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Peștera cu Oase · Anina · 40,000 years ago

A Neanderthal in the family. No sign of Oase 1 in later populations

The Oase 2 cranium seen from the front: a modern forehead, strong cheekbones, a massive jaw.
The Oase 2 cranium, the adolescent from the same cave: a modern braincase with archaic traits. The picture is Oase 2; the 6–9% was measured in Oase 1. The Oase 2 cranium · Ryan Somma · CC BY-SA 2.0 · Wikimedia Commons

A man whose remains were found near Anina, about 40,000 years ago, carried 6–9% Neanderthal DNA from an ancestor 4–6 generations back. DNA from the jawbone allowed the publication of the Oase 1 genome in 2015. His genome shows no detectable contribution to later populations.

Descend into the cave

Three bars on one scale: today's indicative benchmark, Oase 1 at the midpoint of the measured range, and a Neanderthal at 100%.

The 40,000-year descent

From Oase 1 to today is about 1,600 generations, using the rounded age of 40,000 years and 25 years per generation. Scroll and watch the present approach.

today

Final value: today.

  1. 40,440 years ago

    Oase 1 dates to 40,440 ± 1,030 cal BP: the jaw's combined age.

  2. 16 February 2002

    Cavers find the jaw on the cave floor. Oase 1's story begins.

  3. 2003

    The jaw is published in PNAS: the oldest certain modern human in Europe.

  4. 2015

    The genome is read: 6–9% Neanderthal DNA, the 4–6-generations-back ancestor.

The discovery

In February 2002, cavers explored the karstic system of the Miniș Valley, near Anina in the south-western Carpathians. Down a chamber unknown until then, they found a profusion of mammal bones: the cave had been mostly a hibernation chamber for the cave bear, Ursus spelaeus. On the old floor, near today's entrance, lay a complete human jaw.

Ștefan Milota, Adrian Bîlgăr and Laurențiu Sarcina found it on 16 February 2002. The chamber was named Peștera cu Oase, the Cave with Bones, and the jaw Oase 1, also nicknamed Ion din Anina. No excavation was needed: the bone lay out in the open, on the surface, in the chamber full of bear bones.

In June 2003 the same cave yielded the remains of a cranium: the anterior cranial skeleton, the left temporal bone and frontal, parietal and occipital fragments, again from the surface. Ștefan Milota, Ricardo Rodrigo and Mircea Gherase found them on 20 June, while mapping the cave. Field campaigns followed in 2003–2005, with systematic recording and digging, and completed the Oase 2 cranium, the cave's second human.

The Oase 1 mandible was found on February 16, 2002 on the paleosurface near the current entrance to the chamber.

Trinkaus et al., PNAS 100(20), 2003
A complete cave-bear skeleton in a museum case, with its massive skull and long fangs.
The cave bear, Ursus spelaeus, the chamber's master before humans: the cave was its hibernation den. A cave-bear skeleton, Ursus spelaeus · Wikipek · CC BY 3.0 · Wikimedia Commons

The bones: the jaw and the cranium

Two people, not one. The jaw is an adult's, the cranium an adolescent's.

  1. Oase 1: the jaw

    A robust lower jaw from a grown man, modern with a few archaic traits and one Neanderthal trait. Two laboratories dated it directly by radiocarbon: over 35,200 14C years (Oxford) and 34,290, +970, −870 14C years (Groningen), a combined 34,950, +990, −890 14C years, calibrated to 40,440 ± 1,030 years. At publication in 2003, it was the oldest certain modern human in Europe.

  2. Oase 2: the cranium

    An adolescent cranium, found in 2003, with the large round braincase of a modern human, but a large face, a thick crest behind the ear and large molars growing larger toward the back of the mouth: a mosaic of modern and archaic. Direct dating of the cranium gave only a minimum age (28,890, +∞, −170 14C years); the minimum age and the morphological similarities to Oase 1, discussed by Rougier et al., point to about 40,500 calendar years (about 35,000 14C years).

  3. Oase 3: the bone that came home

    A left temporal bone was first described as the cave's third human (Trinkaus et al. 2003). Its later articulation along the squamous suture showed it belongs to the same cranium as the Oase 2 bones (Rougier et al. 2007). The total stays two people.

The rounded age in the title, 40,000, is 40,440 rounded to thousands.

The Oase 2 cranium seen from the front: a modern forehead, strong cheekbones, a massive jaw.
The Oase 2 cranium: a modern braincase, a face and teeth with archaic traits. The Oase 2 cranium · Ryan Somma · CC BY-SA 2.0 · Wikimedia Commons
The reconstructed bust of the Oase 2 adolescent: skin, hair, and eyes meeting the viewer.
The Oase 2 facial reconstruction, shown at the Neanderthal Museum in Mettmann, Germany. The Oase 2 facial reconstruction, Neanderthal Museum, Mettmann · Daniela Hitzemann (photograph) · CC BY-SA 4.0 · Wikimedia Commons

The genome: how Neanderthal he was

In 2015, 13 years after the discovery, Qiaomei Fu's team read the Oase 1 genome. He was male: mitochondrial DNA from a basal branch of haplogroup N, a Y chromosome from haplogroup F, later refined to K2a*, as in the Ust'-Ishim man. The headline stands in three bars:

The Neanderthal share of the genome

Oase 1 is drawn at the midpoint of the measured range, 7.5%: the bar shows 7.5 out of 100.

Against the indicative benchmark of ≈2%, the midpoint of the range (7.5%) is 3.75 times as large.

Three long, unbroken chunks

Three Neanderthal stretches in his genome each run over 50 centimorgans. Long stretches mean recent mixing: recombination would have chopped them small over many generations. Their length dates the Neanderthal ancestor to 4–6 generations before him, about 100–180 years earlier if a generation is 25–30 years.

An enlarged scheme: the three segments, each over 50 centimorgans, in the Oase 1 genome.

The generations, reckoned

The table shows the expected contribution from a single ancestor, as intuition: 1/2 to the power n, where n is the number of generations. Real inheritance varies through recombination, and total Neanderthal DNA can come from more than one ancestor. So the percentage alone does not prove the generation count: the published 4–6-generation inference rests on the length of the three segments over 50 centimorgans.

Expected contribution from a single ancestor (intuition, not proof)
·1/2n%
parent1/250%
grandparent1/425%
great-grandparent1/812.5%
great-great-grandparent1/166.25%
5 generations1/323.12%
6 generations1/641.56%

The table is only intuition: real inheritance varies, and the evidence for 4–6 generations is the segments' length.

A reconstruction of a Neanderthal man: a low forehead, a heavy brow, long hair.
The Neanderthal, the relative that left Oase 1 some 6–9% of his genome. A Neanderthal reconstruction · Luna04 · CC BY 2.5 · Wikimedia Commons
Two skulls side by side: the Neanderthal one, with its thick brow, and the modern human one.
A Neanderthal skull beside a modern human skull: close relatives, distinct species. Skulls compared: Neanderthal and modern human · Jacklee · CC BY-SA 4.0 · Wikimedia Commons

What Oase 1 was not

Three qualifications the genome calls for, one by one:

  1. Not the ancestor of Europeans

    His genome is no closer to present-day Europeans than to East Asians. His population did not contribute substantially to later humans in Europe: the genome shows no detectable contribution to later populations.

  2. Not the only mixing

    We all carry the old traces from the Near East. The Oase 1 mixing came later, in Europe, on a branch with no detectable contribution. His genome itself proves that modern humans and Neanderthals crossed on European ground.

  3. Not the only one with a recent Neanderthal ancestor

    Three early modern humans from Bacho Kiro, Bulgaria, each had Neanderthal ancestors a few generations back: recent mixing was no Oase 1 exception. Their genomes show that Europe's first modern humans mixed with Neanderthals, perhaps commonly (Hajdinjak et al. 2021).

The ancient-DNA timeline: 1997–2022

From the first scrap of Neanderthal DNA to the Nobel Prize: 11 stops.

  1. 1997

    The first trace

    Krings and team publish the first Neanderthal mitochondrial DNA sequence: it falls outside present-day human variation, a sign Neanderthals died out contributing no mtDNA.

  2. 2002

    The find

    16 February: the Oase 1 jaw comes to light in Peștera cu Oase, near Anina.

  3. 2003

    The publication

    The jaw appears in PNAS: the oldest certain modern human in Europe.

  4. 2007

    The cranium

    Oase 2 is described in PNAS: an adolescent with a mosaic of modern and archaic traits.

  5. 2010

    The Neanderthal draft

    Green and team publish the Neanderthal genome draft: present-day people outside Africa carry about 2% Neanderthal DNA.

  6. 2010

    Denisova

    A finger bone from Siberia reveals another extinct relative: the Denisovans.

  7. 2014

    Ust'-Ishim

    The genome of the roughly 45,000-year-old Siberian.

  8. 2015

    The Oase 1 genome

    Fu and team show 6–9% Neanderthal DNA and the 4–6-generations-back ancestor.

  9. 2016

    Ice Age Europe

    The genetic history of Ice Age Europe: the study reconstructs shifts and replacements among European populations.

  10. 2021

    Bacho Kiro

    Genomes from the Bulgarian cave show early modern humans with recent Neanderthal ancestors; Oase 1 shows no detectable affinities to later populations.

  11. 2022

    The Nobel Prize

    Svante Pääbo takes the Medicine Nobel for the genomes of extinct hominins.

The quiz: six statements

True or false? Choose, then open the answer.

  1. 1

    Oase 1 is a direct ancestor of present-day Europeans.

    See the answer

    False. His genome resembles Europeans no more than East Asians: it shows no detectable contribution to later populations.

  2. 2

    Oase 1 had a Neanderthal ancestor 4–6 generations back.

    See the answer

    True. The three over-50-centimorgan segments point to a mixing 4–6 generations before him, about 100–180 years earlier at 25–30 years per generation.

  3. 3

    Archaeologists found the jaw in a dig.

    See the answer

    False. Three cavers found it on 16 February 2002, on the cave floor, in the open. The field campaigns with systematic recording and digging followed in 2003–2005.

  4. 4

    Oase 1 was male.

    See the answer

    True. The Y chromosome was read from the jaw: haplogroup F, later refined to K2a*.

  5. 5

    The Oase 2 cranium comes from the same man as the jaw.

    See the answer

    False. The jaw is a grown adult's, the cranium an adolescent's: two people. The Oase 3 temporal belongs with the cranium.

  6. 6

    In 2015, Oase 1 had the largest share of Neanderthal DNA reported in a sequenced Homo sapiens individual.

    See the answer

    True. At its 2015 publication, the 6–9% share was the largest reported in a sequenced Homo sapiens individual.

Sources and method

Most facts about the bones come from the primary papers: Trinkaus et al. 2003 for the discovery, Rougier et al. 2007 for the cranium, Fu et al. 2015 for the genome. The list below groups them by theme, with links to each paper.

Every number on the page shows its working: rounding 40,440 to 40,000, subtracting 2015 minus 2002, the expected 1/2-to-the-n contribution, the 3.75 ratio to the 2 benchmark, the 100–180 years from 4–6 generations of 25–30 years.

The genome diagram is drawn for this page, to scale, from the published 6–9% range. No published figure is copied. From secondary syntheses come: the Miniș Valley and the proximity to Anina, the 12 galleries, the Ion din Anina nickname.

Images

  • A cave-bear skeleton, Ursus spelaeus · Wikipek · CC BY 3.0 · Wikimedia Commons
  • The Oase 2 cranium · Ryan Somma · CC BY-SA 2.0 · Wikimedia Commons
  • The Oase 2 facial reconstruction, Neanderthal Museum, Mettmann · Daniela Hitzemann (photograph) · CC BY-SA 4.0 · Wikimedia Commons
  • A Neanderthal reconstruction · Luna04 · CC BY 2.5 · Wikimedia Commons
  • Skulls compared: Neanderthal and modern human · Jacklee · CC BY-SA 4.0 · Wikimedia Commons