Birth of Homo erectus - Wood - 2020 - Wiley Online Library
John de Vos, the curator of the collection, was generous with his time and expertise. The hominin fossils were in what looked to be their ...
Birth of Homo erectus - Wood - 2020 - Evolutionary Anthropology: Issues, News, and Reviews - Wiley Online Library

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# Birth of _Homo erectus_

[Bernard Wood](/authored-by/Wood/Bernard),

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Bernard Wood

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- [orcid.org/0000-0003-0273-7332](https://orcid.org/0000-0003-0273-7332)

Center for the Advanced Study of Human Paleobiology, George Washington University, Washington, District of Columbia, USA

**Correspondence**

Bernard Wood, Center for the Advanced Study of Human Paleobiology, George Washington University, Suite 6000, SEH, 800 22nd St NW, Washington, DC 20052.

Email: [\[email protected\]](/cdn-cgi/l/email-protection#0c6e697e626d7e686d7b6363684c6b616d6560226f6361 "Link to email address")

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[Bernard Wood](/authored-by/Wood/Bernard),

Corresponding Author

Bernard Wood

- [\[email protected\]](/cdn-cgi/l/email-protection#f99b9c8b97988b9d988e96969db99e94989095d79a9694 "Link to email address")

- [orcid.org/0000-0003-0273-7332](https://orcid.org/0000-0003-0273-7332)

Center for the Advanced Study of Human Paleobiology, George Washington University, Washington, District of Columbia, USA

**Correspondence**

Bernard Wood, Center for the Advanced Study of Human Paleobiology, George Washington University, Suite 6000, SEH, 800 22nd St NW, Washington, DC 20052.

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First published: 27 November 2020

[https://doi.org/10.1002/evan.21873](https://doi.org/10.1002/evan.21873)

Citations: [6](#citedby-section)

PITHECANTHROPUS ERECTUS, EINE MENSCHENAEHNLICHE UEBERGANGSFORM AUS JAVA. By Eugène Dubois (1894) Batavia: Landesdruckerei. Accessible from the Biodiversity Heritage Library. [https://www.biodiversitylibrary.org/item/132258](https://www.biodiversitylibrary.org/item/132258 "Link to external resource")

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## Abstract

Eugène Dubois was the pioneer of human origins research in South-East Asia, specifically on two of the islands, Sumatra and Java, now included in Indonesia. Dubois was a polymath, whose research interests embraced encephalization and hydrology as well as paleoanthropology. His interpretations of the hominin fossil evidence he collected, which he eventually assigned to _Pithecanthropus erectus_, changed over the years, and he evidently felt defensive about those interpretations, but in his 1894 paper he presents cogent reasons for his decision. The taxon he introduced is still recognized, and while it is no longer seen as “the” link between fossil apes and modern humans, it is currently one of the longest surviving hominin taxa.

In the closing decade of the 19thC Europe was the primary source of fossil evidence for human evolution—the sole exception was Talgai in Australia. Darwin had pointed to Africa as the likely location of the earliest human ancestors, but almost all researchers were convinced Europe was where our own species, _Homo sapiens_, originated. What came before modern humans and the Neanderthals was the stuff of speculation. This changed in 1894 with a publication making the case that three fossils recently excavated at Trinil, on the island of Java, were evidence of _Pithecanthropus erectus_, a creature that provided a link “from the anthropoid apes to Man”.[1](#evan21873-bib-0001) The fossil evidence for what we now know as _Homo erectus_ has been expanded over the years, and ideas about how it relates to modern humans have changed, but Eugène Dubois' paper remains an important milestone in our understanding of human evolution. Eugène Dubois' life and career, and the impact of his research on his contemporaries, have been meticulously researched by a historian of science.[2](#evan21873-bib-0002) This account draws heavily on, and greatly benefits from, Bert Theunissen's meticulous scholarship and assiduous attention to detail. Pat Shipman has also written a biography of Dubois,[3](#evan21873-bib-0003) but by her own admission it is unconventional in that she reconstructs actual, or imagined, conversations, thus making it challenging to tease out fact from informed fiction. Paul Storm worked as Shipman's research assistant on Dubois' biography, and the two of them published a much briefer, and more conventional, assessment of Dubois's contribution to paleoanthropology (Figure [1](#evan21873-fig-0001)).[4](#evan21873-bib-0004)

[![Details are in the caption following the image](/cms/asset/8a138e0e-0fdc-472c-bf10-0321b98bb5d0/evan21873-fig-0001-m.png)](/cms/asset/df953888-45f4-4863-b51b-fecf3e2c4221/evan21873-fig-0001-m.jpg)

**FIGURE 1**

Open in figure viewer [PowerPoint](/action/downloadFigures?id=evan21873-fig-0001&partId=&doi=10.1002%2Fevan.21873)

Plate taken from the 1894 publication of the Trinil hominins. Trinil 1—M3; Trinil 2—Calotte (both found 1891) and Trinil 3—left femur (found in 1892). Note that the irregular bony exostosis on the medial aspect of the proximal end of the shaft is a response to bleeding resulting from trauma involving the adductor muscles

Marie Eugène François Thomas Dubois, the son of an apothecary, was born in 1858 in Eijsden in the province of Limburg in the Netherlands. After attending primary school in his home town, and high school in nearby Roermond, in 1877 Dubois entered the University of Amsterdam to study medicine, qualifying as an M.D. in 1884. Dubois evidently had an aptitude for anatomy, for in 1881, while still a student, he was appointed as an assistant to his anatomy professor Max Fürbringer—who had been trained by Gegenbaur and Haeckel in Jena. He was also invited to teach anatomy at two art schools in Amsterdam, something he continued to do until his departure for what was then referred to as the Dutch East Indies. Dubois became involved in Fürbringer's comparative vertebrate anatomy and embryology research, and he showed that both the fourth and fifth branchial arches—and not just the fourth arch as Fürbringer had suggested—contribute to the development of the thyroid cartilage. In 1886, Dubois was appointed as a lecturer in the department of anatomy in the University of Amsterdam, with the clear expectation that he would eventually succeed Fürbringer as professor.[2](#evan21873-bib-0002) But a little over a year later, Dubois' academic career was upended.

Given the influence of Ernst Haeckel on Dubois' teachers, and the popularity of Haeckel's books, it is inconceivable that Dubois was not influenced by Haeckel's conviction that modern humans are more closely related to the extant apes from South-East Asia (Nusantara in Malay) than they are to the African apes, which was Darwin's position. So, just as Darwin had identified Africa as the place to look for fossil evidence of human ancestors, Haeckel had pushed the case for Nusantara. Dubois failed to persuade the Dutch Colonial Office to support an expedition to the Dutch East Indies to look for fossils, so he joined the Royal Dutch East Indies Army as a Medical Officer Second Class, and on the 28th October, 1887, together with his new wife Anna, he set sail from the Netherlands _en route_ to Sumatra. Despite his exhaustive archival research, Bert Theunissen confesses he is none the wiser about when and why Dubois decided to take the radical step of going halfway around the world in search of the link between the anthropoid apes and modern humans.

Dubois arrived in Padang in Sumatra in the second week of December, and for the first few months he was busy working in the local hospital. In the following May, he asked to be transferred to a convalescent hospital inland, where he had more opportunity for research. It was here that Dubois wrote the article[5](#evan21873-bib-0005) that was intended to convince the Dutch East Indies Government to allow him to focus on paleontological research. His campaign of persuasion was ultimately successful, and in March 1889 he was officially transferred to the Directorate of Education, Religion, and Industry (ERI), and given permission to carry out paleontological research in Sumatra and Java. Dubois' initial research in Sumatra had only turned up the porcupine-gnawed teeth of extant mammals, and the discovery of Holocene modern human crania at Wadjak (now Wajak) in East Java led him to shift the focus of his pursuit of human ancestors, and in April 1890, Dubois was given permission to begin work in Java. Two corporals from the corps of engineers—Gerhadus Kriele and Anthonie de Winter—were assigned to assist him. He was also allocated 50 laborers to undertake any excavations, but the Dutch were harsh colonizers, and it needs to be acknowledged that the labor was forced, not voluntary. In November 1890, Dubois and his team found a more ancient-looking—but frustratingly fragmentary—mandibular fragment at Kedoeng Broebos (now Kedung Brubus) in the Kendeng Hills, but he decided to switch his attention to sediments along the Solo River, so in August 1891 the team began excavating along the banks of the Solo River, near the village of Trinil (Figure [2](#evan21873-fig-0002)).

[![Details are in the caption following the image](/cms/asset/f8e39c96-6915-4e67-9899-35a27815141d/evan21873-fig-0002-m.png)](/cms/asset/85d643ee-4faf-44f3-ac35-f7fd5e47d261/evan21873-fig-0002-m.jpg)

**FIGURE 2**

Open in figure viewer [PowerPoint](/action/downloadFigures?id=evan21873-fig-0002&partId=&doi=10.1002%2Fevan.21873)

Eugène Dubois' excavation at Trinil, Java, in the 1890s. The white “x” marks the spot where the calotte (Trinil 2) was found in 1891 \[Color figure can be viewed at [wileyonlinelibrary.com](http://wileyonlinelibrary.com)\]

In September, 1891, Dubois' field crew unearthed a primate upper third molar. In his third-quarter report for that year Dubois assigned the Trinil 1 tooth to _Anthropopithecus_,[6](#evan21873-bib-0006) which at the time was the genus name used for extant chimpanzees, as well as for the fossil apes Richard Lydekker was recovering in the present-day border area between India and Pakistan. A month later the field crew recovered a large primate calotte, Trinil 2,[7](#evan21873-bib-0007) (Figure [3](#evan21873-fig-0003)) and when they returned to Trinil after the rainy season, in August of the following year they uncovered a complete adult left femur, Trinil 3. Dubois initially stuck with his decision to allocate the material to _Anthropopithecus_, but because he interpreted the femur—which despite an obvious pathology was much more modern human-like than the calotte—as belonging to a creature that stood upright and “used its legs exclusively for locomotion”, he assigned Trinil 1–3 to a novel species, _Anthropopithecus erectus_.[8](#evan21873-bib-0008) But Dubois' views about his discoveries were evolving. Some have suggested that Dubois at one stage believed that the Trinil remains belonged to a fossil gibbon, but that is incorrect. All he suggested was that among the living apes, the Trinil fossils were as close, or even closer, to gibbons than to chimpanzees. But, he also became convinced that they belonged to a taxon that was ancestral to modern humans, which would explain why Dubois abandoned _Anthropopithecus_ in favor of _Pithecanthropus_,[1](#evan21873-bib-0001) a genus name introduced by Ernst Haeckel in his 1868 book _Natürliche Schöpfungsgeschichte_ ( _The History of Creation_).[9](#evan21873-bib-0009) Several of Dubois' teachers—not just Fürbringer—were influenced by Haeckel, who had predicted that a taxon would be found that provided a link between fossil apes and modern humans, and such was Haeckel's confidence in its existence that he proposed a name for the missing taxon, _Pithecanthropus alalus_. In the 1906 English translation of his 1874 _Anthropogenie oder Entwickelungsgeschichte des Menschen_,[10](#evan21873-bib-0010) Haeckel suggested that the penultimate stage in his evolutionary scheme consisted of “men who lacked one of the chief human characteristics, articulate speech and the higher intelligence that goes with it, and so had a less developed brain”, hence the species name _alalus_—meaning “without speech.” The Trinil 2 calotte is now part of an impressive public display in the Naturalis Biodiversity Center in Leiden, but when I visited the Dubois collection in the 1970s it was still in the old Rijksmuseum van Natuurlijke Historie. John de Vos, the curator of the collection, was generous with his time and expertise. The hominin fossils were in what looked to be their original boxes, wrapped in cotton wool. Dubois's papers are with the fossil collection, and it was a privilege to see Dubois' hand-written manuscript of his 1894 paper. Even more impressive was the evidence that Dubois changed his mind as he was writing the manuscript, for in his own hand he had crossed out _Anthropopithecus_ and replaced it with _Pithecanthropus_ (Figure [4](#evan21873-fig-0004)).

[![Details are in the caption following the image](/cms/asset/09ade9d0-6fd8-41d7-9ead-ea0ca7231d99/evan21873-fig-0003-m.png)](/cms/asset/f23cb473-929d-48da-9a93-305590224bfc/evan21873-fig-0003-m.jpg)

**FIGURE 3**

Open in figure viewer [PowerPoint](/action/downloadFigures?id=evan21873-fig-0003&partId=&doi=10.1002%2Fevan.21873)

Dr. John de Vos, curator at the Rijksmuseum van Natuurlijke Historie (now incorporated into the Naturalis Biodiversity Centre), with the original Trinil 2 calotte \[Color figure can be viewed at [wileyonlinelibrary.com](http://wileyonlinelibrary.com)\]

[![Details are in the caption following the image](/cms/asset/7066a9d5-5072-493a-84e7-b9a78fb8f174/evan21873-fig-0004-m.png)](/cms/asset/7b6abf60-87a9-463a-975a-44f145fefc58/evan21873-fig-0004-m.jpg)

**FIGURE 4**

Open in figure viewer [PowerPoint](/action/downloadFigures?id=evan21873-fig-0004&partId=&doi=10.1002%2Fevan.21873)

Detail from Eugène Dubois' handwritten manuscript of his 1894 paper, highlighting the change from “ _Anthropopithecus_” to “ _Pithecanthropus_” \[Color figure can be viewed at [wileyonlinelibrary.com](http://wileyonlinelibrary.com)\]

Early in 1895, Dubois left Java to travel to India, where he had the opportunity to examine Lydekker's fossils in Calcutta, as well as visit the source of the fossils in the Siwalik Hills. In June, 1895, the Dubois family, along with what remained of the fossils collected at Trinil, left Batavia to travel to Amsterdam via Marseilles. Dubois exhibited his discoveries at the Third International Congress of Zoology held in Leiden in September, and he gave talks about _Pithecanthropus erectus_ at the Société Belge de Géologie in Brussels, the Ecole d'Anthropologie in Paris, and then, in November, in Dublin at the Royal Dublin Society. An anonymous report of his Dublin presentation published in _Nature_ under the heading “Dr. Dubois” “Missing Link.”[11](#evan21873-bib-0011) gives a sense of the ground that Dubois covered in these presentations.

Eugène Dubois never returned to Amsterdam to live. For the first 2 years after his return to the Netherlands the family lived in The Hague, and then in 1897—when Dubois was appointed curator of the Palaeontological and Mineralogical Cabinet in the Teylers Museum—Dubois and his family moved to Haarlem. In the same year, Dubois was appointed Professor of Crystallography, Mineralogy, Geology and Paleontology at the University of Amsterdam. Although he kept the hominin fossils with him in Haarlem, the non-hominin fossils he had collected were in the Rijksmuseum van Natuurlijke Historie in Leiden. Dubois spent substantial time and effort juggling his Trinil-related research and his teaching and curatorial obligations, but he was also an expert in water management, and in his archives there were more reprints of papers to do with hydrology than paleontology. This is not evident from most descriptions of Dubois' career.

Dubois never returned to the Dutch East Indies, but his field workers continued to excavate at Trinil until 1900. Two additional isolated teeth, Trinil 4, a left M2, and Trinil 5, a left P3, were found in 1897, and much later in time Dubois' attention was drawn to four other femoral fossils—a substantial part of the shaft and the much-abraded proximal end, designated Trinil 6, and three femoral shaft fragments, Trinil 7–9, that were “found in a collection of fossil bones acquired from excavations executed at Trinil in 1900”—that were also assigned to _P_. _erectus_.[12](#evan21873-bib-0012) A fifth femoral shaft fragment was also allocated to _P_. _erectus_, but while Dubois noted that it “is certainly not from Trinil,” he suggested there is “some probability” that it comes from Kedung Brubus.[13](#evan21873-bib-0013) Prompted by the discovery of OH 28,[14](#evan21873-bib-0014) Michael Day and Theya Molleson[15](#evan21873-bib-0015) examined all of the Trinil femora (i.e., Trinil 3, 7–9, and the fifth femoral shaft fragment) using radiography and microscopy. They concluded that the fifth femoral shaft fragment “is not hominine, hominid or even primate”,[15](#evan21873-bib-0015) p. 151, and expressed doubts about whether the femoral remains from Trinil are the same age as the calotte and the teeth.[15](#evan21873-bib-0015) A decade later, Gail Kennedy[16](#evan21873-bib-0016) addressed the traditional morphometric affinities of the Trinil femora, with a focus on Trinil 3, in a multivariate analysis. Her conclusion was that “although the Trinil femora demonstrate a few characters, particularly in the lower shaft, similar to those in _Homo erectus_, their overall pattern unequivocally allies them with the sapient comparative group.” A more sophisticated recent analysis by Chris Ruff and colleagues[17](#evan21873-bib-0017) came to the same conclusion, suggesting that Trinil 3 “derives from a much more recent period than the calotte”,[17](#evan21873-bib-0017) p. 147. But, Ruff and colleagues also conceded that the Trinil 7–9 femora “may represent _H_. _erectus_”,[17](#evan21873-bib-0017) p. 147.

After re-establishing himself in the Netherland, Dubois continued to write about _P_. _erectus_, and he was evidently planning a more extensive treatment of the fossils, because in 1924 Dubois[18](#evan21873-bib-0018) published 32 photographic images that “are destined for a future memoir”,[18](#evan21873-bib-0018) p. 30. Dubois also took a keen interest in the new discoveries being made in Java by Ralph von Koenigswald, and he was still commenting on—and vigorously contesting—von Koenigswald's interpretations well into his eighth decade.[19](#evan21873-bib-0019), [20](#evan21873-bib-0020) As I have suggested, Dubois' research interests extended beyond paleoanthropology and vertebrate paleontology, and no doubt prompted by the relatively small endocranial volume of Trinil 2, he was one of the first researchers to systematically investigate the relationship between the size of the brain and body size.[21](#evan21873-bib-0021), [22](#evan21873-bib-0022)

As for Trinil, Emil Selenka planned to revive fieldwork and excavations at the site, but he died in 1902 before he could travel to Java, so it was his wife, Margarethe, who led the expedition to Trinil in 1907–8. There was then a long hiatus until a joint Indonesian- Japanese team returned in the 1970s. Neither expedition recovered any hominin fossils. More recently a multi-national research team has once again begun field research at Trinil.[23](#evan21873-bib-0023)

In 1936, Ralph von Koenigswald's exploration of the Sangiran Dome recovered the first of many _P_. _erectus_ remains, but the search for fossils was curtailed in 1940 because of WWII. Since the end of WWII researchers operating in and around the Sangiran Dome have recovered crania, mandibles and some postcranial remains.[24](#evan21873-bib-0024) Many researchers assign the _Homo (Javanthropus) soloensis_ remains from Ngandong, Indonesia, to what we now call _H_. _erectus_. The other major contribution to the hypodigm comes from the _Sinanthropus pekinensis_ fossils from Locality 1 at the Zhoukoudian (formerly Choukoutien) Cave near Beijing. The Locality 1 hominins, which show relatively little morphological variation, resemble, but are not identical to, the _P_. _erectus_ remains recovered from Java. Subsequently, similar-looking evidence has come from sites in Africa and Eurasia.

Until its taxonomy was rationalized, the two main regional subsets of what we now know as _H_. _erectus_ were attributed to three genera: two of them, _Pithecanthropus_ and _Meganthropus_, were from Java, and one, _Sinanthropus_, from China. A fourth genus, _Atlanthropus_, was used for the North African material and a fifth, _Telanthropus_, was introduced to accommodate a _H_. _erectus_-like mandible, SK 15, recovered from Member 2 at Swartkrans in southern Africa. Franz Weidenreich[25](#evan21873-bib-0025) was the first to suggest that the genus _Pithecanthropus_ should be subsumed into _Homo_, and in the same paper he proposed that fossils recovered from what was then called Choukoutien (now called Zhoukoudian), which were initially assigned to _Sinanthropus pekinensis_,[26](#evan21873-bib-0026) should also be transferred to _H_. _erectus_. Subsequently the hypodigms of _Meganthropus_,[27](#evan21873-bib-0027) _Atlanthropus_ [28](#evan21873-bib-0028) and _Telanthropus_ [29](#evan21873-bib-0029) were transferred to _Homo_ as _H_. _erectus_, but a recent analysis of the occlusal morphology of specimens assigned to _Meganthropus palaeojavanicus_, conventionally considered a junior synonym of _H_. _erectus_, suggests that they are more likely hominid and not hominin.[30](#evan21873-bib-0030)

Over the years researchers have proposed changes to the way _H_. _erectus_ is interpreted. Some suggest that what they interpret as morphological continuity between _H_. _erectus_, pre-modern _Homo_ and then _Homo sapiens_ effectively invalidates the specific status of the former. Recent advocates of this course of action include Milford Wolpoff and colleagues,[31](#evan21873-bib-0031) Phillip Tobias,[32](#evan21873-bib-0032) and Darren Curnoe and Tobias.[33](#evan21873-bib-0033) Others have suggested that the hypodigm of _H_. _erectus_ should be expanded to include _Homo habilis_,[34](#evan21873-bib-0034) but the method used to capture skull shape was so lacking in discrimination that it failed to separate Dmanisi 4 from a Neanderthal cranium, nor could it distinguish two crania belonging to _Australopithecus africanus_ (Sts 71 and Sts 5) from crania (KNM-WT 15000 and KNM-ER 3733) belonging to early African _H_. _erectus_ or _H_. _ergaster_.

If the most recently published maximum ages ( _c_.1.4 Ma) for Modjokerto/Perning[35](#evan21873-bib-0035) and Sangiran[36](#evan21873-bib-0036) are confirmed, and an age of _c_.150 ka,[37](#evan21873-bib-0037) or even _c_.120 ka,[38](#evan21873-bib-0038) for the hominins recovered from Ngandong is accepted, then the temporal range of _H_. _erectus_ sensu stricto in Java would be from _c_.1.42 Ma to _c_.120 ka. If _H_. _ergaster_ is included in _H_. _erectus_, then an occipital fragment from Koobi Fora (KNM-ER 2598) would push back the first appearance date to _c_.1.9 Ma, which would be consistent with the age of the _H_. _erectus_-like hominins recovered at Dmanisi.

Although we now think there are many links, and likely more than one chain, in the hominin clade, the evidence Dubois caused to be collected in Java in the 1890s was the first evidence of a long-lived lineage that likely made a significant contribution to the evolution of modern humans. Dubois' detailed 1894 description of _P_. _erectus_ deserves its place in a survey of classic contributions to paleoanthropology.

## ACKNOWLEDGMENTS

I am grateful to John de Vos, for his hospitality in Leiden and for reading and improving this contribution. I am grateful to Josie Joordens for her help, and to two reviewers for their wise suggestions.

## Biography

- **Bernard Wood** is University Professor of Human Origins at George Washington University. He has been interested in paleoanthropology since joining Richard Leakey's first expedition to Lake Turkana, Kenya, in 1968, and is best known for his research on the origins of _Homo_, hominin systematics, phylogeny reconstruction, and comparative morphology.


## REFERENCES

- 1Dubois E. 1894. Pithecanthropus erectus, eine menschenaehnliche Uebergangsform aus Java, Batavia: Landesdruckerei.





[Google Scholar](/action/getFTRLinkout?url=http%3A%2F%2Fscholar.google.com%2Fscholar_lookup%3Fhl%3Den%26publication_year%3D1894%26author%3DE%2BDubois%26title%3DPithecanthropus%2Berectus%252C%2Beine%2Bmenschenaehnliche%2BUebergangsform%2Baus%2BJava&doi=10.1002%2Fevan.21873&linkType=gs&linkLocation=Reference&linkSource=FULL_TEXT)

- 2Theunissen B. 1989. Eugène Dubois and the ape-man from Java. The history of the first ‘missing link’ and its discoverer, Dordrecht: Kluwer. p 1–216.





[Google Scholar](/action/getFTRLinkout?url=http%3A%2F%2Fscholar.google.com%2Fscholar_lookup%3Fhl%3Den%26publication_year%3D1989%26pages%3D1-216%26author%3DB%2BTheunissen%26title%3DEug%25C3%25A8ne%2BDubois%2Band%2Bthe%2Bape%25E2%2580%2590man%2Bfrom%2BJava.%2BThe%2Bhistory%2Bof%2Bthe%2Bfirst%2B%25E2%2580%2598missing%2Blink%25E2%2580%2599%2Band%2Bits%2Bdiscoverer&doi=10.1002%2Fevan.21873&linkType=gs&linkLocation=Reference&linkSource=FULL_TEXT)

- 3Shipman P. 2001. The man who found the missing link. New York: Simon and Schuster. p 1–514.





[Google Scholar](/action/getFTRLinkout?url=http%3A%2F%2Fscholar.google.com%2Fscholar_lookup%3Fhl%3Den%26publication_year%3D2001%26pages%3D1-514%26author%3DP%2BShipman%26title%3DThe%2Bman%2Bwho%2Bfound%2Bthe%2Bmissing%2Blink&doi=10.1002%2Fevan.21873&linkType=gs&linkLocation=Reference&linkSource=FULL_TEXT)

- 4Shipman P, Storm P. 2002. Missing links: Eugène Dubois and the origins of paleoanthropology. _Evol Anthropol_ 11: 108–116.

10.1002/evan.10021



[Web of Science®](/servlet/linkout?suffix=null&dbid=128&doi=10.1002%2Fevan.21873&key=000177049000004&getFTLinkType=true&doiForPubOfPage=10.1002%2Fevan.21873&refDoi=10.1002%2Fevan.10021&linkType=ISI&linkSource=FULL_TEXT&linkLocation=Reference)[Google Scholar](/action/getFTRLinkout?url=http%3A%2F%2Fscholar.google.com%2Fscholar_lookup%3Fhl%3Den%26volume%3D11%26publication_year%3D2002%26pages%3D108-116%26journal%3DEvol%2BAnthropol%26author%3DP%2BShipman%26author%3DP%2BStorm%26title%3DMissing%2Blinks%253A%2BEug%25C3%25A8ne%2BDubois%2Band%2Bthe%2Borigins%2Bof%2Bpaleoanthropology&doi=10.1002%2Fevan.21873&doiOfLink=10.1002%2Fevan.10021&linkType=gs&linkLocation=Reference&linkSource=FULL_TEXT)

- 5Dubois E. 1888. Over de wenschelijkheid van een onderzoek naar de diluviale fauna van Ned. Indië, in het bijzonder van Sumatra. _Naturkundig tijdschrift voor Nederlandsch-indië_ 49: 148–165.





[Google Scholar](/action/getFTRLinkout?url=http%3A%2F%2Fscholar.google.com%2Fscholar_lookup%3Fhl%3Den%26volume%3D49%26publication_year%3D1888%26pages%3D148-165%26journal%3DNaturkundig%2Btijdschrift%2Bvoor%2BNederlandsch%25E2%2580%2590indi%25C3%25AB%26author%3DE%2BDubois%26title%3DOver%2Bde%2Bwenschelijkheid%2Bvan%2Been%2Bonderzoek%2Bnaar%2Bde%2Bdiluviale%2Bfauna%2Bvan%2BNed.%2BIndi%25C3%25AB%252C%2Bin%2Bhet%2Bbijzonder%2Bvan%2BSumatra&doi=10.1002%2Fevan.21873&linkType=gs&linkLocation=Reference&linkSource=FULL_TEXT)

- 6Dubois E. 1892. Palaeontologische onderzoekingen op Java. Verslag van het Mijnwezen. _Extra bijvoegsel de Javasche courant (Batavia) nr_ 3: 12–14.





[Google Scholar](/action/getFTRLinkout?url=http%3A%2F%2Fscholar.google.com%2Fscholar_lookup%3Fhl%3Den%26volume%3D3%26publication_year%3D1892%26pages%3D12-14%26journal%3DExtra%2Bbijvoegsel%2Bde%2BJavasche%2Bcourant%2B%2528Batavia%2529%2Bnr%26author%3DE%2BDubois%26title%3DPalaeontologische%2Bonderzoekingen%2Bop%2BJava.%2BVerslag%2Bvan%2Bhet%2BMijnwezen&doi=10.1002%2Fevan.21873&linkType=gs&linkLocation=Reference&linkSource=FULL_TEXT)

- 7Dubois E. 1892. Palaeontologische onderzoekingen op Java. Verslag van het Mijnwezen. _Extra bijvoegsel de Javasche courant (Batavia) nr._ 36: 12–15.





[Google Scholar](/action/getFTRLinkout?url=http%3A%2F%2Fscholar.google.com%2Fscholar_lookup%3Fhl%3Den%26volume%3D36%26publication_year%3D1892%26pages%3D12-15%26journal%3DExtra%2Bbijvoegsel%2Bde%2BJavasche%2Bcourant%2B%2528Batavia%2529%2Bnr.%26author%3DE%2BDubois%26title%3DPalaeontologische%2Bonderzoekingen%2Bop%2BJava.%2BVerslag%2Bvan%2Bhet%2BMijnwezen&doi=10.1002%2Fevan.21873&linkType=gs&linkLocation=Reference&linkSource=FULL_TEXT)

- 8Dubois E. 1893. Palaeontologische onderzoekingen op Java. _Verslag Mijnwezen (Batavia) nr_ 10: 10–14.





[Google Scholar](/action/getFTRLinkout?url=http%3A%2F%2Fscholar.google.com%2Fscholar_lookup%3Fhl%3Den%26volume%3D10%26publication_year%3D1893%26pages%3D10-14%26journal%3DVerslag%2BMijnwezen%2B%2528Batavia%2529%2Bnr%26author%3DE%2BDubois%26title%3DPalaeontologische%2Bonderzoekingen%2Bop%2BJava&doi=10.1002%2Fevan.21873&linkType=gs&linkLocation=Reference&linkSource=FULL_TEXT)

- 9Haeckel E. 1868. Natürliche Schöpfungsgeschichte: gemeinverständliche wissenschaftliche Vorträge über die Entwickelungslehre im Allgemeinen und Diejenige von Darwin, Goethe und Lamarck im Besonderen, Berlin: G. Reimer.





[Google Scholar](/action/getFTRLinkout?url=http%3A%2F%2Fscholar.google.com%2Fscholar_lookup%3Fhl%3Den%26publication_year%3D1868%26author%3DE%2BHaeckel%26title%3DNat%25C3%25BCrliche%2BSch%25C3%25B6pfungsgeschichte%253A%2Bgemeinverst%25C3%25A4ndliche%2Bwissenschaftliche%2BVortr%25C3%25A4ge%2B%25C3%25BCber%2Bdie%2BEntwickelungslehre%2Bim%2BAllgemeinen%2Bund%2BDiejenige%2Bvon%2BDarwin%252C%2BGoethe%2Bund%2BLamarck%2Bim%2BBesonderen&doi=10.1002%2Fevan.21873&linkType=gs&linkLocation=Reference&linkSource=FULL_TEXT)

- 10Haeckel E. 1874. Anthropogenie oder Entwickelungsgeschichte des Menschen, Leipzig: W. Engelmann.





[Google Scholar](/action/getFTRLinkout?url=http%3A%2F%2Fscholar.google.com%2Fscholar_lookup%3Fhl%3Den%26publication_year%3D1874%26author%3DE%2BHaeckel%26title%3DAnthropogenie%2Boder%2BEntwickelungsgeschichte%2Bdes%2BMenschen&doi=10.1002%2Fevan.21873&linkType=gs&linkLocation=Reference&linkSource=FULL_TEXT)

- 11Anon. 1895. Dr. Dubois' “Missing Link”. _Nature_ 53: 115–116.

10.1038/053115a0



[Google Scholar](/action/getFTRLinkout?url=http%3A%2F%2Fscholar.google.com%2Fscholar%3Fhl%3Den%26q%3D%2BAnon.%2B1895.%2BDr.%2BDubois%2527%2B%25E2%2580%259CMissing%2BLink%25E2%2580%259D.%2BNature%2B53%253A%2B115%25E2%2580%2593116.&doi=10.1002%2Fevan.21873&doiOfLink=10.1038%2F053115a0&linkType=gs&linkLocation=Reference&linkSource=FULL_TEXT)

- 12Dubois E. 1932. The distinct organization of Pithecanthropus, of which the femur bears evidence, now confirmed from other individuals of the described species. _Proc K Ned Akad Wet_ 35: 716–722.





[Google Scholar](/action/getFTRLinkout?url=http%3A%2F%2Fscholar.google.com%2Fscholar_lookup%3Fhl%3Den%26volume%3D35%26publication_year%3D1932%26pages%3D716-722%26journal%3DProc%2BK%2BNed%2BAkad%2BWet%26author%3DE%2BDubois%26title%3DThe%2Bdistinct%2Borganization%2Bof%2BPithecanthropus%252C%2Bof%2Bwhich%2Bthe%2Bfemur%2Bbears%2Bevidence%252C%2Bnow%2Bconfirmed%2Bfrom%2Bother%2Bindividuals%2Bof%2Bthe%2Bdescribed%2Bspecies&doi=10.1002%2Fevan.21873&linkType=gs&linkLocation=Reference&linkSource=FULL_TEXT)

- 13Dubois E. 1935. The sixth (fifth new) femur of _Pithecanthropus erectus_. _Proc K Ned Akad Wet_ 38(8): 850–852.





[Google Scholar](/action/getFTRLinkout?url=http%3A%2F%2Fscholar.google.com%2Fscholar_lookup%3Fhl%3Den%26volume%3D38%26publication_year%3D1935%26pages%3D850-852%26journal%3DProc%2BK%2BNed%2BAkad%2BWet%26issue%3D8%26author%3DE%2BDubois%26title%3DThe%2Bsixth%2B%2528fifth%2Bnew%2529%2Bfemur%2Bof%2BPithecanthropus%2Berectus&doi=10.1002%2Fevan.21873&linkType=gs&linkLocation=Reference&linkSource=FULL_TEXT)

- 14Day MH. 1971. Post-cranial remains of _Homo erectus_ from Bed IV, Olduvai Gorge, Tanzania. _Nature_ 232: 383–387.

10.1038/232383a0



[CAS](/servlet/linkout?suffix=null&dbid=32&doi=10.1002%2Fevan.21873&key=1%3ASTN%3A280%3ADC%252BD2Mvgt1Skug%253D%253D&getFTLinkType=true&doiForPubOfPage=10.1002%2Fevan.21873&refDoi=10.1038%2F232383a0&linkType=COI&linkSource=FULL_TEXT&linkLocation=Reference)[PubMed](/servlet/linkout?suffix=null&dbid=8&doi=10.1002%2Fevan.21873&key=16063052&getFTLinkType=true&doiForPubOfPage=10.1002%2Fevan.21873&refDoi=10.1038%2F232383a0&linkType=PMID&linkSource=FULL_TEXT&linkLocation=Reference)[Web of Science®](/servlet/linkout?suffix=null&dbid=128&doi=10.1002%2Fevan.21873&key=A1971J976400025&getFTLinkType=true&doiForPubOfPage=10.1002%2Fevan.21873&refDoi=10.1038%2F232383a0&linkType=ISI&linkSource=FULL_TEXT&linkLocation=Reference)[Google Scholar](/action/getFTRLinkout?url=http%3A%2F%2Fscholar.google.com%2Fscholar_lookup%3Fhl%3Den%26volume%3D232%26publication_year%3D1971%26pages%3D383-387%26journal%3DNature%26author%3DMH%2BDay%26title%3DPost%25E2%2580%2590cranial%2Bremains%2Bof%2BHomo%2Berectus%2Bfrom%2BBed%2BIV%252C%2BOlduvai%2BGorge%252C%2BTanzania&doi=10.1002%2Fevan.21873&doiOfLink=10.1038%2F232383a0&linkType=gs&linkLocation=Reference&linkSource=FULL_TEXT)

- 15Day MH, Molleson TI. 1976. The puzzle from JK2—a femur and a tibial fragment (OH 34) from Olduvai Gorge, Tanzania. _J Hum Evol_ 5: 455–465.

10.1016/0047-2484(76)90086-5



[Web of Science®](/servlet/linkout?suffix=null&dbid=128&doi=10.1002%2Fevan.21873&key=A1976CM81900005&getFTLinkType=true&doiForPubOfPage=10.1002%2Fevan.21873&refDoi=10.1016%2F0047-2484%2876%2990086-5&linkType=ISI&linkSource=FULL_TEXT&linkLocation=Reference)[Google Scholar](/action/getFTRLinkout?url=http%3A%2F%2Fscholar.google.com%2Fscholar_lookup%3Fhl%3Den%26volume%3D5%26publication_year%3D1976%26pages%3D455-465%26journal%3DJ%2BHum%2BEvol%26author%3DMH%2BDay%26author%3DTI%2BMolleson%26title%3DThe%2Bpuzzle%2Bfrom%2BJK2%25E2%2580%2594a%2Bfemur%2Band%2Ba%2Btibial%2Bfragment%2B%2528OH%2B34%2529%2Bfrom%2BOlduvai%2BGorge%252C%2BTanzania&doi=10.1002%2Fevan.21873&doiOfLink=10.1016%2F0047-2484%2876%2990086-5&linkType=gs&linkLocation=Reference&linkSource=FULL_TEXT)

- 16Kennedy GE. 1983. Femoral morphology in _Homo erectus_. _J Hum Evol_ 12: 587–616.

10.1016/S0047-2484(83)80001-3



[Web of Science®](/servlet/linkout?suffix=null&dbid=128&doi=10.1002%2Fevan.21873&key=A1983RY35100001&getFTLinkType=true&doiForPubOfPage=10.1002%2Fevan.21873&refDoi=10.1016%2FS0047-2484%2883%2980001-3&linkType=ISI&linkSource=FULL_TEXT&linkLocation=Reference)[Google Scholar](/action/getFTRLinkout?url=http%3A%2F%2Fscholar.google.com%2Fscholar_lookup%3Fhl%3Den%26volume%3D12%26publication_year%3D1983%26pages%3D587-616%26journal%3DJ%2BHum%2BEvol%26author%3DGE%2BKennedy%26title%3DFemoral%2Bmorphology%2Bin%2BHomo%2Berectus&doi=10.1002%2Fevan.21873&doiOfLink=10.1016%2FS0047-2484%2883%2980001-3&linkType=gs&linkLocation=Reference&linkSource=FULL_TEXT)

- 17Ruff CB, Puymerail L, Macchiarelli R, Sipla J, Ciochon R. 2015. Structure and composition of the Trinil femora: Functional and taxonomic implications. _J Hum Evol_ 80: 147–158.

10.1016/j.jhevol.2014.12.004



[PubMed](/servlet/linkout?suffix=null&dbid=8&doi=10.1002%2Fevan.21873&key=25681015&getFTLinkType=true&doiForPubOfPage=10.1002%2Fevan.21873&refDoi=10.1016%2Fj.jhevol.2014.12.004&linkType=PMID&linkSource=FULL_TEXT&linkLocation=Reference)[Web of Science®](/servlet/linkout?suffix=null&dbid=128&doi=10.1002%2Fevan.21873&key=000352750800012&getFTLinkType=true&doiForPubOfPage=10.1002%2Fevan.21873&refDoi=10.1016%2Fj.jhevol.2014.12.004&linkType=ISI&linkSource=FULL_TEXT&linkLocation=Reference)[Google Scholar](/action/getFTRLinkout?url=http%3A%2F%2Fscholar.google.com%2Fscholar_lookup%3Fhl%3Den%26volume%3D80%26publication_year%3D2015%26pages%3D147-158%26journal%3DJ%2BHum%2BEvol%26author%3DCB%2BRuff%26author%3DL%2BPuymerail%26author%3DR%2BMacchiarelli%26author%3DJ%2BSipla%26author%3DR%2BCiochon%26title%3DStructure%2Band%2Bcomposition%2Bof%2Bthe%2BTrinil%2Bfemora%253A%2BFunctional%2Band%2Btaxonomic%2Bimplications&doi=10.1002%2Fevan.21873&doiOfLink=10.1016%2Fj.jhevol.2014.12.004&linkType=gs&linkLocation=Reference&linkSource=FULL_TEXT)

- 18Dubois E. 1924. Figures of the calvarium and endocranial cast, a fragment of the mandible and three teeth of Pithecanthropus erectus. _Proc K Ned Akad Wet_ 27(5–6): 459–464.





[Google Scholar](/action/getFTRLinkout?url=http%3A%2F%2Fscholar.google.com%2Fscholar_lookup%3Fhl%3Den%26volume%3D27%26publication_year%3D1924%26pages%3D459-464%26journal%3DProc%2BK%2BNed%2BAkad%2BWet%26issue%3D5%25E2%2580%25936%26author%3DE%2BDubois%26title%3DFigures%2Bof%2Bthe%2Bcalvarium%2Band%2Bendocranial%2Bcast%252C%2Ba%2Bfragment%2Bof%2Bthe%2Bmandible%2Band%2Bthree%2Bteeth%2Bof%2BPithecanthropus%2Berectus&doi=10.1002%2Fevan.21873&linkType=gs&linkLocation=Reference&linkSource=FULL_TEXT)

- 19Dubois E. 1937. On the fossil human skulls recently discovered in Java and _Pithecanthropus erectus_. _Man_ 37: 1–7.

10.2307/2789129



[Google Scholar](/action/getFTRLinkout?url=http%3A%2F%2Fscholar.google.com%2Fscholar_lookup%3Fhl%3Den%26volume%3D37%26publication_year%3D1937%26pages%3D1-7%26journal%3DMan%26author%3DE%2BDubois%26title%3DOn%2Bthe%2Bfossil%2Bhuman%2Bskulls%2Brecently%2Bdiscovered%2Bin%2BJava%2Band%2BPithecanthropus%2Berectus&doi=10.1002%2Fevan.21873&doiOfLink=10.2307%2F2789129&linkType=gs&linkLocation=Reference&linkSource=FULL_TEXT)

- 20Dubois E. 1938. The mandible recently described and attributed to the _Pithecanthropus_ by G.H.R von Koenigswald, compared with the mandible of _Pithecanthropus erectus_ described in 1924 by Eug. Dubois. _Proc K Ned Akad Wet_ 41(2): 139–147.





[Google Scholar](/action/getFTRLinkout?url=http%3A%2F%2Fscholar.google.com%2Fscholar_lookup%3Fhl%3Den%26volume%3D41%26publication_year%3D1938%26pages%3D139-147%26journal%3DProc%2BK%2BNed%2BAkad%2BWet%26issue%3D2%26author%3DE%2BDubois%26title%3DThe%2Bmandible%2Brecently%2Bdescribed%2Band%2Battributed%2Bto%2Bthe%2BPithecanthropus%2Bby%2BG.H.R%2Bvon%2BKoenigswald%252C%2Bcompared%2Bwith%2Bthe%2Bmandible%2Bof%2BPithecanthropus%2Berectus%2Bdescribed%2Bin%2B1924%2Bby%2BEug.%2BDubois&doi=10.1002%2Fevan.21873&linkType=gs&linkLocation=Reference&linkSource=FULL_TEXT)

- 21Dubois E. 1898. Ueber die Abhängigkeit des Hirngewichtes von der Körpergrösse beim Menschen. _Arch Anthropol_ 25: 423–441.





[Google Scholar](/action/getFTRLinkout?url=http%3A%2F%2Fscholar.google.com%2Fscholar_lookup%3Fhl%3Den%26volume%3D25%26publication_year%3D1898%26pages%3D423-441%26journal%3DArch%2BAnthropol%26author%3DE%2BDubois%26title%3DUeber%2Bdie%2BAbh%25C3%25A4ngigkeit%2Bdes%2BHirngewichtes%2Bvon%2Bder%2BK%25C3%25B6rpergr%25C3%25B6sse%2Bbeim%2BMenschen&doi=10.1002%2Fevan.21873&linkType=gs&linkLocation=Reference&linkSource=FULL_TEXT)

- 22Dubois E. 1914. Die gesetzmäbige Beziehung von Gehirmasse zu Körpergrösse bei den Wirbeltieren. _Z Morphol Anthropol_ 18: 323–350.





[Google Scholar](/action/getFTRLinkout?url=http%3A%2F%2Fscholar.google.com%2Fscholar_lookup%3Fhl%3Den%26volume%3D18%26publication_year%3D1914%26pages%3D323-350%26journal%3DZ%2BMorphol%2BAnthropol%26author%3DE%2BDubois%26title%3DDie%2Bgesetzm%25C3%25A4bige%2BBeziehung%2Bvon%2BGehirmasse%2Bzu%2BK%25C3%25B6rpergr%25C3%25B6sse%2Bbei%2Bden%2BWirbeltieren&doi=10.1002%2Fevan.21873&linkType=gs&linkLocation=Reference&linkSource=FULL_TEXT)

- 23Joordens JCA, d'Errico F, Wesselingh FP, et al. 2015. _Homo erectus_ at Trinil on Java used shells for tool production and engraving. _Nature_ 518: 228–231.

10.1038/nature13962



[CAS](/servlet/linkout?suffix=null&dbid=32&doi=10.1002%2Fevan.21873&key=1%3ACAS%3A528%3ADC%252BC2cXitFamsrrP&getFTLinkType=true&doiForPubOfPage=10.1002%2Fevan.21873&refDoi=10.1038%2Fnature13962&linkType=COI&linkSource=FULL_TEXT&linkLocation=Reference)[PubMed](/servlet/linkout?suffix=null&dbid=8&doi=10.1002%2Fevan.21873&key=25470048&getFTLinkType=true&doiForPubOfPage=10.1002%2Fevan.21873&refDoi=10.1038%2Fnature13962&linkType=PMID&linkSource=FULL_TEXT&linkLocation=Reference)[Web of Science®](/servlet/linkout?suffix=null&dbid=128&doi=10.1002%2Fevan.21873&key=000349190300037&getFTLinkType=true&doiForPubOfPage=10.1002%2Fevan.21873&refDoi=10.1038%2Fnature13962&linkType=ISI&linkSource=FULL_TEXT&linkLocation=Reference)[Google Scholar](/action/getFTRLinkout?url=http%3A%2F%2Fscholar.google.com%2Fscholar_lookup%3Fhl%3Den%26volume%3D518%26publication_year%3D2015%26pages%3D228-231%26journal%3DNature%26author%3DJCA%2BJoordens%26author%3DF%2Bd%2527Errico%26author%3DFP%2BWesselingh%26title%3DHomo%2Berectus%2Bat%2BTrinil%2Bon%2BJava%2Bused%2Bshells%2Bfor%2Btool%2Bproduction%2Band%2Bengraving&doi=10.1002%2Fevan.21873&doiOfLink=10.1038%2Fnature13962&linkType=gs&linkLocation=Reference&linkSource=FULL_TEXT)

- 24Aziz F, Kaifu Y, Baba H. 2013. Introduction. In: F Aziz, H Baba, editors. _Homo erectus_ in Indonesia, Bandung: Centre for Geological Survey. p 1–8.





[Google Scholar](/action/getFTRLinkout?url=http%3A%2F%2Fscholar.google.com%2Fscholar_lookup%3Fhl%3Den%26publication_year%3D2013%26pages%3D1-8%26author%3DF%2BAziz%26author%3DY%2BKaifu%26author%3DH%2BBaba%26title%3Din%2BIndonesia&doi=10.1002%2Fevan.21873&linkType=gs&linkLocation=Reference&linkSource=FULL_TEXT)

- 25Weidenreich F. 1940. Some problems dealing with ancient man. _Am Anthrop_ 42(3): 375–383.

10.1525/aa.1940.42.3.02a00010



[Google Scholar](/action/getFTRLinkout?url=http%3A%2F%2Fscholar.google.com%2Fscholar_lookup%3Fhl%3Den%26volume%3D42%26publication_year%3D1940%26pages%3D375-383%26journal%3DAm%2BAnthrop%26issue%3D3%26author%3DF%2BWeidenreich%26title%3DSome%2Bproblems%2Bdealing%2Bwith%2Bancient%2Bman&doi=10.1002%2Fevan.21873&doiOfLink=10.1525%2Faa.1940.42.3.02a00010&linkType=gs&linkLocation=Reference&linkSource=FULL_TEXT)

- 26Black D. 1927. Further hominid remains of lower quaternary age from Chou Kou Tien deposit. _Nature_ 120: 954.

10.1038/120954b0



[Google Scholar](/action/getFTRLinkout?url=http%3A%2F%2Fscholar.google.com%2Fscholar_lookup%3Fhl%3Den%26volume%3D120%26publication_year%3D1927%26pages%3D954%26journal%3DNature%26author%3DD%2BBlack%26title%3DFurther%2Bhominid%2Bremains%2Bof%2Blower%2Bquaternary%2Bage%2Bfrom%2BChou%2BKou%2BTien%2Bdeposit&doi=10.1002%2Fevan.21873&doiOfLink=10.1038%2F120954b0&linkType=gs&linkLocation=Reference&linkSource=FULL_TEXT)

- 27Mayr E. 1944. On the concepts and terminology of vertical subspecies and species. _Natl Res Council Committee Common Probl Genet Paleontol Systemat Bull_ 2: 11–16.





[Google Scholar](/action/getFTRLinkout?url=http%3A%2F%2Fscholar.google.com%2Fscholar_lookup%3Fhl%3Den%26volume%3D2%26publication_year%3D1944%26pages%3D11-16%26journal%3DNatl%2BRes%2BCouncil%2BCommittee%2BCommon%2BProbl%2BGenet%2BPaleontol%2BSystemat%2BBull%26author%3DE%2BMayr%26title%3DOn%2Bthe%2Bconcepts%2Band%2Bterminology%2Bof%2Bvertical%2Bsubspecies%2Band%2Bspecies&doi=10.1002%2Fevan.21873&linkType=gs&linkLocation=Reference&linkSource=FULL_TEXT)

- 28Le Gros Clark WE. 1964. The fossil evidence for human evolution: An introduction to the study of palaeoanthropology, 2nd ed.Chicago: University of Chicago Press.





[Google Scholar](/action/getFTRLinkout?url=http%3A%2F%2Fscholar.google.com%2Fscholar_lookup%3Fhl%3Den%26publication_year%3D1964%26author%3DWE%2BLe%2BGros%2BClark%26title%3DThe%2Bfossil%2Bevidence%2Bfor%2Bhuman%2Bevolution%253A%2BAn%2Bintroduction%2Bto%2Bthe%2Bstudy%2Bof%2Bpalaeoanthropology&doi=10.1002%2Fevan.21873&linkType=gs&linkLocation=Reference&linkSource=FULL_TEXT)

- 29Howell FC. 1978. Hominidae. In: VJ Maglio, HBS Cooke, editors. Evolution of African mammals, Cambridge: Harvard University Press. p 154–248.

10.4159/harvard.9780674431263.c11



[Google Scholar](/action/getFTRLinkout?url=http%3A%2F%2Fscholar.google.com%2Fscholar_lookup%3Fhl%3Den%26publication_year%3D1978%26pages%3D154-248%26author%3DFC%2BHowell%26title%3DEvolution%2Bof%2BAfrican%2Bmammals&doi=10.1002%2Fevan.21873&doiOfLink=10.4159%2Fharvard.9780674431263.c11&linkType=gs&linkLocation=Reference&linkSource=FULL_TEXT)

- 30Zanolli C, Kullmer O, Kelley J, et al. 2019. Evidence for increased hominid diversity in the early to middle Pleistocene of Indonesia. _Nat Ecol Evol_ 3: 755–764.

10.1038/s41559-019-0860-z



[PubMed](/servlet/linkout?suffix=null&dbid=8&doi=10.1002%2Fevan.21873&key=30962558&getFTLinkType=true&doiForPubOfPage=10.1002%2Fevan.21873&refDoi=10.1038%2Fs41559-019-0860-z&linkType=PMID&linkSource=FULL_TEXT&linkLocation=Reference)[Web of Science®](/servlet/linkout?suffix=null&dbid=128&doi=10.1002%2Fevan.21873&key=000466498300014&getFTLinkType=true&doiForPubOfPage=10.1002%2Fevan.21873&refDoi=10.1038%2Fs41559-019-0860-z&linkType=ISI&linkSource=FULL_TEXT&linkLocation=Reference)[Google Scholar](/action/getFTRLinkout?url=http%3A%2F%2Fscholar.google.com%2Fscholar_lookup%3Fhl%3Den%26volume%3D3%26publication_year%3D2019%26pages%3D755-764%26journal%3DNat%2BEcol%2BEvol%26author%3DC%2BZanolli%26author%3DO%2BKullmer%26author%3DJ%2BKelley%26title%3DEvidence%2Bfor%2Bincreased%2Bhominid%2Bdiversity%2Bin%2Bthe%2Bearly%2Bto%2Bmiddle%2BPleistocene%2Bof%2BIndonesia&doi=10.1002%2Fevan.21873&doiOfLink=10.1038%2Fs41559-019-0860-z&linkType=gs&linkLocation=Reference&linkSource=FULL_TEXT)

- 31Wolpoff MH, Thorne AG, Jelínek J, Zhang Y. 1994. The case for sinking _Homo erectus_. 100 years of _Pithecanthropus_ is enough. _Cour Forsch Senckenberg_ 171: 341–361.





[Google Scholar](/action/getFTRLinkout?url=http%3A%2F%2Fscholar.google.com%2Fscholar_lookup%3Fhl%3Den%26volume%3D171%26publication_year%3D1994%26pages%3D341-361%26journal%3DCour%2BForsch%2BSenckenberg%26author%3DMH%2BWolpoff%26author%3DAG%2BThorne%26author%3DJ%2BJel%25C3%25ADnek%26author%3DY%2BZhang%26title%3DThe%2Bcase%2Bfor%2Bsinking%2BHomo%2Berectus.%2B100%25E2%2580%2589years%2Bof%2BPithecanthropus%2Bis%2Benough&doi=10.1002%2Fevan.21873&linkType=gs&linkLocation=Reference&linkSource=FULL_TEXT)

- 32Tobias PV. 1995. The place of _Homo erectus_ in nature with a critique of the cladistic approach. In: Human evolution in its ecological context. Vol. 1: Palaeo-anthropology: Evolution and ecology of _Homo erectus_, Leiden: _Pithecanthropus_ Centennial Foundation. p 31–41.





[Google Scholar](/action/getFTRLinkout?url=http%3A%2F%2Fscholar.google.com%2Fscholar_lookup%3Fhl%3Den%26publication_year%3D1995%26pages%3D31-41%26author%3DPV%2BTobias%26title%3DHuman%2Bevolution%2Bin%2Bits%2Becological%2Bcontext.%2BVol.%2B1%253A%2BPalaeo%25E2%2580%2590anthropology%253A%2BEvolution%2Band%2Becology%2Bof&doi=10.1002%2Fevan.21873&linkType=gs&linkLocation=Reference&linkSource=FULL_TEXT)

- 33Curnoe D, Tobias PV. 2006. Description, new reconstruction, comparative anatomy, and classification of the Sterkfontein Stw 53 cranium, with discussions about the taxonomy of other southern African early _Homo_ remains. _J Hum Evol_ 50(1): 36–77.

10.1016/j.jhevol.2005.07.008



[PubMed](/servlet/linkout?suffix=null&dbid=8&doi=10.1002%2Fevan.21873&key=16243378&getFTLinkType=true&doiForPubOfPage=10.1002%2Fevan.21873&refDoi=10.1016%2Fj.jhevol.2005.07.008&linkType=PMID&linkSource=FULL_TEXT&linkLocation=Reference)[Web of Science®](/servlet/linkout?suffix=null&dbid=128&doi=10.1002%2Fevan.21873&key=000235053500002&getFTLinkType=true&doiForPubOfPage=10.1002%2Fevan.21873&refDoi=10.1016%2Fj.jhevol.2005.07.008&linkType=ISI&linkSource=FULL_TEXT&linkLocation=Reference)[Google Scholar](/action/getFTRLinkout?url=http%3A%2F%2Fscholar.google.com%2Fscholar_lookup%3Fhl%3Den%26volume%3D50%26publication_year%3D2006%26pages%3D36-77%26journal%3DJ%2BHum%2BEvol%26issue%3D1%26author%3DD%2BCurnoe%26author%3DPV%2BTobias%26title%3DDescription%252C%2Bnew%2Breconstruction%252C%2Bcomparative%2Banatomy%252C%2Band%2Bclassification%2Bof%2Bthe%2BSterkfontein%2BStw%2B53%2Bcranium%252C%2Bwith%2Bdiscussions%2Babout%2Bthe%2Btaxonomy%2Bof%2Bother%2Bsouthern%2BAfrican%2Bearly%2BHomo%2Bremains&doi=10.1002%2Fevan.21873&doiOfLink=10.1016%2Fj.jhevol.2005.07.008&linkType=gs&linkLocation=Reference&linkSource=FULL_TEXT)

- 34Lordkipanidze D, Jashashvili T, Vekua A, et al. 2007. Postcranial evidence from early _Homo_ from Dmanisi, Georgia. _Nature_ 449: 305–310.

10.1038/nature06134



[CAS](/servlet/linkout?suffix=null&dbid=32&doi=10.1002%2Fevan.21873&key=1%3ACAS%3A528%3ADC%252BD2sXhtVKiu7jN&getFTLinkType=true&doiForPubOfPage=10.1002%2Fevan.21873&refDoi=10.1038%2Fnature06134&linkType=COI&linkSource=FULL_TEXT&linkLocation=Reference)[PubMed](/servlet/linkout?suffix=null&dbid=8&doi=10.1002%2Fevan.21873&key=17882214&getFTLinkType=true&doiForPubOfPage=10.1002%2Fevan.21873&refDoi=10.1038%2Fnature06134&linkType=PMID&linkSource=FULL_TEXT&linkLocation=Reference)[Web of Science®](/servlet/linkout?suffix=null&dbid=128&doi=10.1002%2Fevan.21873&key=000249557300038&getFTLinkType=true&doiForPubOfPage=10.1002%2Fevan.21873&refDoi=10.1038%2Fnature06134&linkType=ISI&linkSource=FULL_TEXT&linkLocation=Reference)[Google Scholar](/action/getFTRLinkout?url=http%3A%2F%2Fscholar.google.com%2Fscholar_lookup%3Fhl%3Den%26volume%3D449%26publication_year%3D2007%26pages%3D305-310%26journal%3DNature%26author%3DD%2BLordkipanidze%26author%3DT%2BJashashvili%26author%3DA%2BVekua%26title%3DPostcranial%2Bevidence%2Bfrom%2Bearly%2BHomo%2Bfrom%2BDmanisi%252C%2BGeorgia&doi=10.1002%2Fevan.21873&doiOfLink=10.1038%2Fnature06134&linkType=gs&linkLocation=Reference&linkSource=FULL_TEXT)

- 35Morley RJ, Morley HP, Zaim Y, Huffman OF. 2020. Paleoenvironmental setting of Mojokerto _Homo erectus_, the palynological expressions of Pleistocene marine deltas, open grasslands and volcanic mountains in East Java. _J Biogeography_ 47: 566–583.

10.1111/jbi.13770



[Web of Science®](/servlet/linkout?suffix=null&dbid=128&doi=10.1002%2Fevan.21873&key=000507852400001&getFTLinkType=true&doiForPubOfPage=10.1002%2Fevan.21873&refDoi=10.1111%2Fjbi.13770&linkType=ISI&linkSource=FULL_TEXT&linkLocation=Reference)[Google Scholar](/action/getFTRLinkout?url=http%3A%2F%2Fscholar.google.com%2Fscholar_lookup%3Fhl%3Den%26volume%3D47%26publication_year%3D2020%26pages%3D566-583%26journal%3DJ%2BBiogeography%26author%3DRJ%2BMorley%26author%3DHP%2BMorley%26author%3DY%2BZaim%26author%3DOF%2BHuffman%26title%3DPaleoenvironmental%2Bsetting%2Bof%2BMojokerto%2BHomo%2Berectus%252C%2Bthe%2Bpalynological%2Bexpressions%2Bof%2BPleistocene%2Bmarine%2Bdeltas%252C%2Bopen%2Bgrasslands%2Band%2Bvolcanic%2Bmountains%2Bin%2BEast%2BJava&doi=10.1002%2Fevan.21873&doiOfLink=10.1111%2Fjbi.13770&linkType=gs&linkLocation=Reference&linkSource=FULL_TEXT)

- 36Matsu'ura S, Kondo M, Danhara T, et al. 2020. Age control of the first appearance datum for Javanese _Homo erectus_ in the Sangiran area. _Science_ 367: 210–214.

10.1126/science.aau8556



[PubMed](/servlet/linkout?suffix=null&dbid=8&doi=10.1002%2Fevan.21873&key=31919224&getFTLinkType=true&doiForPubOfPage=10.1002%2Fevan.21873&refDoi=10.1126%2Fscience.aau8556&linkType=PMID&linkSource=FULL_TEXT&linkLocation=Reference)[Web of Science®](/servlet/linkout?suffix=null&dbid=128&doi=10.1002%2Fevan.21873&key=000506811300044&getFTLinkType=true&doiForPubOfPage=10.1002%2Fevan.21873&refDoi=10.1126%2Fscience.aau8556&linkType=ISI&linkSource=FULL_TEXT&linkLocation=Reference)[Google Scholar](/action/getFTRLinkout?url=http%3A%2F%2Fscholar.google.com%2Fscholar_lookup%3Fhl%3Den%26volume%3D367%26publication_year%3D2020%26pages%3D210-214%26journal%3DScience%26author%3DS%2BMatsu%2527ura%26author%3DM%2BKondo%26author%3DT%2BDanhara%26title%3DAge%2Bcontrol%2Bof%2Bthe%2Bfirst%2Bappearance%2Bdatum%2Bfor%2BJavanese%2BHomo%2Berectus%2Bin%2Bthe%2BSangiran%2Barea&doi=10.1002%2Fevan.21873&doiOfLink=10.1126%2Fscience.aau8556&linkType=gs&linkLocation=Reference&linkSource=FULL_TEXT)

- 37Indriati E, Swisher CC, Lepre C, et al. 2011. The age of the 20 meter Solo River terrace, Java, Indonesia and the survival of _Homo erectus_ in Asia. _PLoS One_ 6(6):e21562.

10.1371/journal.pone.0021562



[CAS](/servlet/linkout?suffix=null&dbid=32&doi=10.1002%2Fevan.21873&key=1%3ACAS%3A528%3ADC%252BC3MXosFGgu7k%253D&getFTLinkType=true&doiForPubOfPage=10.1002%2Fevan.21873&refDoi=10.1371%2Fjournal.pone.0021562&linkType=COI&linkSource=FULL_TEXT&linkLocation=Reference)[PubMed](/servlet/linkout?suffix=null&dbid=8&doi=10.1002%2Fevan.21873&key=21738710&getFTLinkType=true&doiForPubOfPage=10.1002%2Fevan.21873&refDoi=10.1371%2Fjournal.pone.0021562&linkType=PMID&linkSource=FULL_TEXT&linkLocation=Reference)[Web of Science®](/servlet/linkout?suffix=null&dbid=128&doi=10.1002%2Fevan.21873&key=000292290100039&getFTLinkType=true&doiForPubOfPage=10.1002%2Fevan.21873&refDoi=10.1371%2Fjournal.pone.0021562&linkType=ISI&linkSource=FULL_TEXT&linkLocation=Reference)[Google Scholar](/action/getFTRLinkout?url=http%3A%2F%2Fscholar.google.com%2Fscholar_lookup%3Fhl%3Den%26volume%3D6%26publication_year%3D2011%26journal%3DPLoS%2BOne%26issue%3D6%26author%3DE%2BIndriati%26author%3DCC%2BSwisher%26author%3DC%2BLepre%26title%3DThe%2Bage%2Bof%2Bthe%2B20%2Bmeter%2BSolo%2BRiver%2Bterrace%252C%2BJava%252C%2BIndonesia%2Band%2Bthe%2Bsurvival%2Bof%2BHomo%2Berectus%2Bin%2BAsia&doi=10.1002%2Fevan.21873&doiOfLink=10.1371%2Fjournal.pone.0021562&linkType=gs&linkLocation=Reference&linkSource=FULL_TEXT)

- 38Rizal Y, Westaway KE, Zaim Y, et al. 2019. Last appearance of _Homo erectus_ at Ngandong, Java, 117,000– 108,000 years ago. _Nature_ 577: 381–385.

10.1038/s41586-019-1863-2



[PubMed](/servlet/linkout?suffix=null&dbid=8&doi=10.1002%2Fevan.21873&key=31853068&getFTLinkType=true&doiForPubOfPage=10.1002%2Fevan.21873&refDoi=10.1038%2Fs41586-019-1863-2&linkType=PMID&linkSource=FULL_TEXT&linkLocation=Reference)[Web of Science®](/servlet/linkout?suffix=null&dbid=128&doi=10.1002%2Fevan.21873&key=000509570100035&getFTLinkType=true&doiForPubOfPage=10.1002%2Fevan.21873&refDoi=10.1038%2Fs41586-019-1863-2&linkType=ISI&linkSource=FULL_TEXT&linkLocation=Reference)[Google Scholar](/action/getFTRLinkout?url=http%3A%2F%2Fscholar.google.com%2Fscholar_lookup%3Fhl%3Den%26volume%3D577%26publication_year%3D2019%26pages%3D381-385%26journal%3DNature%26author%3DY%2BRizal%26author%3DKE%2BWestaway%26author%3DY%2BZaim%26title%3DLast%2Bappearance%2Bof%2BHomo%2Berectus%2Bat%2BNgandong%252C%2BJava%252C%2B117%252C000%25E2%2580%2593%2B108%252C000%25E2%2580%2589years%2Bago&doi=10.1002%2Fevan.21873&doiOfLink=10.1038%2Fs41586-019-1863-2&linkType=gs&linkLocation=Reference&linkSource=FULL_TEXT)


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