First Upright Apes Likely Evolved in Europe: New Findings Reveal Evolutionary Origins

Illustration of Grecopithecus freybergi, an early ape.

Illustration of Grecopithecus freybergi, a great ape from southeastern Europe 7.2 million years ago

Velizar Simeonovski, Chicago

Discoveries indicate that 7.2 million years ago, early apes might have been walking upright in present-day Bulgaria. Researchers uncovered ancient leg bones exhibiting characteristics of bipedalism.

These leg bones predate all known human fossils, including those from Africa, suggesting that bipedalism—a critical milestone in human evolution—may have originated in Europe rather than Africa.

“The earliest signs of bipedalism are found in Europe,” states Madeleine Boehme, from the University of Tübingen, Germany.

Boehme and a dedicated team have been excavating at Azmaka near Chirpan, Bulgaria, since 2008, uncovering a sediment layer approximately 20 meters thick, deposited by rivers over time.

A crucial find in 2016 was a right femur, known to be 7.2 million years old and nearly intact, nicknamed “Diva.” This femur, measuring 21.5 centimeters, offers substantial insight into the locomotion of early humans.

“This discovery is incredibly significant,” remarks Clement Zanoli from the University of Bordeaux, France, who was not involved in the study. “The femur’s exceptional preservation is rare in the Miocene hominin fossil record.”

During early epochs, Europe hosted various great apes, yet by 7.2 million years ago, most had become extinct. Today, all closest relatives to modern apes are located in Africa, as are the earliest human ancestors, or hominins.

The only ape confirmed from Azmaka is Grecopithecus freybegii, known primarily from a damaged jawbone found in Greece, and some teeth from North Macedonia and Azmaka. Consequently, Böhme’s team posits that the femur likely belongs to Grecopithecus.

“This is the most reasonable assumption at this time,” states Zanoli, but he stresses that more fossil evidence is necessary for stronger validation.

“The relationships among these fossils remain unclear,” explains Kelsey Pugh from OCAD University, Toronto, Canada.

Femoral bones of Grecopithecus (left), Lucy (Australopithecus afarensis, center), and a chimpanzee (right). Long femoral neck (shown in red) associated with upright walking.

Spasov et al. 2026

Böhme’s team meticulously measured the femur and conducted CT scans, uncovering attributes indicative of a bipedal organism. Notably, a short neck that laterally extends and forms a rounded ball fits into the pelvis, resembling structures found in bipedal hominids but absent in knuckle-walking species. Additionally, the outer bone layer’s thickness at the neck’s base supports vertical loads.

An additional ridge was identified, which would have allowed for the attachment of buttock muscles, playing a vital role in maintaining an upright posture, according to Boehme.

While other scientists find the findings intriguing, they remain unconvinced. “This femur displays several traits,” Zanoli points out. “Some suggest bipedalism, while others imply quadrupedal movement. Hence, determining the locomotion remains complex.”

The nuanced study of extinct great apes has revealed increasing challenges in accurately diagnosing bipedalism based solely on isolated fossil bones. Pugh notes, “Many features once thought exclusive to bipedal hominids have also been noted in quadrupedal apes. Thus, prioritization of diagnostic traits is crucial.”

Much attention has focused on Sahelanthropus tchadensis, known from a solitary location in Chad and considered the earliest hominid. This species lived 7 million years ago, slightly younger than the Azmaka specimen. The femur of Sahelanthropus has long fueled debate about bipedalism.

Boehme and her team are determined to establish that significant steps in early human evolution transpired in Europe. They’ve identified potential hominid features in Grecopithecus, despite past controversies linked to a damaged jawbone. Furthermore, they have also detailed another European ape, Danuvius guggenmosi, which existed approximately 11.6 million years ago.

These early human-like ancestors might have migrated to Africa, potentially driven by climate changes. Research indicates that this migration likely gave rise to all subsequent hominins, including modern humans.

Zanoli comments, “Numerous other animals migrated between Africa and Eurasia. If fauna can do it, why can’t hominins?”

Yet, Professor Pugh asserts that further discoveries are essential to substantiate bipedalism evidence in Grecopithecus, allowing detailed analysis of relationships among other apes and early humans. Without this, she emphasizes that it’s premature to construct comprehensive evolutionary narratives.

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New Study Uncovers How the Pelvis Evolved and Helped Our Ancestors Stand Upright

The pelvis is often described as the keystone for upright locomotion. Over millions of years, it has undergone significant transformations, enabling us to walk on two legs more effectively than any other part of our lower body. However, the specifics of this remarkable adaptation have been largely unknown. Recent research has uncovered two crucial genetic changes that reshaped the pelvis, permitting it to evolve into the upright structure that our ancestors utilized while traversing the Earth.

Ardipithecus ramidus Humanity resided in Africa over 4 million years ago. Illustrations by Arturo Asensio, via Quo.es

Anatomists have long recognized that the human pelvis is distinct among primates.

In our closest relatives, African apes (chimpanzees, bonobos, and gorillas) possess hipbones that are tall, narrow, and flat from front to back. When viewed from the side, they resemble thin blades.

The pelvic structure of an ape supports large muscles essential for climbing.

In contrast, human hip bones rotate sideways, forming a bowl shape. This flaring of the hip bones allows for muscle attachment critical for maintaining balance while shifting weight from one foot to the other during upright locomotion.

Nonetheless, the mechanisms behind this transformation have been elusive until now.

In a recent study, Professor Terrence Capelini from Harvard University and his team pinpointed vital genetic and developmental shifts that facilitated the evolution from the pelvis of tetraleaf monkeys to bipedalism.

“Our findings illustrate a complete mechanistic shift in human evolution,” stated Professor Capelini.

“There is no parallel to this among other primates.”

The researchers analyzed 128 samples of embryonic tissue from humans housed in museums in the US and Europe, along with nearly 20 other primate species.

These collections included specimens over 100 years old, preserved on glass slides or in bottles.

Using CT scans and histological analysis, they investigated pelvic anatomy during the early stages of development.

Their research revealed that the evolution of the human pelvis unfolded in two major phases.

Initially, the growth plate shifted 90 degrees, widening the human ilium instead of extending its height.

Following this adjustment, the timeline for embryonic bone formation was altered.

Typically, bones in the lower body develop when chondrocytes align along the long axis of the growing bone.

This cartilage becomes rigid through a process known as ossification.

At the early stages of development, similar to other primates, human growth plates formed from the head and continued to develop.

However, by day 53, the growth plate had notably shifted vertically from its initial orientation, resulting in a shorter and broader hip joint.

“When I examined my pelvis, it wasn’t initially on my radar,” Professor Capelini remarked.

“I anticipated a gradual modification to shorten and widen it, but histology indicated a complete 90-degree reversal.”

Group of Australopithecus afarensis. Image credit: Matheus Vieira.

A further significant alteration was the timeline of bone formation.

In most cases, bones develop along the primary ossification center located in the center of the bone shaft.

However, in humans, the ilium diverges from this norm, with ossification beginning at the posterior region in the sacrum and expanding radially.

This mineralization remains restricted to the peripheral layer, while internal ossification is postponed by 16 weeks, allowing bones to grow and maintain their shape during their geometric transitions.

To uncover the molecular mechanisms driving these changes, the team employed techniques like single-cell multiomics and spatial transcriptomics.

The researchers identified over 300 active genes, including three with notable roles: Sox9 and PTH1R (which control growth plate shifts) and runx2 (which governs ossification changes).

The significance of these genes is underscored by diseases arising from their dysfunction.

For example, mutations in Sox9 can lead to Campomelic dysplasia, a disorder characterized by an abnormally narrow hip joint lacking lateral flaring. Similarly, mutations in PTH1R result in narrow hip joints and various skeletal disorders.

The scientists propose that these adaptations began with the reorientation of the growth plate around the time our ancestors separated from African apes, estimated to have occurred between 5 and 8 million years ago.

They believe the pelvis has served as a focal point for evolutionary transformations over millions of years.

As brain size increased, the pelvis encountered selective pressures known as the obstetric dilemma—the trade-off between a narrow pelvis for efficient movement and a broader one for accommodating the birth of larger babies.

Researchers suspect that the delay in ossification likely occurred within the last two million years.

The oldest pelvic fossil, dated at 4.4 million years, belongs to Ardipithecus from Ethiopia—a species exhibiting a blend of upright walking and tree-climbing features, with pelvic characteristics akin to those of humans.

The renowned 3.2 million-year-old skeleton of Lucy (Australopithecus afarensis) showcases further adaptations for bipedalism, including the distinctively flaring hip blades.

“From that point onwards, all hominin fossils displayed pelvises that diverged significantly from those of earlier primates,” stated Professor Capelini.

“The implications of brain size and its subsequent changes should not be interpreted through growth models applicable to chimpanzees and unassociated primates.”

“Models should focus on the developments between humans and their own lineage.”

“Post-fetal growth occurred against the backdrop of novel methods for constructing the pelvis.”

This study is set to be published in the journal Nature.

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G. Senevilas et al. The evolution of hominin bipedal walking in two steps. Nature Published online on August 27th, 2025. doi:10.1038/s41586-025-09399-9

Source: www.sci.news