Orrorin tugenensis is one of the earliest known hominin species. Recovered fossils date to around six million years ago--so far back in time that they may have lived at the same time as the very earliest known hominin ancestor, Sahelanthropus tchadensis.
"Orrorin tugenensis" means "original man from Tugen", which is the name given to the species by Martin Pickford and Brigitte Senut, the paleoanthropology team leaders who discovered and identified it in 2001. Pickford and Senut chose this name in part because O. tugenesis fossils were discovered in the Tugen Hills, Kenya, in the Rift Valley. The "original man" part of the name has a more complicated explanation.
Where did Orrorin tugenesis live? Its fossils have been found in the Tugen Hills, on the Kenyan side of the Rift Valley.
The Rift Valley is famous as a site with abundant fossil deposits, especially fossils of ancient hominids. Its geology has been well documented for over 100 years. (Hill and Ward, 1988) According to Hill and Ward, the oldest strata of sediment, dated to about 14 million years, has no fossils whatsoever. The Muruyur layer, at 13 million years old, is the oldest to contain fossils. Orrorin tugenesis doesn't appear in the fossil record until the sedimentary Lukeino Formation, which can be dated precisely because it is bracketed by easily-dated volcanic layers.
The Lukeino Formation, which is about 120 meters thick, was thoroughly investigated by the Baringo Paleontological Research Project using K-Ar and Ar-Ar and paleomagnetic stratigraphy absolute dating techniques. The fossilized faunal remains, including O. tugenensis, were found in tuff-, siltstone-, and sandstone-rich uppermost part of the formation. K-Ar dating indicates the formation is 6.06 million years old, give or take 130,000 years. (Dieno et al, 2002) Ar-Ar sampling conducted by Dieno and colleagues dates the upper strata of the site at 5.73 million years old, give or take 50,000 years. Paleomagnetic results corroborate this date range as well. All results support that O. tugenensis lived in the Tugen Hills approximately 6 million years ago.
The Rift Valley saw a lot of volcanic activity before, during, and after O. tugenesis's time. This accumulation is what encased and preserved the faunal remains that survive today, fossilizing them. (Dericquebourg et al, 2019)
Interestingly, faunal remains of a gorilla-sized ape and a chimp-like hominoid were discovered in the same Lukeino Formation at O. tugenesis, spurring speculation that the three hominoids may have co-existed. (Senut, Pickford, et al, 2001) Fossils of monkeys and impalas have also been recovered. It is believed this area's paleoenvironment was woodland and savanna, with many ruminant mammals and monkeys populating a landscape containing shallow lakes. (Pickford and Senut, 2001) The area only became the ecosystem we know today roughly 700 thousand years ago. (Faith, Rowan, and Du, 2019).
Senut and Pickford not only discovered a new species, but an entirely new genus. How this new genus fits into the grand phylogentic scheme is not resolved, but Senut and Pickford list descriptions and comparisons for the species:
"Order Primates Linnæus, 1758
Suborder Anthropoidea Mivart, 1864
Superfamily Hominoidea Gray, 1825
Family Hominidae Gray, 1825
Genus Orrorin Senut et al., 2001
Orrorin tugenensis Senut et al., 2001"
(Senut, Pickford, et al, 2001)
Orrorin tugenesis is a very early hominid with very few fossils surviving to clue us in on what it may have looked like. Nonetheless, paleoanthropologists have been able to determine some important traits that distinguish it from other hominids and hominins.
Orrorin tugenesis's dentition is a great place to observe some of these differences. O. tugenesis had apelike dentition--incisors, canines and premolars--and small molars in comparison to its front teeth. A trait that defies an otherwise ancestral-looking mouth of teeth, is the derived trait of having thick enamel.
Like its ancestors, Orrorin tugenesis likely spent a lot of time in the trees. The recovered phalanx fossils are curved, which would have given it the ability to easily hang from branches. This is especially true when paired with the shape of humerus (upper arm bone) found, which is adapted to climbing.
Though it is contentious, there is indication in the shape of Orrorin tugenesis's femur that it had the ability to walk on two legs at least some of the time. The round/spherical head is rotated at a slight angle that would facilitate 2-legged walking. Another clue is the length of the femoral neck. This is important because there are researchers who believe Orrorin tugenesis was bipedal often enough to be considered the first hominin. Others are not convinced. For example, there are some who strongly argue the cortical bone distribution point to bipedalism. (Davers et all, 2022) There are those who argue compellingly that there is not enough evidence for anything more than infrequent bipedalism at most. (Cazenave, Pina, et al, 2024)
Beyond these traits, there simply is not enough fossil material to make a reasonably educated guess about how Orrorin tugenesis may have looked, behaved, or how it was related to other hominids and hominins.
Was Orrorin tugenensis was a hominid or a hominin? This extensively-argued question hinges entirely on whether or not O. tugenensis was bipedal and therefore considered a hominin. Some scientists do not believe there is enough evidence to conclude O. tugenensis was bipedal. On the other end of the spectrum, there are those who believe O. tugenensis was not only a bipedal hominin, but a direct ancestor to the obligate bipedal genus Homo. The latter hypothesis requires a radical re-configuration of the phylogenetic tree, moving australopithecines to their own branch. The team who discovered and identified Orrorin tugenensis advocate for this model.
Some researchers fall on the side of Orrorin tugenesis being a biped based on femur morphology. When exploring evidence for bipedalism in Miocene apes, G. Davers and colleagues note how some Miocene apes' femurs, including Orrorin tugenesis's, have "... proximal platymeria immediately distal to the lesser trochanter ... a trait observed in O. tugenensis and later hominins and suggests a correlation with neck elongation, an indicator of hominin bipedalism" (Davers et all, 2022). Davers also links the femur's cortical thickness and the presence of calcar femorale, all found in O. tugenensis, with bipedalism.
Marine Cazenave and colleagues have independently tested Davers et al's findings and have disagreed with their conclusion on nearly every point. For example, Cazenave has determined O. tugenesis's femur's cortical thickness and distribution, which the Davers publication associated with bipedalism, is similar to Pongo (chimpanzee) and therefore cannot be used as evidence for habitual bipedalism. (Cazenave, Pina, et al, 2024)
Orrorin tugenensis's status as a hominin and position on the the phylogenetic tree is still debated.
Why does it matter so much if Orrorin tugenesis was a hominin or a hominid? Does it really make a difference to science if it is an ancestor to Homo? There are many disagreements about ancient hominin phylogeny, but the answer to this question has particularly impactful ramifications. If Orrorin tugenesis is Homo's ancestor, the model for how we order and understand hominin lineage is radically reconfigured. In fact, the australopithecines, who we currently believe are Homo's ancestor, would be relegated to a dead-end branch of the phylogenetic tree.
Very few fossil specimens have been found, and those that have been found are fragments. What Orrorin tugenensis may have looked like as a living creature is a mystery due to the lack of cranial and post-cranial fossils.
Why don't researchers use ancient DNA extracted from Orrorin tugenensis's remains to answer questions about Orrorin tugenensis, like they have with more recent extinct hominins? To accomplish that, scientists need viable organic matter like that found in long bones and hard teeth. Unfortunately, all the recovered Orrorin tugenensis remains have long since fossilized. Fossilization is the replacement of organic tissue with minerals over a long period of time. There is simply nothing left to test but rocks in the shape of O. tugenensis bone fragments. This transformation from bone to mineral happened over a very long time as a result of the specific conditions of the Lukeino Formation in which the bones were trapped six million years ago. (Dericquebourg et al, 2019)
Though fossil materials are sparse, researchers have been able to extrapolate some information about Orrorin tugenesis by piecing together the fossils that have been found.
BAR 1000a'00 and BAR 1000b'00 are the Orrorin tugenensis type specimen and are left and right mandibular fragments, respectively. They demonstrate how small Orrorin tugenesis's molars are, and how thick tooth its enamel is.
BAR 1000a'00 has the roots of molar 1, and complete molars 2 and 3. BAR 1000b'00 has complete molar 3, which is not well preserved and doesn't provide any new information BAR 1000a'00 hadn't already provided. (Senut, Pickford, et al, 2001)
Later on, BAR 201’01, a mandibular symphasis bone that only fits awkwardly into place, was added to the holotype. (Senut, Pickford, et al, 2001)
All components of this specimen were found by a team lead by Brigitte Senut and Martin Pickford.
BAR 1002'00 is the infamous bone of contention whose traits, and the implications of those traits, are hotly debated.
The overall size of BAR 1002'00 is the same as an ape's femur. However, the shape of the head and neck have differences that hint at bipedalism--and so do the morphology of the muscle attachments. It is the only femur specimen with a well preserved femoral head. (Galik, Senut, et al, 2004)
This fossil was found by a team lead by Brigitte Senut and Martin Pickford.
The moniker "Millennium Man" was given to this assemblage because Orrorin tugenensis fossils were discovered around the turn of the 21st century--the beginning of a new millennium. "Millennium Man" is a composite of 12 specimens from at least 5 individuals. Together, these fossils help researchers do what is impossible individually: They give a tiny glimpse into what a theoretical Orrorin tugenensis might have been like.
All components of this specimen were found by a team lead by Brigitte Senut and Martin Pickford.
There are only a handful of Orrorin tugenesis fossils, so it has proven difficult to arrive at satisfying conclusions for a host of questions regarding this ancient creature. More fossils could mean more answers--or at least different questions.
Though protected under Kenya's Museums and Antiquities Heritage Act of 2006, Ministry of Culture, Arts and Sports; Tugen Hills' archaeological sites are at great risk of being destroyed by human activities like farming, foraging, and other clandestine actions related to widespread poverty. (Kipkorir and Wandibba, 2015)
Cazenave, Marine, Marta Pina, et al. 2024. "Postcranial evidence does not support habitual bipedalism in Sahelanthropus tchadensis: A reply to Daver et al. (2022)" Journal of Human Evolution: 103557. doi:10.1016/j.jhevol.2024.103557. hal-04632880.
Daver, G., F. Guy, H.T. Mackaye, et al. 2022. "Postcranial evidence of late Miocene hominin bipedalism in Chad." Nature 609: 94–100. https://doi.org/10.1038/s41586-022-04901-z.
Deino, Alan L., Lisa Tauxe, Marc Monaghan, and Andrew Hill. 2002. “40Ar/39Ar Geochronology and Paleomagnetic Stratigraphy of the Lukeino and Lower Chemeron Formations at Tabarin and Kapcheberek, Tugen Hills, Kenya.” Journal of Human Evolution 42 (1–2): 117–40. doi:10.1006/jhev.2001.0521.
Dericquebourg, Perrine, et al. 2019. “Bone Diagenesis and Origin of Calcium Phosphate Nodules from a Hominid Site in the Lukeino Formation (Tugen Hills, Kenya).” Palaeogeography, Palaeoclimatology, Palaeoecology 536 (January). doi:10.1016/j.palaeo.2019.109377.
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Galik, K., B. Senut, M. Pickford, and et al. 2004. “External and Internal Morphology of the BAR 1002’00 Orrorin Tugenensis Femur.” Science 305, no. 5689: 1450–53. http://www.jstor.org/stable/3837821.
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