ACKNOWLEDGMENT: Many thanks to @mionectes, who brings valuable commentary on genetics to this Journal.
INTRODUCTION
Early in the Holocene, many genera of large-bodied ‘Pleistocene megafauna’ became extinct in the Americas.
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This constitutes one of the most important episodes in the evolutionary history and biogeography of mammals. Furthermore, it remains to be fully explained.
By comparison, there were remarkably few megafaunal extinctions at similar latitudes in Eurasia.
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These contemporaneous extinctions, in the early Holocene, consisted – at the generic level – only of
- Megaloceros, and
- Mammuthus (a proboscidean genus shared with North America).
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An additional possible genus is Coelodonta, which may perhaps have persisted slightly into the Holocene.
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The nearly categorical difference between the western and eastern hemispheres can be explained partly by the historical biogeography of Homo.
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In Eurasia, large-bodied mammals co-evolved with Homo, whereas in the Americas predation by Homo came into effect only at the end of the Pleistocene.
Furthermore, several Eurasian clades of mammalian megafauna (e.g. Equus, Camelus, Bos, Bubalus) were partly domesticated, which meant extinction of ancestral spp. by anthropogenic modification, as opposed to the outright exterminations seen in the Americas.
A recent paper by Liang et al. (2026, ) has raised the possibility of an additional species – and possibly genus – of ‘Pleistocene megafauna’ that became extinct in Eurasia during the Holocene.
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The relevant species is Bubalus mephistopheles.
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The palaeontological significance of B. mephistopheles is limited by the continued existence of the genus Bubalus, which is represented today by at least two spp..
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However, B. mephistopheles seems to belong to a clade distinct from Bubalus kerabau, and possibly also distinct from Bubalus bubalis.
Instead, B. mephistopheles may be most closely related to subgenus Anoa, represented in the extant fauna by two spp. in Sulawesi.
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Anoa possibly warrants its own genus (Groves and Grubb 2011, and ).
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If so,
- our new species should be called Anoa mephistopheles, and
- the genus survives not in its original Eurasia, but instead in Wallacea, across the permanent sea-barrier called the Wallace Line.
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In turn, this would mean that A. mephistopheles represents a new genus of extinct ‘Pleistocene megafauna’ for Eurasia.
This is as opposed to the current assumption, which is that Anoa evolved as a result of
- Bubalus swimming across the Wallace Line, about 1.5 million years ago, and
- subsequent differentiation of a new, insular, genus by virtue of geographical isolation.
The result would be a revision of our list of megafaunal genera that became extinct during the Holocene in Eurasia, to:
- Megaloceros,
- Anoa (remaining extant in Indonesia, in diminutive form), and
- Mammuthus (shared with North America, ).
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Taking my interpretation to its furthest, tentative conclusion:
Anoa may be the only Eurasian genus to emulate somewhat the anthropogenic extinctions of the Americas.
Mammuthus, Megaloceros, and Coelodonta were ‘bizarre ice age giants’ in the conception of Valerius Geist. Thus, their extinction in Eurasia may have been owing more to climatic warming than to predation by humans.
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By contrast, A. mephistopheles was hardly aberrant among bovins.
It was cold-adapted (with long, dense pelage year-round?) only relative to extant ‘water buffaloes’ – which are
- tropical to subtropical, and
- notoriously prone to overheating in warm weather, unless able to wet their bodies.
It is likely that predation by humans during the Holocene caused the extinction of several spp. of Equus.
However, if extinction of A. mephistopheles was similarly caused, the difference is that a whole genus disappeared recently from Eurasia.
Posted on June 22, 2026 01:38 AM by
milewski
The author added the following in comments beneath the original post.
Chatgpt
MY QUESTION
Bubalus wangsjoki and Bubalus mephistopheles are conspecific. Which name takes taxonomic priority/precedence? Supply full references.
ANSWER FROM CHATGPT
If Bubalus wangsjoki (more commonly spelled Bubalus wansijocki or Bubalus wansjocki in the literature) and Bubalus mephistopheles are regarded as the same biological species, then the valid name is determined by the Principle of Priority of the International Commission on Zoological Nomenclature.
Taxonomic priority
Name Author Year Priority
Bubalus mephistopheles Hopwood 1925 Senior synonym
Bubalus wansijocki (often misspelled wangsjoki) Teilhard de Chardin 1928 Junior synonym
Therefore:
If the two taxa are conspecific, Bubalus mephistopheles Hopwood, 1925 has nomenclatural priority and is the correct name. Bubalus wansijocki Chardin, 1928 becomes its junior subjective synonym.
Original descriptions
Hopwood, A. T. (1925).
“A new species of buffalo from the Pleistocene of China.”
Annals and Magazine of Natural History, Series 9, 15: 277–281.
This paper established the name Bubalus mephistopheles.
Teilhard de Chardin, P. (1928).
Description of Bubalus wansijocki from the Chinese Pleistocene. The name was introduced three years after Hopwood’s publication.
Recent evidence
Recent ancient-DNA work has indicated that the traditional Chinese fossil buffalo species are less distinct than previously supposed and that B. mephistopheles and B. wansijocki belong to the same evolutionary lineage. Under such a taxonomic interpretation, the earlier name necessarily has precedence under the ICZN rules.
References
Hopwood, A. T. 1925. A new species of buffalo from the Pleistocene of China. Annals and Magazine of Natural History, ser. 9, 15: 277–281.
Teilhard de Chardin, P. 1928. Original description of Bubalus wansijocki from the Chinese Pleistocene.
International Commission on Zoological Nomenclature. 1999. International Code of Zoological Nomenclature, 4th ed. Article 23 (Principle of Priority).
So, assuming the synonymy is accepted, the correct taxonomic name is Bubalus mephistopheles Hopwood, 1925, with Bubalus wansijocki Teilhard de Chardin, 1928 as its junior synonym.
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Chatgpt
MY QUESTION
List all spp. of mammals with mean adult female body mass >100 kg that went extinct in Eurasia at the end of the Pleistocene or early in the Holocene. Supply full references.
ANSWER FROM CHATGPT
If the criterion is:
mammals
mean adult female body mass >100 kg
native to Eurasia
species that became globally extinct during the Late Pleistocene or Early Holocene (roughly 50–5 ka)
then the list is surprisingly short and reasonably well established.
Species Approximate mean adult female mass Approximate extinction
Palaeoloxodon antiquus 3–6 t Late Pleistocene (~40–30 ka in most of range)
Mammuthus primigenius 2.5–4 t mainland ~11.7 ka; Wrangel Island ~4 ka
Coelodonta antiquitatis 1.8–2.2 t ~14–13.8 ka
Stephanorhinus kirchbergensis 1.5–2.5 t Late Pleistocene
Stephanorhinus hemitoechus 1.2–1.8 t Late Pleistocene
Elasmotherium sibiricum 3.5–4 t ~39–35 ka
Hippopotamus antiquus 1.5–3 t Late Pleistocene
Megaloceros giganteus 350–450 kg Early Holocene (~7.7 ka in W. Siberia)
Ursus spelaeus 180–250 kg females ~30–27 ka
Panthera spelaea 130–180 kg females ~14 ka
Homotherium latidens 110–170 kg females Late Pleistocene
Species excluded
Several large Eurasian mammals do not qualify because they survived into historical times or are still extant:
Bos primigenius – survived until AD 1627.
Equus ferus ferus – survived into the eighteenth–nineteenth centuries.
Bison priscus – generally regarded as ancestral to living bison rather than an end-Pleistocene Eurasian extinction, with survival into the Holocene in Siberia.
Ovibos moschatus – extirpated from Eurasia but survives in North America.
Borderline cases
The following satisfy the body-mass criterion but are more uncertain because of taxonomy or chronology:
Palaeoloxodon namadicus (>8 t) probably persisted into the Late Pleistocene in South Asia, but the precise extinction date remains poorly constrained.
Stegodon orientalis exceeded 2 t but most accepted last occurrences pre-date the terminal Pleistocene, and its final chronology is debated.
Principal references
Stuart, A. J. (1991). Mammalian extinctions in the Late Pleistocene of northern Eurasia and North America. Biological Reviews, 66, 453–562.
Stuart, A. J., & Lister, A. M. (2012). Extinction chronology of the woolly rhinoceros Coelodonta antiquitatis in the context of late Quaternary megafaunal extinctions in northern Eurasia. Quaternary Science Reviews, 51, 1–17.
Stuart, A. J., Kosintsev, P. A., Higham, T. F. G., & Lister, A. M. (2004). Pleistocene to Holocene extinction dynamics in giant deer and woolly mammoth. Nature, 431, 684–689.
Lemoine, M. et al. (2024). The late-Quaternary megafauna extinctions: Patterns, causes, ecological consequences and implications for ecosystem management in the Anthropocene. Cambridge Prisms: Extinction.
This list is intentionally conservative and includes only species for which the available evidence indicates (1) female body mass comfortably exceeds 100 kg and (2) global extinction occurred during the Late Pleistocene or early Holocene in Eurasia.
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STEGODON
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Chatgpt
MY QUESTION
How many thousands of years ago did the last species of Stegodon become fully extinct? Supply full references.
ANSWER FROM CHATGPT
On present evidence, the last known species (or subspecies) of Stegodon became fully extinct approximately 50,000 years ago, i.e. about 50 thousand years ago (50 ka).
There has been some historical debate because an influential 2008 paper argued that the dwarf Flores form Stegodon florensis insularis survived until about 12 ka (12,000 years BP), but subsequent excavations and improved chronologies have shown that the youngest securely dated Stegodon remains on Flores are around 50 ka, with no convincing Holocene survival.
Chronology
Taxon Last accepted occurrence Approximate age
Stegodon florensis insularis (Flores) Late Pleistocene ~50 ka
Earlier interpretation (now generally superseded) Volcanic extinction ~12 ka
Why the 12 ka date changed
The classic study by van den Bergh and colleagues described Stegodon florensis insularis from Liang Bua and concluded:
“Both became extinct during a volcanic eruption around 12 ka BP.”
This referred to both the stegodon and the diminutive hominin Homo floresiensis.
However, subsequent re-evaluation of the Liang Bua stratigraphy and dating demonstrated that the fossil-bearing deposits are substantially older than originally thought. The last appearance of Stegodon is now placed at approximately 50,000 years ago, coinciding with major environmental change and the arrival of modern humans on Flores.
Full references
van den Bergh, G. D., Kaifu, Y., Kurniawan, I., Kono, R. T., Brumm, A., Setiyabudi, E., Aziz, F., & Morwood, M. J. (2008). The youngest stegodon remains in Southeast Asia from the Late Pleistocene archaeological site Liang Bua, Flores, Indonesia. Quaternary International, 182(1), 16–48.
(Original proposal that Stegodon florensis insularis survived until ~12 ka.)
Saegusa, H. (2020). Stegodontidae and Anancus: Keys to understanding dental evolution in Elephantidae. Quaternary Science Reviews, 246, 106527.
(Modern review of stegodont evolution and extinction.)
Stegodon. Current synthesis of the fossil record, summarizing that “the last records of Stegodon florensis date to around 50,000 years ago.”
Therefore, the best current answer is:
The last species of Stegodon became fully extinct approximately 50 thousand years ago. The earlier estimate of 12 thousand years ago is of historical interest but is no longer the generally accepted chronology.
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Chatgpt
MY QUESTION
Consider all megafaunal extinctions in the American supercontinent during the early Holocene. How many genera became extinct in this episode? Supply full references.
ANSWER FROM CHATGPT
The answer depends on how one defines megafauna and the extinction episode.
If one follows the conventional palaeontological definition of the continental American Late Pleistocene–early Holocene megafaunal extinction (generally mammals with adult body mass >44 kg, excluding island endemics and excluding taxa that merely became locally extirpated), the best-supported estimate is:
Approximately 83 genera became globally extinct across the American supercontinent.
This figure is obtained by summing the accepted continental losses:
Region Extinct megafaunal genera
North America 35 genera
South America 48 genera
Total 83 genera
The numbers above are those summarized by standard reviews of the Late Quaternary extinction.
North America: approximately 35 genera disappeared between about 16,000 and 11,500 years BP, with most losses concentrated near the Pleistocene–Holocene transition.
South America: approximately 48 genera disappeared during the last 100,000 years, with the majority lost during the terminal Pleistocene and earliest Holocene extinction pulse. Four genera are known to have survived into the early Holocene before finally becoming extinct.
Because no megafaunal genus is shared between the North and South American extinct lists, the continental total is simply:
35+48= 83 extinct genera
Principal references
Barnosky, A. D., Koch, P. L., Feranec, R. S., Wing, S. L. & Shabel, A. B. (2004). Assessing the causes of Late Pleistocene extinctions on the continents. Science 306: 70–75.
Stuart, A. J. (2015). Late Quaternary megafaunal extinctions on the continents: a short review. Geological Journal 50: 338–363. DOI: 10.1002/gj.2633.
Elias, S. A. & Schreve, D. C. (2019). Late Pleistocene Megafaunal Extinctions. In: Encyclopedia of Quaternary Science (2nd ed.). Elsevier. This chapter explicitly summarizes the losses as 35 extinct genera in North America and 48 in South America, for a total of 83 extinct genera.
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Chatgpt
MY QUESTION
When did the last species of Mammuthus become extinct in North Americas? Supply full references.
ANSWER FROM CHATGPT
The last species of the genus Mammuthus to become extinct in North America was the woolly mammoth, Mammuthus primigenius. The final surviving North American population lived on St. Paul Island, Alaska (Pribilof Islands) and disappeared approximately 5,600 ± 100 years before present (BP), equivalent to roughly 3600 BCE.
This extinction occurred several centuries before the final global extinction of the species on Wrangel Island, Russia, where mammoths survived until about 4,000 years BP (ca. 2000 BCE).
Chronology of the last North American mammoths
Taxon Last known occurrence Approximate calendar age
Mammuthus exilis Santa Rosa Island, California ca. 9,100–9,000 BCE
Mammuthus columbi Continental North America ca. 8,100–7,700 BCE
Mammuthus primigenius St. Paul Island, Alaska 5,600 ± 100 BP (ca. 3600 BCE)
Evidence
The most comprehensive multiproxy study of the St. Paul population concluded:
“The timing of extinction is tightly constrained to 5,600 ± 100 y ago.”
The authors argued that extinction was driven primarily by loss of freshwater habitat as postglacial sea-level rise progressively reduced the island and concentrated mammoths around dwindling water sources, rather than by human hunting or simple food limitation.
Earlier ancient-DNA work on a mammoth from Qagnax̂ Cave had already shown that remains dated to approximately 5725 radiocarbon years BP represented the youngest mammoth remains then known from North America, confirming the existence of a relict Holocene population.
Full references
Graham, R. W., Belmecheri, S., Choy, K., Culleton, B. J., Davies, L. J., Froese, D., Heintzman, P. D., Hritz, C., Kapp, J. D., Newsom, L. A., Rawcliffe, R., Saulnier-Talbot, É., Shapiro, B., Wang, Y., Williams, J. W., & Wooller, M. J. (2016). Timing and causes of mid-Holocene mammoth extinction on St. Paul Island, Alaska. Proceedings of the National Academy of Sciences, 113(33), 9310–9314..
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Debruyne, R., Chu, G., King, C. E., Bos, K., Kuch, M., Schwarz, C., Szpak, P., Gröcke, D. R., Matheus, P., Zazula, G., Guthrie, D., Froese, D., Buigues, B., de Marliave, C., Flemming, C., Poinar, D., & Poinar, H. N. (2009). Phylogeographic analysis of the mid-Holocene mammoth from Qagnax̂ Cave, St. Paul Island, Alaska. Palaeogeography, Palaeoclimatology, Palaeoecology, 273(1–2), 184–190..
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Vartanyan, S. L., Arslanov, K. A., Tertychnaya, T. V., & Chernov, S. B. (2008). Collection of radiocarbon dates on the mammoths (Mammuthus primigenius) and other genera of Wrangel Island, northeast Siberia, Russia. Quaternary International, 182(1), 185–205..
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Chatgpt
MY QUESTION
How long ago did Bubalus cross the Wallace Line to reach Sulawesi for the first time? Supply full references.
ANSWER FROM CHATGPT
The best current evidence suggests that the ancestor of the Sulawesi buffaloes (anoa) first crossed the Wallace Line during the Early Pleistocene, approximately 1–2 million years ago (Ma). The most likely estimate is about 1.4 Ma, although the exact date remains uncertain.
Molecular evidence
The most recent dedicated biogeographic analysis is:
Priyono, D.S., Solihin, D.D., Farajallah, A., & Purwantara, B. (2024). Biogeographic history of the endangered dwarf buffalo, subgenus Anoa (Bovidae: Bubalus quarlesi and Bubalus depressicornis): a perspective based on mitochondrial DNA phylogeny. Evolutionary Systematics, 8(2), 311–321.
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Their principal findings are:
the two living anoa species diverged 1.42 Ma (95% HPD 1.14–1.71 Ma);
the ancestral area of the Indonesian Bubalus lineage is reconstructed as Sundaland;
colonization of Sulawesi therefore necessarily predates the 1.42 Ma split;
the authors conclude that dispersal into Sulawesi occurred during the Pleistocene, with the ancestor subsequently diversifying into mountain and lowland anoa.
Thus the crossing of the Wallace Line is inferred to have occurred slightly before 1.42 Ma, probably around 1.5–2 Ma.
Genomic evidence
A genome-wide phylogenetic study reached a very similar conclusion:
Low, W.Y., et al. (2021). The genome of the lowland anoa (Bubalus depressicornis) illuminates the origin of river and swamp buffalo. Molecular Phylogenetics and Evolution, 161, 107170.
CitedDOI
This study found that:
anoa,
river buffalo, and
swamp buffalo
underwent a rapid radiation during the Pleistocene, with separation occurring at approximately 0.84 Ma.
This date is for the divergence among the major Bubalus lineages, not necessarily the initial arrival in Sulawesi, but it is entirely consistent with a Pleistocene colonization.
Fossil evidence
The fossil record is compatible with an Early Pleistocene arrival but does not yet provide an exact first-occurrence date.
The most comprehensive review is:
Rozzi, R. (2017). A new extinct dwarfed buffalo from Sulawesi and the evolution of the subgenus Anoa: An interdisciplinary perspective. Quaternary Science Reviews, 157, 188–205.
CitedDOI
Rozzi describes the extinct dwarf buffalo Bubalus grovesi and reviews the evolution of Sulawesi buffaloes, concluding that:
all Sulawesi buffaloes derive from a single colonization by a continental Bubalus ancestor;
after arrival, they underwent insular dwarfing and endemic diversification into several extinct and living forms.
Geological context
Sulawesi has never been connected to the Sunda Shelf, even during glacial sea-level minima. Therefore, the ancestral buffalo necessarily crossed a substantial marine barrier—the classic Wallace Line—by overwater dispersal (probably swimming between emergent islands and narrowed straits), rather than by a continuous land bridge.
Conclusion
The present evidence indicates that:
Earliest crossing of the Wallace Line by Bubalus: Early Pleistocene.
Most probable age: approximately 1.5–2.0 million years ago.
Minimum age implied by living anoa divergence: older than 1.42 Ma.
No convincing evidence currently supports a Miocene or Pliocene colonization, and no fossil has yet demonstrated an older presence on Sulawesi.
Full references
Low, W.Y., et al. 2021. The genome of the lowland anoa (Bubalus depressicornis) illuminates the origin of river and swamp buffalo. Molecular Phylogenetics and Evolution 161: 107170..
Priyono, D.S., Solihin, D.D., Farajallah, A., & Purwantara, B. 2024. Biogeographic history of the endangered dwarf buffalo, subgenus Anoa (Bovidae: Bubalus quarlesi and Bubalus depressicornis): a perspective based on mitochondrial DNA phylogeny. Evolutionary Systematics 8(2): 311–321..
Rozzi, R. 2017. A new extinct dwarfed buffalo from Sulawesi and the evolution of the subgenus Anoa: An interdisciplinary perspective. Quaternary Science Reviews 157: 188–205..
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Chatgpt
MY QUESTION
Which spp. of Equus became extinct in Eurasia in the Holocene? Supply full references.
ANSWER FROM CHATGPT
If the question is interpreted strictly as species of the genus Equus that survived into the Holocene in Eurasia but subsequently became globally extinct within the Holocene, the answer is remarkably short.
Species Holocene occurrence Extinction
Equus hydruntinus Europe and southwestern Asia Mid–late Holocene (Neolithic–Iron Age)
Equus ovodovi Siberia, Mongolia and northern China Late Holocene (ca. 3900–3400 years BP)
No other widely accepted Eurasian Equus species are known to have become extinct during the Holocene. The wild horse, Equus ferus, survived into historical times (although many wild populations disappeared locally), and the Asiatic wild asses (E. hemionus complex), kiang (E. kiang), and Przewalski’s horse (E. przewalskii) remain extant.
Equus hydruntinus (European wild ass)
This species occupied open habitats throughout much of Europe and southwestern Asia during the Late Pleistocene. During the Holocene its range became fragmented into isolated refugia in southern Europe and adjacent regions before disappearing entirely.
The most comprehensive analysis of its extinction concludes that:
it survived well into the Holocene;
extinction was temporally staggered, occurring in different refugial populations between the Neolithic and Iron Age;
the evidence does not support survival into the medieval period.
The authors write that E. hydruntinus is “one of Europe’s very few globally extinct Holocene megafaunal mammal species.”
Principal references
Burke, A., Eisenmann, V. & Ambler, G. K. (2003). The systematic position of Equus hydruntinus, an extinct species of Pleistocene equid. Quaternary Research 59: 459–469. DOI: 10.1016/S0033-5894(03)00059-0.
Crees, J. J. & Turvey, S. T. (2014). Holocene extinction dynamics of Equus hydruntinus, a late-surviving European megafaunal mammal. Quaternary Science Reviews 91: 16–29. DOI: 10.1016/j.quascirev.2014.03.003.
Equus ovodovi (Ovodov horse; last Sussemionus)
Originally described from Siberian fossils, this species was later shown by radiocarbon dating and ancient genomics to have survived far longer than previously suspected.
Current evidence indicates:
occurrence from southern western Siberia to northern China;
extinction in Siberia near the end of the Pleistocene;
survival in Mongolia until approximately 3900 years BP;
survival in northern China until approximately 3400 years BP, making it a genuine Holocene extinction.
Genomic analyses also demonstrate a long-term decline in genetic diversity before extinction.
Principal references
Eisenmann, V. & Vasiliev, S. (2011). Unexpected finding of a new Equus species (Mammalia, Perissodactyla) belonging to a supposedly extinct subgenus in Late Pleistocene deposits of Khakassia (Southwestern Siberia). Geodiversitas 33: 519–530.
Cai, D., Zhu, S., Gong, M., Zhang, N., Wen, J., et al. (2022). Radiocarbon and genomic evidence for the survival of Equus (Sussemionus) until the late Holocene. eLife 11: e73346.
Taxonomic note
Some older literature treated Equus hydruntinus as a subspecies of the Asiatic wild ass (Equus hemionus hydruntinus), but modern morphological and palaeontological work generally recognizes it as a distinct extinct species closely related to the hemiones.
Thus, under the current mainstream taxonomy, only two accepted species of Equus became globally extinct in Eurasia during the Holocene: Equus hydruntinus and Equus ovodovi.
TRANSCRIPT FROM AI
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