America. At least on three occasions, primitive horses
Stegolophodon and the Elephantinae (Kalb et al. 1996; dispersed across Bering into the Old World: around
Gheerbrant/Tassy 2oo9). This is impossible if Stegolo- 2o Ma Anchitherium (coincident with the »Proboscidean
phodon appeared about 16,5 Ma ago (see below) and Datum Event«), at 11 Ma Hipparion and 2,6 Ma Equus
Tetralophodon about 11–12 Ma ago. It seems thus, that (coincident with the Great American Interchange). The
the origin and classification of these taxa is not yet dispersal of Hipparion from North America to virtually
settled. all parts of the Old World is a very important event
and coincides with the dispersal of Tetralophodon in
Glaciation on the northern hemisphere and faunal
the opposite direction and may have coincided with a
exchange Asia – North America
dispersal of Mammut, but the direction of this disper-
The formation of glaciers on land lowers global sea sal is not clear.
level, since the source of the ice is the water of the
oceans, but the floating arctic ice does not affect sea Stegotetrabelodon
level. Whereas the »Proboscidean Datum Event« was Stegotetrabelodon is a large bunodont tetralophodont
triggered by glaciations on Antarctica, glaciation on form, with a complex third molar with the cusps trans-
the northern hemisphere allowed the Great Ameri- formed in transverse ridges (Fig. 2). However, its most
can Interchange. However, this was not the first time characteristic feature consists of two long and straight
that a glaciation on the northern hemisphere occurred. tusks in the lower jaw inserted in a long and narrow
Around 11 Ma, possibly for the first time extensive ter- symphysis (Fig. 4). The incisors have a large round
restrial glaciers formed on the northern hemisphere on section. There are also two long and straight upper
Greenland (Helland/Holmes 1998). At that time, some incisors. The genus is recognized in Africa and Arabia
very important faunal exchange between North Amer- (Tassy 1999).
ica and the Old World occurred. There has been much discussion on the origin of
Primitive horses originated when Europe and North this genus (Tassy 1999). Material commonly assigned
America were still connected. Following the separation to Tetralophodon is variable as to size and some have

350 Jan van der Made // The evolution of the elephants and their relatives
large lower tusks and others none and narrow elon-
gated symphyses. Whether or not this represents
intraspecific polymorphism or different species is not
resolved, but the forms with large tusks suggest affini-
ties of Stegotetrabelodon with Tetralophodon.
The previous question is related to the different
views on the classification of Stegotetrabelodon in
the Gomphotheriidae or in the Elephantidae. Its molar
structure suggests affinities with primitive Elephanti-
dae. The insertion in a cladogram by J. E. Kalb et al.
(1996) of Stegotetrabelodon, with its huge lower tusks,
between Stegolophodon, with small lower tusks since
early Middle Miocene and Stegodon with very advanced
tusk reduction is curious in this respect and a descent
from Tetralophodon seems more plausible.
The genus Stegodibelodon is usually classified along
with Stegotetrabelodon, but does not seem to be well
known and has occasionally been included in other
genera like Loxodonta (Beden 1983).
Atmospheric carbon dioxide, photosynthesis and mammal
evolution
Today we are burning fossil fuels and increasing the
carbon dioxide in the atmosphere, creating thereby a
global »green house«. Between 6–8 Ma ago, just the
opposite may have occurred by natural causes. Apart
from the effects that this may have had on global tem-
perature, this had a profound effect on the vegetation
for another reason.
There are different pathways of photosynthesis, one
is called C3 and is efficient when there is much carbon
dioxide in the atmosphere, and the other is called C4 and
is efficient when there is less CO2 . Most trees and shrubs
use C3 type of photosynthesis, while most grasses use
the C4 pathway. The normal isotope of carbon has an
atom weight 12, where as there are also isotopes with
weights of 13 and 14. The latter is well known for radio-
13
carbon dating, but here the stable isotope C is of inter-
est. A small proportion of this isotope is present in the
carbon dioxide in the atmosphere. Photosynthesis is
selective to CO2 with this heavy isotope, and C3 and C4
13
plants take C up in different proportions to the normal
12
C . By analyzing carbon isotopes in carbonate nodules
in ancient soils, it is possible to know the kind of plants
that grew on that soil. But even more interesting here,
by analyzing this isotope in fossil tooth enamel, it is
possible to know what kind of plants these animals ate.
Between 6–8 Ma ago, C4 grasslands became widespread
at low latitudes, while at higher latitudes this occurred
later or not at all (Cerling et al. 1997).
// Die Evolution der Elefanten und ihrer Verwandten
The appearance of open landscapes might have trig-
gered the evolution of bipedal walking in our ancestors
at that time. Most herbivorous animals adapted rela-
tively rapidly to the environmental changes by evolv-
ing different dentitions. Depending on the growth
stage, grass may have a low nutritive value and more
of it has to be eaten. For this reason, grazers tend to
increase the occlusal surface of the teeth. In ruminants
and pigs this is seen in an increase in the size of the last
molar. Many grasses have phytoliths, which consist of
silicium oxide, just like glass: one can cut one’s hand
with a leaf of a certain type of grass. These phytoliths
cause much wear on the teeth, and as a result teeth
with higher crowns evolved to compensate for the wear.
These and other adaptations started to evolve about 6
13
to 8 Ma ago and the C isotopes of these teeth confirm
that these animals indeed changed their diets (Cerling
et al. 1997; 1999).
These changes had a profound effect on the evo-
lution of the proboscideans and are the cause of the
appearance of true elephants. Shortly after 8 Ma, pro-
boscideans started to include more C 4 plants in their
diets and not later than about 7 Ma ago Stegodon
and Primelephas evolved from Stegolophodon by the
progressive addition of lobes to the molars (Fig. 2). In
addition to this trend, the lobes evolved into flattened
transverse plates, while the space between the plates
became filled in by cementum. The occlusal surface
changed functionally and became a nearly flat grind-
ing surface. Chewing changed from crushing to grind-
ing. This trend continued, but also crown height started
to increase in Loxodonta, Mammuthus and Elephas
(compare side views of the molars of Primelephas and
Elephas in Fig. 2). Other proboscideans changed their
diet, but maintained bunodont dentitions until they
became extinct. Deinotherium did not change its diet
or its morphology and remained one of the few large
browsers in Africa, until it became extinct close to the
Plio-Pleistocene transition (Cerling et al. 1997; 1999).
Stegodonts
The family Stegodontidae (or subfamily Stegodonti-
nae within the Elephantidae in other classifications)
contains the genera Stegolophodon, Stegodon and the
recently described Selenetherium.
Stegolophodon is the first to appear and has very
early records in Japan (Koda 2oo3; Kalb et al. 1996a).
Some of the oldest specimens are relatively bunodont,
but at an early stage the cusp and cusplets became
arranged in transverse ridges (Fig. 2). This suggests that
351
the genus evolved from a bunodont form in northern sidered it ancestral to Elephas and Mammuthus, but not
Asia. The skull is still long and low, but at a very early to Loxodonta, which he believed to have evolved from
stage the lower tusk became much reduced (Fig. 5). Stegotetrabelodon. In the more recent analyses, Prim­
Stegodon is widely assumed to have arisen from elephas is in a more basal position than Loxodonta.
Stegolophodon. The skull is shortened and the upper Loxodonta has cheek teeth with higher crowns. Con-
tusks are huge (Fig. 5). The lower tusks are generally trary to the other Elephantinae, it has rhomb shaped
absent and the anterior part of the mandible is short- lophs, which are very thick (Fig. 2). The primitive spe-
ened. The ridges or plates of the cheek teeth are well cies L. adaurora and L. exoptata are recognized. The
developed and increased in numbers (Fig. 2), a trend morphology of the dentition suggests it is a special-
which was to continue within this genus. The molars ized grazer, but T. E. Cerling et al. (1999) found that
of some forms became high crowned. There are many the recent Loxodonta africana is a C3 browser. Around
dwarfed endemic insular species on the different 1,8 Ma ago, the genus disappeared from east Africa, but
islands of Japan, the Philippines and Indonesia. survived in north and south Africa. In a Mediterra-
Selenotherium lacks a lower canine, its molars have nean type of environment, C3 vegetation is dominant.
low crowns with cusp pairs that are fused into curved Loxodonta atlantica may have become again a browser
transverse crests, with a convex anterior surface and and when it dispersed into east Africa at the end of the
it has a third lower molar with six of these (MacKaye Pleistocene, it may have maintained this habit. Some
et al. 2oo5). consider the smaller Loxodonta cyclotis a different spe-
Stegodon has traditionally been seen as ancestral cies, but most consider it a subspecies.
or close to Loxodonta, Elephas and Mammuthus. In the The genus Elephas contains the subgenera Elephas
cladograms by H. T. MacKaye et al. (2oo5), the Afri- and Palaeoloxodon. The earliest representative is E. na‑
can Selenotherium is generally inserted between Ste- watensis, which had still low crowns, a low number
golophodon and Stegodon, which would suggest that of plates and a low plate frequency. This form may
the origin of the latter genus is in Africa. However, have given rise to the lineage of Elephas recki, which
H. Saegusa et al. (2oo5) argues that the evolution of Ste- consists of the sequence of species or subspecies eko-
golophodon to Stegodon is documented in Asia. These rensis, brumpti, shungurensis, atavus, ileretensis, recki
views are mutually exclusive. and iolensis, covering more than 4 Ma. Some consider
that this sequence, with the possible exception of E. (P.)
Elephants: Primelephas, Loxodonta and Elephas
iolensis represents an anagenetic lineage, while others
Generally, the subfamily Elephantinae is considered think that these forms have overlapping chronologic
to include the genera Primelephas, Loxodonta, Mam- distributions (Beden 1983; Todd 2oo5). This sequence
muthus and Elephas (including the subgenus Palae- of forms is characterized by an increase in plate num-
oloxodon) (Shoshani/Tassy 1996, appendix C1; 2oo5). bers and lamellar frequence and in skull shape.
In a cladistic analysis by J. E. Kalb et al. (1996), two Elephas ekorensis, an early member of the lineage or
characters unique to these genera are presented: one branch leading to E. recki, is believed to have given rise
of them is a detail of tooth morphology and the other to Elephas (Elephas) planifrons (Beden 1983), which dis-
is a minimum plate number on the lower third molar persed into southern Asia. In Pakistan it is recorded in
of eight. Apart from Primelephas, the remaining three strata with ages from 3,3–2,7 Ma. This is still a primi-
genera are all clearly hypsodont, but this occurs also tive form with low crowns, a low lamellar frequency
in some stegodonts. The elongation of the third molar and thick enamel. After 2,7 Ma it is replaced by E. (E.)
is an important tendency in this group. Within this hysudricus (Fig. 2; 6), which arrived by dispersal and
group the following parameters increase: crown height, which has higher crowns, a higher lamellar frequency
number of plates per tooth, and the lamellar frequency and thinner enamel (Hussain et al. 1992). The arrival
(number of plates per ten centimetres). The plates, of this species coincides with the first appearance
lamellas, lobes, or lophs become flatter and enamel of deer and lagomorphs in the Indian Subcontinent
thickness decreases. and is probably related to the changes in global cli-
Primelephas is the most primitive of these four mate at that time (see section Astronomy, ice ages and
genera in that it has still very low crowned molars with elephants). Elephas hysudricus may have given rise to
a limited number of thick plates (Fig. 2), but much of Elephas hysundrindicus in Java and Elephas (Elephas)
this genus, like its skull, is unknown. Beden (1983) con- maximus on the mainland of southern Asia. During

352 Jan van der Made // The evolution of the elephants and their relatives
 Africa S Asia Europe N Asia N America
Africa S Asia Europe N Asia N America

P. namadicus
E. maximus
0

Loxodonta

Elephas (Palaeoloxodon) recki-lineage

Mammuthus columbii
M. primigenius

M. primigenius
M. primigenius
Elephas planifrons

M.. trogontherii
1

M. trogontherii

Mammuthus meridionalis
P. antiquus
E. hysudricus

E. (Pal.) namadicus

E. (Pal.) antiquus
Elephas hysudrindicus
6
2 Phylogeny and distribution of
the Elephantinae (insular spe-
Loxodonta
cies omitted). Simplified pre-

M. meridionalis
Mammuthus meridionalis
africana
3 sumed phylogenetic relation-

Mammuthus rumanus
ships mostly after Kalb et al.
Selenotherium

1996; Tassy 2003; Saegusa/

Primelephas

4
Elephas nawatensis

Gilbert 2008; Lister et al. 2005.

Conventions as in Fig. 3.
Mammuthus subplanifrons-africanavus

Mammuthus
5
Elephas hysudricus Phylogenese und Verteilung der
Elephas Elephantinae (Inselarten sind
(Palaeoloxodon)
Elephas maximus weggelassen). Vereinfachte
6 antiquus
angenommene phylogenetische
Verwandtschaftsbeziehungen
7 mehrheitlich nach Kalb et al.
1996; Tassy 2003; Saegusa/
Gilbert 2008; Lister et al. 2005.
8
Angaben wie in Abb. 3.

the Late Pleistocene it dispersed into Java and replaced sion is stronger in arid environments and the wind
there E. hysundrindicus (Van den Bergh 1999). may carry dust far away. Windblown dust in ocean
sediments can be recognized and shows that during
Africa isolated again by increasing aridity in the Middle
the Late Pliocene and Pleistocene there were impor-
East and Sahara
tant fluctuations in aridity in north Africa and Arabia,
For a long time after the Mid-Miocene Climatic Opti- which correspond to the different Milankovich cycles
mum no proboscideans dispersed out of Africa, but (see section Astronomy, Ice ages and Elephants), but
around 3,2 Ma two dispersals occurred: Mammuthus that there was an overall trend in increasing aridity
dispersed to northern Eurasia and Elephas planifrons (deMenocal 1995).
to the Indian Subcontinent. These dispersals may have Due to this increasing aridity, the faunal exchange
been possible, because these proboscideans became between Africa and Eurasia decreased and became
adapted to grazing in Africa, which made it possible for more selective. It had probably a major effect on the
them to disperse across the Middle East and, in the case dispersal of humans. The earliest humans with a primi-
of Mammuthus, into the more seasonal higher latitudes. tive type of lithic industry (called Oldowan) dispersed
The Middle East had probably a dryer climate and in out of Africa before faunal exchange became very
seasonal climates it is more difficult for a browser to restricted at 1,8 Ma. A more advanced type of lithic
survive during the winter. industry developed in Africa shortly later, but its dis-
The geographic distribution of mammals suggests persal out of Africa was halted in the Levantine area,
that there was a zone of dry or open landscapes run- until about 1– o,9 Ma ago, when it dispersed into Asia
ning across northern Africa and the Middle East to and subsequently into Europe. This dispersal coincides
central Asia, at least since the Middle Miocene, though with a short period of faunal exchange between Africa
the extent, position and degree of dryness of this zone and Eurasia and with new species dispersing into (but
may have fluctuated (Van der Made in press; Van der not through) the Levantine corridor (Van der Made
Made/Mateos in press). A study of phytoliths suggests in press). Elephas (Palaeoloxodon) antiquus was one of
that open landscapes were common in Turkey at least those species.
since 2o Ma ago, but that these were not dominated
Elephants: Palaeoloxodon
by C4 grasses (Strömberg et al. 2oo7). Sand dunes in
Chad testify the existence of extremely arid conditions Palaeoloxodon is a subgenus of Elephas and is discussed
as much as 7 Ma ago (Schuster et al. 2oo6). Wind ero- in detail by Saegusa/Gilbert 2oo5. It is characterized

// Die Evolution der Elefanten und ihrer Verwandten 353

Unterkiefer von Phiomia wintoni,
Oase El Fayum, Ägypten; Jabal
Qatrani-Formation, ca. 33 Milli-
onen Jahre vor heute. Leihgabe:
Naturhistorisches Museum Wien
(oben). Unterkiefer von Palaeo-
mastodon beadnelli, Oase El
Fayum, Ägypten; Jabal Qatrani-
Formation, ca. 33 Millionen Jahre
vor heute. Leihgabe: Museum für
Naturkunde Berlin (unten).
Mandible of Phiomia wintoni, El
Fayum Oasis, Egypt; Jabal Qatra-
ni-Formation, about 33 million
years BP. Item on loan from:
Naturhistorisches Museum Wien
(above). Mandible of Palaeomasto-
don beadnelli, El Fayum Oasis,
Egypt; Jabal Qatrani-Formation,
about 33 million years BP. Item
on loan from: Museum für Natur­
kunde Berlin (below).
Unterkiefer eines Wollhaarmam-
muts (Mammuthus primigenius),
Speyer, Rheinland-Pfalz; ca.
14 000 Jahre vor heute (oben).
Unterkiefer eines Steppenele-
fanten (Mammuthus trogontherii),
Grube Dyckerhoff/Mosbacher
Sande, Wiesbaden, Hessen; ca.
400 000 Jahre vor heute (unten).
Beide Leihgaben: Staatliches
Museum für Naturkunde
Stuttgart.
Mandible of a woolly mammoth
(Mammuthus primigenius), Speyer,
Rhineland-Palatia; about 14,000
years BP (above). Mandible of a
steppe elephant (Mammuthus tro-
gontherii). Pit Dyckerhoff/Mosba-
cher Sande, Wiesbaden, Hesse;
about 400,000 years BP (below).
Both items on loan from: Staat­
liches Museum für Naturkunde
Stuttgart.
by skull morphology (Fig. 6) and may have derived
from the Elephas recki lineage about 1– o,9 Ma ago and
at that time may have spread into Eurasia. This might
have occurred because glacial cyclicity became more
pronounced and the genus Mammuthus adapted to the
cold environments, leaving a niche at mid latitudes
for a proboscidean adapted to warm environments
(Van der Made/Mazo 2oo1). Traditionally European
fossils have been assigned to Elephas (P.) antiquus
and Asian fossils to E. (P.) namadicus. Saegusa/Gilbert
(2oo5) however, noted that two morphologies, corre-
sponding to different species, are found in Europe and
offers three alternative scenarios to explain this. In
one of these, a primitive morphology, corresponding
to E. antiquus, is replaced after 3oo ka (Steinheim) by
a morphology, which corresponds to E. namadicus.
This may have occurred also in the mainland of East
Asia. Before this replacement occurred, Palaeoloxodon
with the primitive morphology, reached the Japanese
islands, dwarfed, and became the endemic E. (P.) nau-
manni. It survived in isolation, while on the main-
land the more advanced morphology (P. namadicus)
replaced the primitive morphology. The dispersal into
Japan may have occurred around 4oo ka. Palaeoxodon
became extinct before 2o ka in Europe and before the
last glacial maximum in Japan. It is possible that the
latest Elephas from Africa is conspecific with the Eura-
sian species.
The subgenus Palaeoloxodon colonized several Medi‑
terranean islands and gave rise there to endemic dwarf
species, which will be discussed elsewhere (Palombo,
this volume).
Astronomy, Ice ages and Elephants
It is well known that the attraction of the moon causes
the tides. This is a cycle of astronomic origin. Other
cycles of astronomic origin, but of a much longer dura-
tion shaped the evolution of the mammoths.
Like the moon, which attracts and is attracted by
the earth, other planets, when their course brings them
near the earth, may deform the orbit of the earth
around the sun. There are several cycles of astronomic
origin, which are known as Milankovich Cycles. The
eccentricity cycle has a periodicity of 1oo,ooo years (or
1oo kiloannum or 1oo ka) and consists in that the orbit
of the earth around the sun is more circular or a little
more elliptic. This cyclicity is related to the ice ages.
The obliquity cycle has a periodicity of 4o ka and con-
sists in a variation of the angle between the plane of the
orbit of the earth around the sun and the axis around
356 Jan van der Made //
which the earth rotates. The more inclined the axis of
the earth, the stronger seasonality, which is especially
noted at higher latitudes. There is also a 2o ka cycle in
the precession of the equinoxes. The effects of these
cycles interfere.
Before 2,6 Ma, the 2o ka cycle was dominant, but
after that date the 4o ka cycle became dominant,
resulting in a periodically increased seasonality at
higher latitudes. As a result many evergreen plants
disappeared from higher latitudes and fruit produc-
tion was restricted during part of the year. Folivo-
rous and frugivorous animals that were deprived of
their preferred food during winter also disappeared
from higher latitudes. For instance tapirs and Mam-
mut borsoni went extinct in northern Eurasia. Grazers
extended their range to the north and Mammuthus
meridionalis reached central and Western Europe.
Mammuthus had become a grazer in Africa due to
environmental change and was pre-adapted to the
new environments that appeared in Eurasia. Around
this time, also glaciations occurred on the northern
continents. Sea level lowered, the Great American
Interchange occurred, the horse Equus dispersed from
North America to Europe and Mammuthus did so in
the opposite direction.
Between 1,2– o,9 Ma, the 1oo ka eccentricity cycle
became dominant leading to the well known glacial
cycles. During these cycles large parts of the north-
ern continents were covered by glaciers. Up to that
time, there were east-west oriented vegetation zones
in northern Eurasia: the tundra in the north which is
humid, but where trees do not grow because of the low
temperatures, the wooded taiga, and the dry and open
steppe in the south. During glacial periods, tempera-
tures dropped and the tundra expanded southward
until it fused with the steppe, forming an open habitat
that extended from the Atlantic to Alaska (Due to the
glacial low sea levels, the Bering Strait became land).
This environment is called the »mammoth steppe«.
During glacial periods, the »interglacial« animals sur-
vived in refuge in the south; in the Iberian Peninsula,
Italy, and the Balkans and in China. During intergla-
cial periods, the »glacial« animals survived in the tun-
dra, steppe or mountains.
The onset of the 1oo ka glacial cyclicity caused much
successive dispersal of species into Europe, leading
to an apparent increase in diversity. Whereas there
was just one elephant species in Europe (Mammuthus
meridionalis) before o,9 Ma, after that date Mammuthus
became typical of the cold stages and Elephas antiquus
The evolution of the elephants and their relatives
dispersed from the south and became the typical ele-
phant of the warm stages.
Elephantidae: mammoths
The mammoths or Mammuthus differ from other ele-
phants in the shape of their skull and in the strongly
curved and twisted tusks being placed close together
and diverging little at the basis (Fig. 6). There is some
discussion on the affinities of Mammuthus: it was seen
as a direct descendant of Primelephas (e. g. Beden 1983),
more closely related to Elephas (Kalb et al. 1996) or to
Loxodonta (Debruyne et al. 2oo3). In the evolution of
this genus, hypsodonty, plate number and lamellar
frequency increased, which is documented by A. M.
Lister (1996) for M. meridionalis, M. trogontherii and
M. primigenius.
The oldest species that is placed in Mammuthus is
the African M. subplanifrons, which in Africa gave rise
to M. africanavaus. Around 3,2 Ma, there is a faunal
turnover in Europe, which probably is related to cli-
matic change. It may have been at this time that Mam-
muthus rumanus arrived in southeastern Europe. This
animal was probably a grazer and its dispersal further
into Europe may have been blocked by a forested envi-
ronment in central Europe. This is a common pattern
and occurred in many different species at many dif-
ferent times (Van der Made/Mateos in press). When
around 2,6 Ma the climate became more seasonal and
possibly a glaciation occurred, Mammuthus meridiona-
lis dispersed further into Europe and possibly at this
time further into Asia as well. The evolution of Eurasian
mammoth is described by A. M. Lister et al. (2oo5). In
Western Europe, M. meridionalis, M. trogontherii and
M. primigenius form a sequence in time, but this is not
a simple anagenetic lineage. Between 2–1,5 Ma, Mam-
muthus trogontherii evolved in Asia from M. meridi-
onalis, and around 65o ka, during a particularly severe
glacial (the »Elsterian«), dispersed into Western Europe
and replaced M. meridionalis there. This pattern may
have been repeated by Mammuthus primigenius, which
may have arisen around 7oo ka in north-eastern Sibe-
ria, and dispersed into Western Europe shortly after
2oo ka. These species were cold adapted and at mid-
latitudes were present in the glacial faunas, but were
replaced by Elephas antiquus during the interglacials.
Mammuthus meridionalis may have given rise to the
endemic insular Mammuthus lamarmorae of Sardinia
(see Palombo in this volume). Mammuthus meridiona-
lis did not only disperse into northern Asia, but around
2,6 Ma it also reached North America. There it evolved
// Die Evolution der Elefanten und ihrer Verwandten
into the local species Mammuthus columbi. Later, Mam-
muthus primigenius dispersed from Asia into North
America, replacing the previous species.
Mammuthus primigenius, the woolly mammoth, is
extremely well studied because many frozen cadavers
were found in the permafrost. It was a cold adapted
species with dense pelage and small ears (extremities
can become frozen in cold climates, so they tend to be
reduced in the course of evolution). At the end of the
last ice age, its range contracted in Siberia (Stuart et al.
2oo4; Nogués-Bravo et al. 2oo8) and species numbers
dropped. This was a natural process and probably had
happened before, followed by an expansion during the
following glacial. However, this time a new human
species was present, which probably was a more able
hunter and had the capacity to live and hunt further to
the north. The youngest fossils of this species are from
the island of Wrangel, north of Siberia and date from
about 4ooo years ago, being roughly contemporary to
the Egyptian pyramids (Vartanyan et al. 1993).
Megafauna extinctions in the wake of human dispersal
The title of this section refers to a chapter in this vol-
ume with the same title. Our species, Homo sapiens,
arose in Africa some 2oo ka ago and subsequently
spread over the world. Its appearance in a continent
or island coincided with extinctions of large mam-
mals, and frequently also of other animals and plants.
There is discussion on the causes of these extinctions:
human hunting as well as other human activities, cli-
mate change or a combination of these factors.
Proboscideans that went extinct around the time
that the human species arrived in the area where they
and their ancestors had lived for a long time, in some
cases millions of years, include: Mammut (North Amer-
ica) Stegomastodon (South America) and Cuvieronius
(America), Stegodon (Asia), Elephas (Palaeoloxodon)
namadicus (Eurasia), Elephas (Palaeoloxodon) recki
lineage (E. iolensis; Africa), Mammuthus primigenius
(North America, Eurasia), as well as endemic insular
species on Cyprus, Sicily, Crete and other Mediterra-
nean islands, Japan, Philippines, Sulawesi and other
Indonesian islands, and the islands off the coast of
California. According to current classification, these
species represent four different families. They are not
only taxonomically diverse, but also ecologically and
they have survived many climatic changes previous
to their extinction. There are still two living species,
but for how long?
357
Acknowledgements
I thank Prof. D. Mania for inviting me to participate
in the Neumark-Nord working group and Dr. H. Meller
for inviting me to participate in this volume as well as
Source of figures
Tab. 1 J. van der Made, Madrid
1–6 J. van der Made, Madrid
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