Palaeoloxodon recki

Palaeoloxodon recki; Temporal range: Late Pliocene–Middle Pleistocene PreꞒ Ꞓ O S D C P T J K Pg N ↓
	Size comparison of a 40 year old adult male Palaeoloxodon recki atavus from Koobi Fora
	Life restoration by Mauricio Antón
Scientific classification
Domain:	Eukaryota
Kingdom:	Animalia
Phylum:	Chordata
Class:	Mammalia
Order:	Proboscidea
Family:	Elephantidae
Genus:	† Palaeoloxodon
Species:	†P. recki
Binomial name
	†Palaeoloxodon recki; (Dietrich, 1894)
Synonyms
	Elephas recki Dietrich, 1894

Last updated March 24, 2024

Palaeoloxodon recki, often known by the synonym Elephas recki is an extinct species of elephant native to Africa and West Asia from the Pliocene or Early Pleistocene to the Middle Pleistocene. During most of its existence, the species (in its broad sense) represented the dominant elephant species in East Africa.^[1] The species is divided into five roughly chronologically successive subspecies. While the type and latest subspecies P. recki recki as well as the preceding P. recki ileretensis are widely accepted to be closely related to Eurasian Palaeoloxodon, the relationships of the other, chronologically earlier subspecies to P. recki recki and P. recki ileretensis are uncertain, with it being suggested they are unrelated and should be elevated to separate species.

Taxonomy

The species was initially named from specimens found at Bed IV in Olduvai Gorge, Tanzania by Wilhelm Otto Dietrich in 1915, originally as a subspecies of the European straight-tusked elephant, what is now called Palaeoloxodon antiquus, as Elephas antiquus recki.^[2] It was named after Hans Reck, a German paleontologist and geologist who had done the initial surveying of the gorge in 1913, and had collected a considerable number of fossils from the locality.^[3] Camille Arambourg in 1942 described additional specimens of the species from Omo Valley in Ethiopia, and suggested that they were distinctive enough that they warranted being placed as the distinct species E. recki. The two deposits are not contemporaneous and the specimens from each locality are morphologically distinctive from each other, which has led to confusion about which locality represents the "typical" morphology of the species.^[2] The placement of Elepas recki in the genus Elephas was contested as early as 1942, when in a publication by Douglas Gordon MacInnes and Henry Fairfield Osborn's posthumous monograph on fossil proboscideans it was suggested that Elephas recki should instead be placed in Palaeoloxodon, though many later authors continued to place the species in the genus Elephas, treating Palaeoloxodon as a subgenus of Elephas.^[3]

Michel Beden's publications during the 1980s on Elephas recki primarily focusing on molar morphology^[4]^[5]^[6] identified five successive subspecies, from oldest to youngest with ages according to Sanders (2023)^[3]:

E. r. brumpti Beden, 1980 Early Pliocene-~3.2 million years ago (mya)
E. r. shungurensis Beden, 1980 ~3.2-2.3 mya
E. r. atavus Arambourg, 1947 ~2.3-1.8 mya
E. r. ileretensis Beden, 1987 ~1.8-1.4/1.6 mya
E. r. recki (Dietrich, 1916) 1.6/1.4 mya-late Middle Pleistocene

For decades after Beden's publications, his view of "Elephas recki" as a succession of subspecies was accepted as orthodoxy. However, Beden's views of Elephas recki evolution were challenged by the work of Nancy Todd published in 2001 and 2005.^[2] Todd found that the different subspecies exhibited high variability in molar dimensions within subspecies, but low variability between the subspecies, and that not all of the subspecies clustered together with each other in cladistic analysis, as would be expected if they formed a single species, with E. r. brumpti E. r. shungurensis and E. r. atavus being placed separately from E. r. ileretensis andE. r. recki, and that the supposed subspecies substantially chronologically overlapped, though later work suggested that this chronological overlap was likely overstated.^[3]

A 2020 PhD thesis by Steven Zhang, focusing primarily on skull morphology again challenged the monophyly of Elephas recki as a whole, finding that while E. r. ileretensis and E. recki recki has a close relationship with Eurasian Palaeoloxodon. which genetic data shows is closely related to African elephants ( Loxodonta ), the other named E. recki subspecies were likely unrelated, and more closely related to true members of the genus Elephas (which contains the living Asian elephant). Zhang suggested that E. r. brumpti was synonymous with "Elephas" planifrons , primarily known from the Late Pliocene-Early Pleistocene of the Indian subcontinent, and that this species should be placed into Phanagoroloxodon , while E. r. atavus should be elevated to full species status as a true member of the genus Elephas. However, while Sanders (2023) accepted that E. r. brumpti was unlikely to be closely related to E. recki recki, he rejected the synonymity of E. r. brumpti with "Elephas" planifrons (though he suggested that the two may be sister species), and questioned their placement in Phanagoroloxodon, while he considered the relationship of E. r. atavus to E. recki recki to be uncertain.^[3]

Outside its core East African distribution. It has also been suggested that material from West Asia, including that from the earliest Middle Pleistocene (c. 780,000 years ago) Paleolithic archaeological site Gesher Bnot Ya'akov (otherwise attributed to P. antiquus) in northern Israel, the Middle Pleistocene (c. 500,000 years ago) Ti's al Ghadah site in northern Saudi Arabia, and the late Middle Pleistocene Shishan Marsh site in Jordan, belongs to P. recki recki.^[7]^[8]^[9]^[3]

Description

Members of the species were larger than any living elephant. A large mostly complete male specimen of P. recki atavus^[3] from Koobi Fora, Kenya, suggested to have been approximately 40 years old when it died, was estimated in a 2016 study to have measured 4.27 metres (14.0 ft) tall at the shoulder and weighed 12.3 tonnes (12.1 long tons; 13.6 short tons),^[10] with the tusks of some P. recki individuals reaching 4 metres (13 ft) in length, and masses likely considerably in excess of 100 kilograms (220 lb).^[11] In comparison to most Eurasian species of Palaeoloxodon, the parieto-occipital crest of P. recki recki at the top of the skull is only weakly developed. The frons (forehead) is tall and biconvex. Like Eurasian Palaeoloxodon species, the premaxillary bones of P. recki recki containing the tusks flare laterally outwards.^[7] Over time the molar teeth of P. recki show an increasing number of lamellae, and taller crown height (hypsodonty).^[12]

Ecology

All named subspecies of P/E. recki, regardless of true evolutionary relationships, are thought to have been dedicated grazers, with the molar teeth of later subspecies showing greater adaption to grazing than earlier subspecies.^[1]

Evolutionary history

Following the emergence of P. recki in Africa, at the end of the Early Pleistocene, around 800,000 years ago, a population of P. recki recki migrated out of Africa, giving rise to the Eurasian radiation of Palaeoloxodon.^[13] Its descendant taxon or last evolutionary stage, Palaeoloxodon iolensis , is known from remains found across Africa of late Middle Pleistocene to Late Pleistocene age. Following the extinction of P. iolensis it was replaced by the modern African bush elephant (Loxodonta africana).^[1]

Relationship with humans

At several sites across Africa, remains of P. recki have been found associated with stone tools. In some cases like Olduvai FLK, these are likely coincidental, but in others which bears cut marks, these likely represent evidence of butchery by archaic humans. Sites containing P. recki remains with cut marks and/or stone tools include Upper Bed II at the Bell's Korongo site in Olduvai Gorge, dating to around 1.35 million years ago, which has been suggested to be the oldest site in the world with reliable evidence of elephant butchery, associated with Oldowan-type stone tools, and the Olorgesailie Basin Member 1 Site 15 in Kenya, dated to 992–974,000 years ago, and the Nadung’a 4 site near Lake Turkana, Kenya, dating to approximately 700,000 years ago. The Barogali site in Djibouti, dating to 1.6-1.3 million years ago, where a disassociated specimen of P. recki (suggested to be E. recki atavus^[3]) was found with numerous stone tools (probably Oldowan) created onsite, has also been suggested to be a butchery site. The P. antiquus/P. recki specimen from Gesher Bnot Ya'akov is associated with an Acheulean stone handaxe and other bifaced tools, and displays cut marks and fracture marks indicative of butchery, though the fracturing of the skull, which has been suggested to be the result of an attempt to extract the brain, may alternatively be the result of postmortem trampling.^[14]

Gallery

Skull at the Gallery of Paleontology and Comparative Anatomy, France
Underside of the skull and lower jaws at the Gallery of Paleontology and Comparative Anatomy
Lower jaw at the Museum für Naturkunde. Berlin

Related Research Articles

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Proboscidea is a taxonomic order of afrotherian mammals containing one living family (Elephantidae) and several extinct families. First described by J. Illiger in 1811, it encompasses the elephants and their close relatives. Proboscideans include some of the largest known land mammals. The largest land mammal of all time may have been a proboscidean; the elephant Palaeoloxodon namadicus has been estimated to be up to 5.2 m (17.1 ft) at the shoulder and may have weighed up to 22 t, surpassing the paraceratheres, the otherwise largest known land mammals, though this estimate was made based on a single fragmentary femur and is speculative. The largest extant proboscidean is the African bush elephant, with a record of size of 4 m (13.1 ft) at the shoulder and 10.4 t. In addition to their enormous size, later proboscideans are distinguished by tusks and long, muscular trunks, which were less developed or absent in early proboscideans.

A mastodon is a member of the genus Mammut, which strictly defined, was endemic to North America and lived from the late Miocene to the early Holocene. Mastodons belong to the order Proboscidea, the same order as elephants and mammoths. Mammut is the type genus of the extinct family Mammutidae, which diverged from the ancestors of modern elephants at least 25 million years ago, during the Oligocene.

Elephantidae is a family of large, herbivorous proboscidean mammals collectively called elephants and mammoths. These are large terrestrial mammals with a snout modified into a trunk and teeth modified into tusks. Most genera and species in the family are extinct. Only two genera, Loxodonta and Elephas, are living.

Elephas is one of two surviving genera in the family of elephants, Elephantidae, with one surviving species, the Asian elephant, Elephas maximus. Several extinct species have been identified as belonging to the genus, extending back to the Pliocene era.

Palaeoloxodon is an extinct genus of elephant. The genus originated in Africa during the Early Pleistocene, and expanded into Eurasia at the beginning of the Middle Pleistocene. The genus contains some of the largest known species of elephants, over 4 metres (13 ft) tall at the shoulders, including the African Palaeoloxodon recki, the European straight-tusked elephant and the South Asian Palaeoloxodon namadicus. P. namadicus has been suggested to be the largest known land mammal by some authors based on extrapolation from fragmentary remains, though these estimates are highly speculative. In contrast, the genus also contains many species of dwarf elephants that evolved via insular dwarfism on islands in the Mediterranean, some only 1 metre (3.3 ft) in height, making them the smallest elephants known. The genus has a long and complex taxonomic history, and at various times, it has been considered to belong to Loxodonta or Elephas, but today is usually considered a valid and separate genus in its own right.

Dwarf elephants are prehistoric members of the order Proboscidea which, through the process of allopatric speciation on islands, evolved much smaller body sizes in comparison with their immediate ancestors. Dwarf elephants are an example of insular dwarfism, the phenomenon whereby large terrestrial vertebrates that colonize islands evolve dwarf forms, a phenomenon attributed to adaptation to resource-poor environments and lack of predation and competition.

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Mammuthus meridionalis, sometimes called the southern mammoth, is an extinct species of mammoth native to Eurasia, including Europe, during the Early Pleistocene, living from around 2.5 million years ago to 800,000 years ago.

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<i>Sinomastodon</i> Extinct genus of gomphothere proboscidean

Sinomastodon is an extinct gomphothere genus known from the Late Miocene to Early Pleistocene of Asia, including China, Japan, Thailand, Myanmar, Indonesia and probably Kashmir.

Palaeoloxodon namadicus is an extinct species of prehistoric elephant known from the early Middle to Late Pleistocene of the Indian subcontinent, and possibly also elsewhere in Asia. The species grew larger than any living elephant, and some authors have suggested it to have been the largest known land mammal based on extrapolation from fragmentary remains, though these estimates are speculative.

Palaeoloxodon cypriotes is an extinct species of dwarf elephant that inhabited the island of Cyprus during the Late Pleistocene. Remains comprise 44 molars, found in the north of the island, seven molars discovered in the south-east, a single measurable femur and a single tusk among very sparse additional bone and tusk fragments. The molars support derivation from the large straight-tusked elephant (Palaeoloxodon antiquus). The species is presumably derived from the older, larger P. xylophagou from the late Middle Pleistocene which reached the island presumably during a Pleistocene glacial maximum when low sea levels allowed a low probability sea crossing between Cyprus and Asia Minor. During subsequent periods of isolation the population adapted within the evolutionary mechanisms of insular dwarfism, which the available sequence of molar fossils confirms to a certain extent. The fully developed Palaeoloxodon cypriotes weighed not more than 200 kg (440 lb) and had a height of around 1 m (3.28 ft). The species became extinct around 12,000 years ago, around the time humans first colonised Cyprus.

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<i>Phanagoroloxodon</i> Extinct genus of mammals

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References

1 2 3 Manthi, Fredrick Kyalo; Sanders, William J.; Plavcan, J. Michael; Cerling, Thure E.; Brown, Francis H. (September 2020). "Late Middle Pleistocene Elephants from Natodomeri, Kenya and the Disappearance of Elephas (Proboscidea, Mammalia) in Africa". Journal of Mammalian Evolution. 27 (3): 483–495. doi:10.1007/s10914-019-09474-9. ISSN 1064-7554. S2CID 198190671.
1 2 3 Todd, Nancy E. (January 2005). "Reanalysis of African Elephas recki: implications for time, space and taxonomy". Quaternary International. 126–128: 65–72. doi:10.1016/j.quaint.2004.04.015.
1 2 3 4 5 6 7 8 Sanders, William J. (2023-07-07). Evolution and Fossil Record of African Proboscidea (1 ed.). Boca Raton: CRC Press. pp. 267–293. doi:10.1201/b20016. ISBN 978-1-315-11891-8.
↑ Beden, M. 1980. Elephas recki Dietrich, 1915 (Proboscidea, Elephantidae). Èvolution au cours du Plio-Pléistocène en Afrique orientale. Geobios 13(6): 891-901. Lyon.
↑ Beden, M. 1983. Family Elephantidae. In J. M. Harris (ed.), Koobi Fora Research Project. Vol. 2. The fossil Ungulates: Proboscidea, Perissodactyla, and Suidae: 40-129. Oxford: Clarendon Press.
↑ Beden, M. 1987. Les faunes Plio-Pléistocène de la basse vallée de l’Omo (Éthiopie), Vol. 2: Les Eléphantidés (Mammalia-Proboscidea) (directed by Y. Coppens and F. C. Howell): 1-162. Cahiers de Paléontologie-Travaux de Paléontologie est-africaine. Centre National de la Recherche Scientifique (CNRS). Paris.
1 2 Larramendi, Asier; Zhang, Hanwen; Palombo, Maria Rita; Ferretti, Marco P. (February 2020). "The evolution of Palaeoloxodon skull structure: Disentangling phylogenetic, sexually dimorphic, ontogenetic, and allometric morphological signals". Quaternary Science Reviews. 229: 106090. Bibcode:2020QSRv..22906090L. doi:10.1016/j.quascirev.2019.106090. S2CID 213676377.
↑ Stimpson, Christopher M.; Lister, Adrian; Parton, Ash; Clark-Balzan, Laine; Breeze, Paul S.; Drake, Nick A.; Groucutt, Huw S.; Jennings, Richard; Scerri, Eleanor M.L.; White, Tom S.; Zahir, Muhammad; Duval, Mathieu; Grün, Rainer; Al-Omari, Abdulaziz; Al Murayyi, Khalid Sultan M. (July 2016). "Middle Pleistocene vertebrate fossils from the Nefud Desert, Saudi Arabia: Implications for biogeography and palaeoecology". Quaternary Science Reviews. 143: 13–36. doi:10.1016/j.quascirev.2016.05.016. hdl: 10072/142575 .
↑ Pokines, James T.; Lister, Adrian M.; Ames, Christopher J. H.; Nowell, April; Cordova, Carlos E. (March 2019). "Faunal remains from recent excavations at Shishan Marsh 1 (SM1), a Late Lower Paleolithic open-air site in the Azraq Basin, Jordan". Quaternary Research. 91 (2): 768–791. doi:10.1017/qua.2018.113. ISSN 0033-5894.
↑ Larramendi, A. (2016). "Shoulder height, body mass and shape of proboscideans" (PDF). Acta Palaeontologica Polonica. 61. doi: 10.4202/app.00136.2014 .
↑ Larramendi, Asier (2023-12-10). "Estimating tusk masses in proboscideans: a comprehensive analysis and predictive model". Historical Biology: 1–14. doi:10.1080/08912963.2023.2286272. ISSN 0891-2963.
↑ Lister, Adrian M. (2013-06-26). "The role of behaviour in adaptive morphological evolution of African proboscideans". Nature. 500 (7462): 331–334. Bibcode:2013Natur.500..331L. doi:10.1038/nature12275. ISSN 0028-0836. PMID 23803767. S2CID 883007.
↑ Lister, Adrian M. (2004), "Ecological Interactions of Elephantids in Pleistocene Eurasia", Human Paleoecology in the Levantine Corridor, Oxbow Books, pp. 53–60, ISBN 978-1-78570-965-4 , retrieved 2020-04-14
↑ Haynes, Gary (March 2022). "Late Quaternary Proboscidean Sites in Africa and Eurasia with Possible or Probable Evidence for Hominin Involvement". Quaternary. 5 (1): 18. doi: 10.3390/quat5010018 . ISSN 2571-550X.

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[:1-1] 1 2 3 Manthi, Fredrick Kyalo; Sanders, William J.; Plavcan, J. Michael; Cerling, Thure E.; Brown, Francis H. (September 2020). "Late Middle Pleistocene Elephants from Natodomeri, Kenya and the Disappearance of Elephas (Proboscidea, Mammalia) in Africa". Journal of Mammalian Evolution. 27 (3): 483–495. doi:10.1007/s10914-019-09474-9. ISSN 1064-7554. S2CID 198190671.

[:0-2] 1 2 3 Todd, Nancy E. (January 2005). "Reanalysis of African Elephas recki: implications for time, space and taxonomy". Quaternary International. 126–128: 65–72. doi:10.1016/j.quaint.2004.04.015.

[:2-3] 1 2 3 4 5 6 7 8 Sanders, William J. (2023-07-07). Evolution and Fossil Record of African Proboscidea (1 ed.). Boca Raton: CRC Press. pp. 267–293. doi:10.1201/b20016. ISBN 978-1-315-11891-8.

[4] Beden, M. 1980. Elephas recki Dietrich, 1915 (Proboscidea, Elephantidae). Èvolution au cours du Plio-Pléistocène en Afrique orientale. Geobios 13(6): 891-901. Lyon.

[5] Beden, M. 1983. Family Elephantidae. In J. M. Harris (ed.), Koobi Fora Research Project. Vol. 2. The fossil Ungulates: Proboscidea, Perissodactyla, and Suidae: 40-129. Oxford: Clarendon Press.

[6] Beden, M. 1987. Les faunes Plio-Pléistocène de la basse vallée de l’Omo (Éthiopie), Vol. 2: Les Eléphantidés (Mammalia-Proboscidea) (directed by Y. Coppens and F. C. Howell): 1-162. Cahiers de Paléontologie-Travaux de Paléontologie est-africaine. Centre National de la Recherche Scientifique (CNRS). Paris.

[:4-7] 1 2 Larramendi, Asier; Zhang, Hanwen; Palombo, Maria Rita; Ferretti, Marco P. (February 2020). "The evolution of Palaeoloxodon skull structure: Disentangling phylogenetic, sexually dimorphic, ontogenetic, and allometric morphological signals". Quaternary Science Reviews. 229: 106090. Bibcode:2020QSRv..22906090L. doi:10.1016/j.quascirev.2019.106090. S2CID 213676377.

[8] Stimpson, Christopher M.; Lister, Adrian; Parton, Ash; Clark-Balzan, Laine; Breeze, Paul S.; Drake, Nick A.; Groucutt, Huw S.; Jennings, Richard; Scerri, Eleanor M.L.; White, Tom S.; Zahir, Muhammad; Duval, Mathieu; Grün, Rainer; Al-Omari, Abdulaziz; Al Murayyi, Khalid Sultan M. (July 2016). "Middle Pleistocene vertebrate fossils from the Nefud Desert, Saudi Arabia: Implications for biogeography and palaeoecology". Quaternary Science Reviews. 143: 13–36. doi:10.1016/j.quascirev.2016.05.016. hdl: 10072/142575 .

[9] Pokines, James T.; Lister, Adrian M.; Ames, Christopher J. H.; Nowell, April; Cordova, Carlos E. (March 2019). "Faunal remains from recent excavations at Shishan Marsh 1 (SM1), a Late Lower Paleolithic open-air site in the Azraq Basin, Jordan". Quaternary Research. 91 (2): 768–791. doi:10.1017/qua.2018.113. ISSN 0033-5894.

[proboscideans-10] Larramendi, A. (2016). "Shoulder height, body mass and shape of proboscideans" (PDF). Acta Palaeontologica Polonica. 61. doi: 10.4202/app.00136.2014 .

[11] Larramendi, Asier (2023-12-10). "Estimating tusk masses in proboscideans: a comprehensive analysis and predictive model". Historical Biology: 1–14. doi:10.1080/08912963.2023.2286272. ISSN 0891-2963.

[:6-12] Lister, Adrian M. (2013-06-26). "The role of behaviour in adaptive morphological evolution of African proboscideans". Nature. 500 (7462): 331–334. Bibcode:2013Natur.500..331L. doi:10.1038/nature12275. ISSN 0028-0836. PMID 23803767. S2CID 883007.

[13] Lister, Adrian M. (2004), "Ecological Interactions of Elephantids in Pleistocene Eurasia", Human Paleoecology in the Levantine Corridor, Oxbow Books, pp. 53–60, ISBN 978-1-78570-965-4 , retrieved 2020-04-14

[14] Haynes, Gary (March 2022). "Late Quaternary Proboscidean Sites in Africa and Eurasia with Possible or Probable Evidence for Hominin Involvement". Quaternary. 5 (1): 18. doi: 10.3390/quat5010018 . ISSN 2571-550X.

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