Paleoloxodon

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Paleoloxodon
Skeletal reconstruction of Palaeoloxodon mnaidriensis

Skeletal reconstruction of Palaeoloxodon mnaidriensis

Temporal occurrence
Lower Pliocene to Middle Holocene
5 million years to 4,000 years
Locations
Systematics
Russell animals (Proboscidea)
Elephantimorpha
Elephantida
Elephants (Elephantidae)
Elephantinae
Paleoloxodon
Scientific name
Paleoloxodon
Matsumoto , 1924

Palaeoloxodon is an extinct genus of elephants that wascommonin Africa and over large parts of Eurasia during the Pliocene and Pleistocene . Its best-known member is the European forest elephant . This occurred mainly in the warm periods in western Eurasia, in the cold periods it was replaced by representatives of the genus Mammuthus . The genus includes numerous species, some of which are among the largest known proboscis . In addition,dwarfed forms occurredon numerous islands in the Mediterranean , which in turn represent the smallest elephants that have been handed down to date. The origin of the genus lies in the Pliocene of Africa, only in the Lower Pleistocene did it reach Eurasia. The mainland populations largely died out in the Upper Pleistocene; individual groups on the Mediterranean islands survived into the Holocene . Palaeoloxodon was scientifically introduced in 1924, but was originally considered a subgenus of Loxodonta , whichunitesthe African elephants . In the last quarter of the 20th century there was a synonymization with the genus Elephas , in which the Asian elephant is. Here was palaeoloxodon sometimes performed as a subgenus. However , according to genetic studies from 2017, paleoloxodon is actually more closely related to the genus Loxodonta .

features

Skull of Palaeoloxodon namadicus

The representatives of the genus Palaeoloxodon represent partly large to very large elephants , which sometimes exceeded the dimensions of today's African elephant ( Loxodonta africana ). For Palaeoloxodon antiquus , the European forest elephant, and Palaeoloxodon recki , an African form, sizes of up to 420 cm shoulder height are available, for Palaeoloxodon namadicus from Asia up to 450 cm is assumed. The weight of the animals should have been between 11 and 15 t in any case. On the other hand, dwarf forms emerged on several islands in the Mediterranean , which sometimes had less than 10% of the body dimensions of the large mainland animals. The smallest known member is Palaeoloxodon falconeri , the Sicilian dwarf elephant, which was found in Sicily and Malta and which weighed 170 kg with a shoulder height of around one meter. Other dwarf forms are passed down with Palaeoloxodon cypriotes from Cyprus , Palaeoloxodon lomolinoi from Naxos or Palaeoloxodon tiliensis from Tilos . Their size varied between 250 kg and 650 kg. In addition to these extremely dwarfed species, there were also individual intermediate forms, including Palaeoloxodon creutzburgi from Crete and Palaeoloxodon mnaidriensis from Malta.

The skull of Palaeoloxodon was extremely massive and tall and remarkably wide. It could reach a height from the alveoli of the upper tusks to the roof of the skull of up to 140 cm. At the top of the skull, the length of large forms was between 47 and 70 cm, the width between 76 and 114 cm. A striking feature was the massive parietal-occipital bone bulge that ran across the roof of the skull and had a deep indentation between two lateral bulges. The feature, especially the hump-like bulges, is shared by Palaeoloxodon with the Asian elephant ( Elephas maximus ), but the former was much more massive than the latter. The bone bulge resulted from the clear elevation of the skull and the forward displacement of the occiput . As a result, the area of ​​the frontal and parietal bones was also shifted forward and was almost vertical. In connection with individual other features, such as the strikingly parallel course of the temporal lines to the occlusal plane of the teeth, the upper part of the skull of Palaeoloxodon looked box-like overall. The parietal-occipital bone bulge showed both a certain phylogenetic and ontogenetic variation. Historically older representatives such as Palaeoloxodon recki had a rather intermediate bony bulge, in younger ones such as Palaeoloxodon namadicus and Palaeoloxodon antiquus it was much more pronounced. But even here it varied with the individual age of an animal and reached its greatest massiveness in the fully grown stage. Then the bulge could partially overhang the front skull and thus considerably expand the attachment points for the thick neck muscles. Another characteristic is the lateral inclination of the alveoli of the upper tusks . This was particularly pronounced in the developed Eurasian forms, but is also found in early African representatives. The short zygomatic arch and the relatively high position of the orbit can also be mentioned here.

Molar of Palaeoloxodon antiquus

Palaeoloxodon's teeth were similar to those of other elephants. It consisted of the two upper tusks and six molars per jaw arch, which included three deciduous premolars and three molars . As with all other elephants, the molars changed horizontally , so that there were a total of six generations of teeth. The molars had a lamellodontic tooth structure, that is, the tooth was composed of individual enamel lamellae , which were integrated in a matrix of dental cement . The edges of the enamel lamellae ran largely parallel to one another. Occasionally, however, diamond-shaped folds of the tooth enamel appeared in the middle of each lamella , which are somewhat reminiscent of the pattern of the African elephant. The first two molars consisted of seven to eleven lamellae, the last one of eleven to 19. The number of lamellae increased in the evolutionary history . In contrast, the enamel thickness of each fold decreased from 3 to 4 mm in early forms down to 1 mm in late forms. This also led to the fact that the lamellar frequency (number of enamel lamellae per 10 cm tooth length) increased from 3.8 to 4.8 in the case of more original members to 4.4 to 7.7 in more modern relatives. Generally the molars were high crowned ( hypsodont ). The height of the teeth sometimes exceeded their width by 150%. The longest tooth, the last molar, was up to 34 cm long in large individuals.

distribution

The representatives of Palaeoloxodon were spread over large parts of Africa and Eurasia . The origin lies in Africa, where the genus first appeared in the transition from the Miocene to the Pliocene around 5 million years ago. She was also present in Eurasia at the latest since the Lower Pleistocene , whereby the emigration probably took place via the Levant . Evidence of paleoloxodon is available from Europe as well as from West , Central , South and East Asia . Coming from East Asia, the elephant representative then also reached Japan . In contrast to the representatives of the genus Mammuthus native to Eurasia at about the same time , Palaeoloxodon never entered the American continent. Palaeoloxodon used largely wooded, warm-temperate landscapes. As a result, its occurrence in western Eurasia was largely limited to the warm periods under the influence of the changing climatic conditions of the Ice Age . In the cold phases in between , Palaeoloxodon was replaced by Mammuthus . The latter was already present in Eurasia with Mammuthus meridionalis at least since the end of the Pliocene and initially colonized similar landscapes as the newly immigrated Palaeoloxodon . The clear ecological differentiation between Palaeoloxodon and Mammuthus could therefore be a result of an initially existing competition between the two representatives of the proboscis, as a result of which a clear niche separation developed. A similar ecological sequence can also be demonstrated for the Upper Pleistocene with the last glaciation phase in Japan, where Palaeoloxodon is mainly detectable in the warmer interstadials , whereas Mammuthus primigenius can be found in the colder ones .

Paleobiology

Live reconstruction of Palaeoloxodon antiquus (European forest elephant)

The construction of the molars with their moderately high crowns distinguishes paleoloxodon as a predominant consumer of mixed vegetable foods. For Palaeoloxodon antiquus from western Eurasia, this is also proven by stomach contents and plant remains on the teeth. As a result, a wide variety of woody plants such as maple , hornbeam , hazelnut , alder , ash , beech and ivy as well as herbs such as artemisia and grasses such as sedges could be identified as food plants. Investigations into the traces of tooth wear of Palaeoloxodon antiquus gave a similar picture . There were also individual differences, as in some animals, due to tooth abrasion, there was a certain dominance of leaves or grasses. It is conceivable that, as with today's elephants, there were locally and seasonally different food compositions. Isotope analyzes on teeth of Palaeoloxodon recki led to basically comparable results . In contrast to most of the other representatives of Palaeoloxodon , Palaeoloxodon jolensis , also from Africa, had extremely high molars. Here isotope analyzes revealed a dominance of grasses in the food spectrum. Similar to today's elephants, there was also a great need for water.

Systematics

Internal systematics of the elephants according to Meyer et al. 2017
 Elephantidae  


 Loxodonta


   

 Paleoloxodon



   

 Elephas


   

 Mammothus




Template: Klade / Maintenance / Style

Palaeoloxodon is a genus from the elephant family (Elephantidae). It thus forms part of the modern line of development from the recent radiation phase of the order of mammoths (Proboscidea). The genus has a certain variety of shapes. Both gigantic representatives as well as dwarfed species are documented, which were distributed over large areas of Eurasia and Africa . Like all elephants, the representatives of Palaeoloxodon are characterized by molars consisting of individual lamellas ( lamellodontic tooth structure) that are changed horizontally , the loss of the lower jaw tusks and an upper skull consisting of air-filled chambers. From today's systematic point of view, Palaeoloxodon is more closely related to the African elephant ( Loxodonta ) than to the Asian elephant ( Elephas ), which has been proven above all by molecular genetics . The genus Loxodonta , which currently comprises two species with the African elephant ( Loxodonta africana ) and the forest elephant ( Loxodonta cyclotis ), is often referred to in a tribe called Loxodontini .

The following types are generally recognized today:

Often Palaeoloxodon namadicus is given as the nominate form , the species was introduced in 1847 together with Palaeoloxodon antiquus . However, Palaeoloxodon naumanni represents the actual type species. The reason for the discrepancy is that Palaeoloxodon naumanni was listed as a subspecies of Palaeoloxodon namadicus for a long time and was also specified by Matsumoto. In later analyzes, however, it turned out to be a separate form.

Regardless of the species, two morphotypes were mainly distinguished for the Eurasian mainland forms. One is an animal with a rather delicate parietal-occipital bone bulge on the skull, which is called "Stuttgart-Morphe", named after a skull from the travertines near Bad Cannstatt . The “Stuttgart-Morphe” accordingly included a large part of the finds of Palaeoloxodon antiquus , but also of Palaeoloxodon turkmenicus and Palaeoloxodon naumanni . The second type had a very robust parietal-occipital bone bulge and is called " namadicus morph". It was the species Palaeoloxodon namadicus godfather. However, in further research, published in 2020, these two morphotypes did not turn out to be special groups of shapes. Rather, the shape of the parietal-occipital bone bulge reflects, among other things, an individual stage of development, but can also indicate a sexual dimorphism to a certain extent .

Tribal history

Lower jaw of Palaeoloxodon recki

The earliest finds of the genus Palaeoloxodon came to light in Africa . Here in East Africa Palaeoloxodon ekorensis is the oldest species in the transition from Miocene to Pliocene around 5 million years ago (an older form called Palaeoloxodon nawataensis was dissolved in 2010 and equated with other early elephant forms). The species was named after the Ekora beds in Kenya , from where the type material consisting of the rear molars comes from. The teeth of the mold were not as clearly high-crowned as in later representatives and had a low frequency of lamellae (3 8 to 4.8). The tusks were also more parallel to each other and far apart, and the parietal-occipital bone bulge was generally poorly developed. The most important African representative is Palaeoloxodon recki . This also occurred in the Lower Pliocene and was mainly present in eastern Africa. With its moderately high-crowned molars and higher lamellar frequency of 4 to 5.5, Palaeoloxodon recki was a bit more modern than its predecessor. The gigantic proboscis was found at important stations such as Olduvai in Tanzania and Olorgesailie in Kenya, but also in Barogali in Djibouti , where a dismantled skeleton was discovered in association with numerous stone tools . There are also individual finds from southern and central Africa. Originally five subspecies of Palaeoloxodon recki were defined, but these could not be clearly confirmed in recent studies. Around 1 million years ago in the course of the Lower Pleistocene Palaeoloxodon recki disappeared again and was replaced by Palaeoloxodon jolensis , of which only dental finds are available so far. Of all African elephant forms , Palaeoloxodon jolensis had the teeth with the highest tooth crowns with a lamella frequency of up to 6.3. The species was found in much of Africa including the northern part, but was not very common. Their last evidence dates back to the transition from the Middle to the Upper Pleistocene around 130,000 years ago.

Skeletal reconstruction of Palaeoloxodon antiquus

In Eurasia , Palaeoloxodon appeared for the first time at the end of the Lower Pleistocene, the immigration possibly took place via the Levant . It is often assumed that they are descendants of Palaeoloxodon recki . The Western Eurasian finds are generally assigned to Palaeoloxodon antiquus , which is also the best-known species, commonly referred to as the European forest elephant. Early finds in Europe come from the southern continental area such as Isernia la Pineta and Slivia in Italy , possibly also from the southwest. The European forest elephant probably did not reach the northern Alpine areas before the younger Cromer complex around 600,000 years ago, as is shown by the finds from the Mosbacher Sands in Hesse and from Pakefield in England, while in the slightly older deposits of Voigtstedt and Süßenborn , both Thuringia, is missing. At the latest in the late Middle Pleistocene, the species was quite common. The various travertine deposits in Central Europe such as Bad Cannstatt in Baden Württemberg, Bilzingsleben as well as Ehringsdorf and Taubach near Weimar, all of Thuringia, are among the most important sites of that time . From a historical research point of view, the travertines of Burgtonna , again Thuringia, where the first fossils of a European forest elephant were found in 1695, are important. The interpretation of these remains as the remains of an elephant on the one hand, and as a "mineral plant" or "fossil unicorn" on the other hand led to a scholarly dispute that lasted for years. The Geiseltal in Saxony-Anhalt is also of outstanding importance , where several complete skeletons were uncovered in lakeshore deposits from the end of the Middle Pleistocene. Find sites far to the north can be found in England and Denmark, the northernmost known distribution limit in Russia is reached around 55 degrees north. Palaeoloxodon antiquus disappeared from the Western Eurasian landscapes at the latest before the maximum of the last glacial period around 35,000 years ago . The most recent finds came to light in south-western Europe.

Skull of Palaeoloxodon turkmenicus

In eastern Eurasia, Palaeoloxodon namadicus then replaced Palaeoloxodon antiquus , a relatively modern form with narrow molars, the last of which had 12 to 19 enamel ridges and a lamella frequency of up to 7.7. It is probably the largest representative of the genus, but its physique was not quite as compact as that of its Western Eurasian cousins, which is documented very well by the slimmer limbs. Both Palaeoloxodon namadicus and Palaeoloxodon antiquus received their first scientific description in 1847 , both forms were for a time synonymous with the former due to the first mentioning as a valid species. Today, the separation of the two on a species level is hardly disputed. Palaeoloxodon namadicus is also documented for the first time in the Lower Middle Pleistocene; numerous finds were made in the valley of the eponymous river Narmada . An almost complete skeleton was reported from the Godavari valley in 1905, the reconstructed thighbone of around 165 cm long and belonging to an animal with a shoulder height of 450 cm. Individual finds, which were discovered in the 19th century and are sometimes very fragmented, such as those from Sagauni, suggest even larger individuals. Palaeoloxodon namadicus was still present in the region at least until the Upper Pleistocene. Young data from around 56,000 years ago were obtained from the sites of a skull found in the Dhasan river basin in the peripheral catchment area of ​​the Ganges . From Kuday-Dag in Turkmenistan the type skull of Palaeoloxodon turkmenicus came to light , which has only a weakly developed parietal-occipital bone bulge. This form is generally considered to have been little studied, but is often considered to be identical to the European forest elephant. However, it could also represent a separate species, which requires further investigation.

Skeletal reconstruction of Palaeoloxodon naumanni

The Japanese island world was populated in the Middle and Upper Pleistocene by Palaeoloxodon naumanni , a comparatively small representative that reached shoulder heights of up to 2.8 m in males and 1.9 m in females. As for the molars, it was more modern. The last molars usually had 18 to 19 lamellae and a lamella frequency of 5.2 to 6.2. Its name dates back to 1924 and is based on teeth found near Shizuoka . Finds of Palaeoloxodon naumanni scatter over large areas of Japan up to the north island of Hokkaido . The Nojiri Lake in the central Japanese prefecture of Nagano is of outstanding importance . Numerous remains of Palaeoloxodon naumanni in association with human stone artifacts were found here on an area of ​​several thousand square meters . The finds belong to the middle section of the last glacial period . The used habitat of the species consisted of coniferous forests. The most recent finds of Palaeoloxodon naumanni are around 23,000 years old, so the species apparently died out before the last cold peak. Some authors also consider all East Asian Palaeoloxodon mainland forms to belong to Palaeoloxodon naumanni . They were initially referred to the subspecies P. n. Huaihoensis in the 1970s with the description of a partial skeleton from Huaiyuan in the east Chinese province of Anhui , but this received species status a good 20 years later. The opposite is the case with Palaeoloxodon tokunagai , which was named in 1929 on the basis of teeth from the central Japanese province of Etchū , but was subsequently united with Palaeoloxodon naumanni . In general, the East Asian finds of palaeoloxodon require extensive revision.

Skeletal reconstruction of Palaeoloxodon falconeri
Molars of Palaeoloxodon lomolinoi

During the Pleistocene, several dwarfed species emerged on various islands in the Mediterranean , each of which is considered to be the European forest elephant. This island dwarfing has progressed differently in the individual species. The best-known form is likely to be Palaeoloxodon falconeri or the Sicilian pygmy elephant, which has been found in Sicily and Malta . Both islands reached the ancestors of Palaeoloxodon falconeri during the Middle Pleistocene. They then developed an extreme dwarfing effect, which led to the smallest known elephant species. For male animals, a shoulder height of 1 m and a weight of around 300 kg are reconstructed, corresponding values ​​for female animals are 0.8 m and 165 kg. The dwarfism also influenced the general physique. The skull of Palaeoloxodon falconeri is much more rounded and the limbs are more reminiscent of those of not fully grown elephants with a less strictly columnar structure. Compared to the mainland forms, Palaeoloxodon falconeri also had a significantly larger brain in relation to body weight. The Spinagallo cave in south-east Sicily is one of the most important sites with more than 3000 bone elements , and Għar Dalam in Malta should not go unmentioned . Palaeoloxodon falconeri was scientifically named in 1869 by George Busk based on finds from Malta. At the end of the Middle Pleistocene, another wave of colonization hit Sicily and Malta, in whose Zuig Palaeoloxodon mnaidriensis arose. This species was not quite as dwarfed as Palaeoloxodon falconeri . The animals measured about 1.8 to 2 m at the shoulder and weighed between 1.1 and 2.5 t. Possibly a third species, Palaeoloxodon melitensis , was intermediate in size between the other two. It was also named by Busk in 1869, but is considered to be partially identical to Palaeoloxodon falconeri . The other dwarf forms are mainly native to the eastern Mediterranean. They mainly affect the Aegean islands of the Dodecanese and the Cyclades . Among other things, Palaeoloxodon tiliensis from Tilos , a 1.4 m high and 650 kg heavy form that was scientifically introduced in 2007, is of importance. Some recent finds of the form are only 3300 years old and are therefore among the last records of the genus Palaeoloxodon . In contrast, the findings from Naxos in 2014 were described as Palaeoloxodon lomolinoi . The animals were only around 10% the size of a European forest elephant. Remains of elephants from other Aegean islands have not yet been described. As in Sicily and Malta, at least two differently sized forms of palaeoloxodon occurred in Cyprus . One is Palaeoloxodon cypriotes , which weighed only around 250 kg and was therefore extremely small, the larger one was given the scientific name Palaeoloxodon xylophagou in 2015 . While Palaeoloxodon cypriotes appeared in the Upper Pleistocene and possibly only disappeared in the Lower Holocene , the remains of Palaeoloxodon xylophagou are middle-Pleistocene. On Crete, in turn, Palaeoloxodon creutzburgi was at home in the Upper Pleistocene , a comparatively large form with an estimated 3 t body weight, the main location of which is the Kaló Chorafi cave in the north of the island. Possibly the species reached the neighboring island of Kassos to the east . A second form from Crete, Palaeoloxodon chaniensis , slightly larger than Palaeoloxodon creutzburgi and introduced in 2000, is controversial in its taxonomic status. The island of Kefalonia in the Gulf of Patras and belonging to the Ionian Islands housed a form that was introduced in 2018 as Palaeoloxodon cephallonicus in the transition from the Middle Pleistocene . At the moment only a single upper jaw, enclosed in a rock matrix, has survived, which suggests an animal the size of Palaeoloxodon creutzburgi or Palaeoloxodon mnaidriensis .

Regardless of the often divided opinion that the continental populations of Palaeoloxodon became extinct at the end of the Pleistocene, Chinese scientists published a study in 2012 in which they struck two teeth from Dingjiabu in the Chinese province of Hebei of the genus and postulated an age in the Middle Holocene. As further evidence of the survival of Palaeoloxodon up to this time, they cited several bronze cauldrons with elephant representations, which belong to the period of the Xia to Zhou dynasties 4100 to 2300 years ago. The representations of the elephants led the study's authors to assume that the trunk had two finger-like appendages, while the Asian elephant had only one. However, these are highly stylized representations that do not allow any taxonomic statements. New examinations of the elephant teeth refer to remains of the Asian elephant with the high number of enamel lamellae (23) and the high lamella frequency (7). In addition, radiometric age dating of the teeth showed that the teeth were in the middle phase of the last glacial around 50,300 years ago and are therefore much older than originally assumed.

Research history

The closer relationship of palaeoloxodon was discussed for a long time. The genus was scientifically introduced in 1924 by Hikoshichirō Matsumoto . In his on Japanese held first description Matsumoto took palaeoloxodon as a subgenus of the African elephant ( Loxodonta on), whereupon also the scientific name indicates. He united Palaeoloxodon namadicus and Palaeoloxodon antiquus within the new subgenus. Henry Fairfield Osborn followed Matsumoto's view in the 1930s. However, he raised Palaeoloxodon at the genus level and separated from it Hesperoloxodon , to which he attributed the Western Eurasian finds. He distinguished an eastern group around Palaeoloxodon namadicus and a western one around Palaeoloxodon antiquus , the differentiating criterion of which was essentially expressed in the different skull design. George Gaylord Simpson repealed this in his general taxonomy of mammals in 1945 and again classified Palaeoloxodon as a subgenus of Loxodonta .

At the end of the 1960s, Emiliano Aguirre first presented the African representatives of Palaeoloxodon as precursors to the Eurasian. He also advocated a closer relationship between Palaeoloxodon and Loxodonta . In contrast, Vincent J. Maglio , who saw Palaeoloxodon much closer to Elephas in an essay published almost at the same time . In his extensive overview of the development of the Elephantidae from 1973, Maglio then combined Palaeoloxodon with Elephas , the genus of the Asian elephant, and listed the former as a subgenus. The common feature was the parietal-occipital bone bulge, which occurs more or less clearly in Elepahs and Palaeoloxodon , but is absent in Loxodonta . Maglio's decision persisted for the next four decades and was shared by numerous scholars.

The picture only changed again with the inclusion of molecular genetic studies, which were presented in 2017 and which included fossils of the European forest elephant in addition to the recent elephant species. These showed a closer relationship with Loxodonta , with a closer connection to the forest elephant ( Loxodonta cyclotis ) than to the African elephant ( Loxodonta africana ). Palaeoloxodon is therefore again largely run as an independent genus. In-depth analyzes also revealed a far more complex relationship between the various elephant species. According to this, there are genetic similarities within the various elephant groups, these are composed of three components: The first concerns alleles that are shared by the African elephant and the (African) forest elephant and which possibly go back to the parent group of the two species. As a second component, the European forest elephant and the West African forest elephant have a significant number of common gene components. Ultimately, however, genetic similarities with the woolly mammoth ( Mammuthus primigenius ) could also be shown. Various hybridization events in the early phase of the phylogenetic development of the individual elephant forms can be assumed to be the cause of this gene mixture . Both palaeoloxodon and Loxodonta and Mammuthus emerged each in Africa, where all three species in the Pliocene before about 5 million years are detectable. Hybridization then took place in a time phase before the genetic isolation of the individual species began to take effect.

literature

  • Asier Larramendi, Hanweng Zhang, Maria Rita Palombo and Marco P. Ferretti: The evolution of Palaeoloxodon skull structure: Disentangling phylogentic, sexually dimorphic, ontogenetic, and allometric morphological signals. Quaternary Science Review 229, 2020, p. 106090, doi: 10.1016 / j.quascirev.2019.106090
  • Jan van der Made: The evolution of the elephants and their relatives in the context of a changing climate and geography. In: Harald Meller (ed.): Elefantenreich. A fossil world in Europe. Halle / Saale 2010, pp. 340-360

Individual evidence

  1. ↑ a b Asier Larramendi: Shoulder height, body mass, and shape of proboscideans. Acta Palaeontologia Polonica 61 (3), 2016, pp. 537-574
  2. ^ Per Christiansen: Body size in proboscideans, with notes on elephant metabolism. Zoological Journal of the Linnean Society 140, 2004, pp. 523-549
  3. ↑ a b c d e f g Maria Rita Palombo: Elephants in miniature. In: Harald Meller (Hrsg.): Elefantenreich - Eine Fossilwelt in Europa. Halle / Saale 2010, pp. 275–295
  4. ↑ a b c d Sevket Sen Eric Barrier and Xavier Crété: Late Pleistocene Dwarf Elephants from the Aegean Islands of Kassos and Dilos, Greece. Annales Zoologici Fennici 51 (1/2), 2014, pp. 27-42
  5. ↑ a b c d Sevket Sen: A review of the Pleistocene dwarfed elephants from the Aegean islands, and their paleogeographic context. Fossil Imprint 73 (1/2), 2017; Pp. 76-92
  6. ↑ a b c Athanassios Athanassiou, Alexandra AE van der Geer and George A. Lyras: Pleistocene insular Proboscidea of ​​the Eastern Mediterranean: A review and update. Quaternary Science Reviews 218, 2019, pp. 306-321
  7. ↑ a b c d e Haruo Saegusa and W. Henry Gilbert: Elephantidae. In: W. Henry. Gilbert and Berhane Asfaw (eds.): Homo erectus: Pleistocene Evidence from the Middle Awash, Ethiopia. Berkeley, Los Angeles, 2008, pp. 193-226
  8. ↑ a b c d e Asier Larramendi, Hanweng Zhang, Maria Rita Palombo and Marco P. Ferretti: The evolution of Palaeoloxodon skull structure: Disentangling phylogentic, sexually dimorphic, ontogenetic, and allometric morphological signals. Quaternary Science Review 229, 2020, p. 106090, doi: 10.1016 / j.quascirev.2019.106090
  9. ^ A b c Vincent J. Maglio: Origin and Evolution of the Elephantidae. Transactions of the American Philosophical Society 63 (3), 1973, pp. 1–149 (pp. 31–50)
  10. ↑ a b c William J. Sanders, Emmanuel Gheerbrant, John M. Harris, Haruo Saegusa, Cyrille Delmer: Proboscidea. In: Lars Werdelin, William Joseph Sanders (eds.): Cenozoic Mammals of Africa. University of California Press, Berkeley / London / New York 2010, pp. 161–251
  11. ^ A b c d Adrian Lister: Ecological interactions of elephantids in Pleistocene Eurasia: Palaeoloxodon and Mammuthus. In: N. Goren-Inbar and JD Speth (Eds.): Human Paleoecology in the Levantine Corridor. Oxford, 2004, pp. 53-60
  12. ↑ a b Florent Rivals, Gina Semprebon and Adrian Lister: An examination of dietary diversity patterns in Pleistocene proboscideans (Mammuthus, Palaeoloxodon, and Mammut) from Europe and North America as revealed by dental microwear. Quaternary International 255, 2012, pp. 188-195
  13. ↑ a b c Keiichi Takahashi, Yuji Soeda, Masami Izuho, ​​Kaori Aoki, Goro Yamada and Mono Akamatsu: A New Specimen of Palaeoloxodon naumanni from Hokkaido and its Significance. Quaternary Research 43 (3), 2004, pp. 169-180
  14. ↑ a b Keiichi Takahashi, Yuji Soeda, Masami Izuho, ​​Goro Yamada, Morio Akamatsu and Chun-Hsiang Chang: The chronological record of the woolly mammoth (Mammuthus primigenius) in Japan, and its temporary replacement by Palaeoloxodon naumanniduring MIS 3 in Hokkaido (northern Japan). Palaeogeography, Palaeoclimatology, Palaeoecology 233, 2006, pp. 1-10
  15. ↑ René Grube: Vegetable food remains of the fossil elephants and rhinos from the interglacial of Neumark-Nord. In: Jan Michal Burdukiewicz, Lutz Fiedler, Wolf-Dieter Heinrich, Antje Justus and Enrico Brühl (eds.): Knowledge hunters . Festschrift for Dietrich Mania. Publications of the State Museum for Prehistory in Halle 57 Halle / Saale, 2003, pp. 221–236
  16. ^ René Grube, Maria Rita Palombo, Paola Iacumin and Antoinette Di Matteo: What did the fossil elephants from Neumark-Nord eat? In: Harald Meller (Hrsg.): Elefantenreich - Eine Fossilwelt in Europa. Halle / Saale, 2010, pp. 253-273
  17. ↑ MR Palombo, ML Filippi, P. Iacumin, A. Longinelli, M. Barbieri and A. Maras: Coupling tooth microwear and stable isotope analyzes for palaeodiet reconstruction: the case study of Late Middle Pleistocene Elephas (Palaeoloxodon) antiquus teeth from Central Italy (Rome area). Quaternary International 126-128, 2005, pp 153-170
  18. ↑ Thure E. Cerling, John M. Harris and Meave G. Leakey: Browsing and grazing in elephants: the isotope record of modern and fossil proboscideans. Oecologia 120, 1999, pp. 364-374
  19. ↑ a b Fredrick Kyalo Manthi, William J. Sanders, J. Michael Plavcan, Thure E. Cerling and Francis H. Brown: Late Middle Pleistocene Elephants from Natodomeri, Kenya and the Disappearance of Elephas (Proboscidea, Mammalia) in Africa. Journal of Mammalian Evolution, 2019, doi: 10.1007 / s10914-019-09474-9
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