The majority of this section will examine the temporal distribution o f Palaeoloxodon antiquus. Discussion of the distribution of the Asian representatives of the genus will also be given although in less detail; these are currently being researched in detail by others (Chang, in prep.). The following sections are based on data gathered in the present study and also from examination of the literature.
3.3.1. First appearances
3.3.1.1. European species of Palaeoloxodon
Only Palaeoloxodon antiquus, the mainland European Palaeoloxodon species will be discussed below. The distribution of the Mediterranean dwarf elephants is discussed in detail in Chapter 7.
Southern Europe
One of the earliest, previously recorded occurrences of Palaeoloxodon antiquus in Europe is from deposits in the Guadix-Baza basin complex at Huéscar-1 in southern Spain. This site is composed of a number of deposits, of various ages from around 800,000 - 600,000 years old, and has yielded a number of large mammal fossils (Mazo 1989). The amount of possible P. antiquus material recovered is small, consisting of several tooth plate fragments and small fragments of tusks. However, a fairly complete mandible with molars M2 and M3 has also been recovered and it is on this specimen which species identification has been made by Mazo (1989).
Comparison of the teeth to those of mammoth species {Mammuthus meridionalis and
Mammuthus trogontherii) by Mazo (1989) shows them to be referable to P. antiquus,
this identification is based mainly upon the general appearance of the occlusal surface. Mazo (1989) comments that the teeth have a primitive structure, the plates being thick with a rhombic shape.
During the present study it was not possible to observe the Huéscar-1 material and no measurements of the teeth are available. However, after examination of the specimen by Stuart in 2000 (pers. comm.) and from observations of photographs of the specimen by the author it is concluded that the mandible should be referred to M. trogontherii. This is based on the relative width of the teeth and the occlusal surface pattern. The teeth of P. antiquus are characteristically narrow for their length and have a very distinct occlusal surface pattern (see Chapter 4 for more detail). The mandible from Huéscar-1 has very wide teeth for their length with an occlusal pattern characteristic of Mammuthus
Several early sites in Italy have yielded Palaeoloxodon antiquus remains. These sites have a faimal assemblage regarded as characterising the middle Galerian mammal age and broadly correlated to the “Cromerian Complex” (Gliozzi et al 1997). It includes; the rhinoceros, Stephanorhinus hundsheimensis, large forms of the aurochs. Bos and the horse, Equus cahallus as well as P. antiquus (Azzaroli 1977,1983). The following middle Galerian sites have yielded P. antiquus remains: Slivia (Bon et al 1992), Isemia La Pineta (Sala 1996) and Ponte Galeria (a complex of sites) (Caloi and Palombo 1994).
These sites are all thought to be of similar age. Both the Ponte Galeria complex and Isemia, have been dated by potassium-argon analysis to approximately 730,000 yBP (Capasso and Petronio 1984). Slivia has been correlated with both these sites based on geological and molluscan biostratigraphic evidence (Gliozzi et al 1997) although Bon
et a l (1992) suggest it may be slightly older. Together these sites represent some of the earliest definite records of P. antiquus is Europe.
Northern Europe
The oldest sites to yield P. antiquus remains in northern Europe are Pakefield-
Kessingland, and Gorton, Suffolk in the Cromer Forest bed Formation. As discussed in Chapter 2, these sites have a similar fauna to West Runton, including Mimomys savini
but as Stuart and Lister (2001) point out, P. antiquus makes its first appearance here and is found with Hippopotamus sp. and Megaloceros dawkinsi both of which are not seen at West Runton. Pollen from both sites indicates the deposits were laid down in pollen zone n and so this rules out the Suffolk sites representing a different stage of the same interglacial as West Runton. As explained in Chapter 2 the dating of this site is
problematic but the presence of Mimomys means it must predate the Arvicola sites of Boxgrove, Westbury and Ostend correlated with MIS 13. The change in fauna from West Runton (which could be assigned to MIS 19) means the Pakefield-Kessingland site must date to at least MIS 15 and possibly MIS 17. This thus represents the oldest
occurrence of P. antiquus in northern Europe.
Palaeoloxodon antiquus has also been recovered from the Cromer Forest Bed site of Ostend in Norfolk. Here two teeth (associated upper M3s) were collected by Green (1842) and are now housed in the Natural History Museum, London. One of these teeth
was originally identified as belonging to M. trogontherii by Stuart and West (1976) although previously Adams (1878) had identified both as being fi’om P. antiquus, which is in agreement with the present study. Pollen analysis firom sediment adhering to one of these teeth is unlike the pollen spectra obtained firom other samples in the Green
Collection, having high levels of Corylus and containing grains referable to ‘type X ’, known fi*om the Hoxnian (Lister, pers. comm.). This contrasts to pollen firom cores at the site and adhering to other mammalian specimens, which indicated an open
environment interpreted as the end of an interglacial (Stuart and West 1976). This suggests that either the remains come from a later interglacial than the other Ostend material, or that they correspond to a different temperate phase of the main Ostend interglacial. Recently (Stuart and Lister 2001), the main deposits at Ostend have been referred to MIS 13.
Germany has yielded a number of important early Middle Pleistocene sites firom which
P. antiquus has not been recovered. Of these, the most significant are Siissenbom and Voigtstedt, both of which are roughly correlated with West Runton. Both of these sites have Mimoyms savini, with closed roots, as opposed to the more advanced Arvicola,
characteristic of “Cromerian Complex” sites of a later age (Koenisgwald and
Koefschoten 1996). At West Runton and Voigtstedt a small number of elephant remains have been recovered, together with the important record of a fairly complete skeleton of
Mammuthus trogontherii at West Runton (Stuart 1996) and so the absence of P.
antiquus here may be due to the lack of material. However, at Siissenbom over 1000 elephant molars have been found, all of which are referable to M. trogontherii. Dietrich (1958) identifies a single molar as being fi-om Palaeoloxodon antiquus; however, examination of the tooth in question by Adam (1961), Aguirre (1969), Lister (pers. comm.) and personal observation, leads to the conclusion that the tooth is in fact a veiy worn lower M3 of M. trogontherii. Thus, P. antiquus was almost certainly absent fi*om Germany during the earlier part of the “Cromerian Complex”.
The first record of P. antiquus in Germany is in the later “Cromerian Complex” sites of Mauer and Mosbach H (the present study) where they are associated with Arvicola rather
iheeoMimomys. P. antiquus is found at low frequencies at these sites compared to the other elephant species present at the site, M. trogontherii. The reasons for this are
actual low number of P. antiquus at that time. A small number of P. antiquus remains have also been recorded from the German site of Karlich II. The age of this deposit is currently under investigation but, as discussed in Chapter 2, it is probably correlated with Stage 11 or 9 on the oxygen isotope record.
3.3.1.2. Middle Eastern species of Palaeoloxodon
As is discussed in detail in Chapter 8, the genus Palaeoloxodon almost certainly evolved in Africa from Palaeoloxodon recki atavus, a Pleistocene sub-species of the
Palaeoloxodon recki lineage. As will be discussed below, section 3.3.2, the only suitable route out of Africa that Palaeoloxodon, and a large number of other genera, could have taken was through the Levant. Thus, the Pleistocene fossils of the Levant have an important role in determining the radiation of Palaeoloxodon.
Unfortunately the Levant has relatively few fossiliferous Pleistocene sites, which is almost certainly related to taphonomic problems caused by the rocky terrain of a majority of the area (Horowitz 1976). The details of only two Middle Pleistocene sites, that have produced remains of P. antiquus, in the Levant, have been published; Gesher Benot Ya’aqov and Holon, both in Israel. However, there is probably a considerable amount of unprovenanced and unidentified fossil elephant material housed within collections (L. Horowitz, pers. comm.).
The most important of these sites, regarding the arrival of Palaeoloxodon in the Levant, is Gesher Benot Ya’aqov which is situated in the valley of the River Jordan, north of Jerusalem (Goren-Inbar et al 1994; Goren-Inbar et al. 2000). In 1989 a partial skull of
P. antiquus was discovered in association with stone and wooden artefacts indicative of an Acheulian living-fioor. The skull was positioned in such a way as to suggest
deliberate breakage for brain extraction; the significance of this is discussed later. The site was originally dated, based on geochronological evidence, to around 500,000 yBP. However, recent examination of the site now suggests a much older date of around 780,000 yBP, at the Brunhes-Matayama boundary (Goren-Inbar et al. 2000). The site is currently being reinvestigated and is thought to contain a long sequence spanning approximately 100,000 -120,000 years. The Palaeoloxodon skull was recovered from the top part of the sequence and it is unknown yet whether older material is present in
lower sections of the sequence. At present, this site represents the oldest occurrence of
P. antiquus in the Levant.
The site of Holon is situated on the coast west of Jerusalem (Yom-Tov and Tchemov 1988.). It has yielded an Acheulian industry with typical lower Palaeolithic lithics together with a fauna consisting of: Bos primigenius. Dama sp. Gazella sp.,
Hippopotamus amphibius and several teeth of Palaeoloxodon antiquus (identified by the author). Electron-spin resonance on two of the bovid teeth and thermoluminescence analysis has dated the site to between 198,000 - 201,000 yBP placing it in MIS 7 (Porat
et al 1999; Chazan et al. 2001).
Another important Middle Pleistocene site in the Levant is Latemné situated in the valley of the River Orontes, Syria. Hooijer (1961) described elephant remains here as being of Mammuthus trogontherii, however this has been challenged by Maglio (1973) who concludes the remains (teeth only) are referable to Palaeoloxodon antiquus (P. namadicus therein). Hooijer (1961) concluded that the teeth were from M. trogontherii
because of the lack of medial expansions on the occlusal surface, a characteristic of
Palaeoloxodon (see Chapter 4) and because the lateral rings of the plates in early wear are equal in size to the median, a characteristics o f Mammuthus. Maglio (1973) states that the teeth of Palaeoloxodon are highly variable and that the presence of the
Mammuthus characters should not preclude the specimen being referable to
Palaeoloxodon. Investigation in the present study has shown that the teeth of
Palaeoloxodon are highly variable (see Chapter 5) but almost always have the classic pattern of large median rings and small lateral rings in early worn plates. Since none of the teeth collected from Latemné have this type of occlusal pattern (personal
observations from photographs in Hooijer 1961 : p i 20) it is concluded that the teeth are referable to Mammuthus trogontherii.