Neanderthal-like Features in 450,000-Year-Old Fossil Teeth from Italy

Fossil teeth from Italy, among the oldest human remains on the Italian Peninsula, show that Neanderthal dental features had evolved by around 450,000 years ago, according to a study published October 3, 2018 in the open-access journal PLOS ONE by Clément Zanolli of the Université Toulouse III Paul Sabatier in France and colleagues. These teeth also add to a growing picture of a period of complex human evolution that we are only beginning to understand.

Virtual rendering of the Visogliano and Fontana Ranuccio teeth.

Zanolli and colleagues examined dental remains from the sites of Fontana Fanuccio, located 50km southeast of Rome, and Visogliano, located 18km northwest of Trieste. At around 450,000 years old, these teeth join a very short list of fossil human remains from Middle Pleistocene Europe. Using micro-CT scanning and detailed morphological analyses, the authors examined the shape and arrangement of tooth tissues and compared them with teeth of other human species. They found that the teeth of both sites share similarities with Neanderthals and are distinct from modern humans.

There has been much debate over the identities and relationships of Middle Pleistocene ancient humans in Eurasia. The discovery of Neanderthal-like teeth so early in the record adds support to the suggestion of an early divergence of the Neanderthal lineage from our own, around the Early-Middle Pleistocene transition. The teeth are also notably different from other teeth known from this time in Eurasia, suggesting that there may have been multiple human lineages populating the region at this time, adding to a growing list of evidence that the Middle Pleistocene was a time of more complex human evolution than previously recognized.

“The remains from Fontana Ranuccio and Visogliano represent among the oldest human fossil remains testifying to a peopling phase of the Italian Peninsula.” Zanolli said, “Our analyses of the tooth internal structural organization reveal a Neanderthal-like signature, also resembling the condition shown by the contemporary assemblage from Atapuerca Sima de los Huesos, indicating that an overall Neanderthal morphological dental template was preconfigured in Western Europe at least 430 to 450 ka ago.”

Citation: Zanolli C, Martinón-Torres M, Bernardini F, Boschian G, Coppa A, Dreossi D, et al. (2018) The Middle Pleistocene (MIS 12) human dental remains from Fontana Ranuccio (Latium) and Visogliano (Friuli-Venezia Giulia), Italy. A comparative high resolution endostructural assessment. PLoS ONE 13(10): e0189773.

Funding: Publication of this work was supported by the Centro Fermi (Roma). The ICTP/Elettra. EXACT Project has been funded by the Regione Friuli-Venezia Giulia. We acknowledge the excavation team from Atapuerca-Sima de los Huesos for their arduous work. The excavation, scanning and analysis of the Atapuerca sample has been funded by the CGL2015-65387-C3-1, 3-P project of the Ministerio de Economía y Competitividad (MINECO) and CEN074A12 from Consejería de Educación from Junta de Castilla y León Government, Spain. Gordon Getty (2013) and Dub Crook (2014, 2015, 2016) of The Leakey Foundation provided financial support to one of the authors (MMT). MMP has a predoctoral contract from the Junta de Castilla y León (BOCYL-D-20052013-14), co-financed by the European Social Funds and LMF had a postdoctoral research grant from Fundación Atapuerca.

Grantee Spotlight: Sofya Dolotovskaya

Sofya Dolotovskaya of the German Primate Center was awarded a Leakey Foundation Research Grant during our spring 2018 cycle for her project entitled “Does pair-living translate into genetic monogamy in a Neotropical primate?”

Sofya Dolotovskaya recording titi monkey vocalizations in the Peruvian Amazon rainforest.

In my project, I want to find out if pair-living is associated with genetic monogamy in a small Neotropical primate, red titi monkey (Callicebus cupreus). Titi monkeys are a textbook example of a “monogamous” primate. They live in apparently perfect families: mother, father, and several offspring. But are these families really that perfect, or do mates cheat on each other? That’s the main question of my project.

Social monogamy, or pair-living, is a system where male and female live as pairs. It is very common among birds, but extremely rare in mammals (only 3-9% of species). Social monogamy, however, doesn’t imply genetic monogamy – a mating system in which a male and female copulate exclusively within the pair. For example, 90% of passerine birds were originally described as monogamous – but genetic studies revealed that most of them engage in extra-pair copulations. In mammals, genetic monogamy has been so far reported for only seven species: the Malagasy giant jumping rat (Hypogeomys antimena), California mouse (Peromyscus californicus),  oldfield mouse (Peromyscus polionotus), Azara’s owl monkey (Aotus azarae), Eurasian beaver (Castor fiber), pack-living coyotes (Canis latrans), and Kirk’s dik-dik (Madoqua kirkii).

Top row: the Malagasy giant jumping rat (CC-BY-SA 3.0 Manfred Werner), California mouse (CC-BY-SA 3.0 Whatiguana), Azara’s owl monkey (CC-BY 2.0 Rich Hoyer). Bottom row: Eurasian beaver (CC-BY-SA 3.0 Per Harald Olsen), coyote (CC-BY-SA 3.0 Manfred Werner), and Kirk’s dik-dik (CC-BY-SA 3.0 Yathin S Krishnappa).

In primates, social monogamy is more common than in other mammalian orders. But so far, genetic monogamy has been confirmed only for Azara’s owl monkey. Like owl monkeys, red titis have stable pair bonds and extensive rates of paternal care: the infants are carried exclusively by males. The home ranges are small and have very little overlap between the groups, and direct intergroup encounters were never observed. Overall, mated pairs seem to have very few opportunities for extra-pair matings. Therefore, I expect genetic monogamy or very low rate of extra-pair paternities in red titis.

A red titi monkey resting in a tree. Photo: Sofya Dolotovskaya

Apart from the general question of genetic monogamy, I’m interested in more specific mechanisms of monogamy maintenance. How is the spacing between groups defined? How do the individuals disperse, and what are the relatedness patterns within and between the pairs? Is there interbreeding avoidance as a spacing mechanism? Are more closely related pairs less territorial then more distantly related ones?

Two red titi monkeys sitting together with their tails intertwined at the London Zoo. Photo CC-BY-SA 3.0 by Steven G. Johnson.

To address all these questions, I study a wild population of titi monkeys in north-eastern Peruvian Amazonia. Together with field assistants, we habituate and then follow titi monkey families. We collect behavioral, vocal, and spatial data, along with fecal samples for genetic analyses. Once back from the field, I will use DNA extracted from the fecal samples to do the parentage and relatedness analyses using microsatellite genotyping.

Information on the genetic consequences of pair-living in titi monkeys can help us to understand the evolution of monogamy in New World primates and eventually in primates in general. This question is of particular interest to human evolution because monogamy has evolved multiple times in humans, especially in recent centuries. Understanding the evolutionary history of social monogamy in other mammals may provide insights into the selective pressures that led to the emergence of stable pair bonds in humans.

Mountain Gorilla Population Passes 1,000

September 24 marks World Gorilla Day, a day to celebrate our close ape cousins. It also commemorates the day Dian Fossey established the Karisoke Research Center in the Virunga Mountains of Rwanda. 

All species of gorillas are critically endangered according to the Red List maintained by the International Union for the Conservation of Nature (IUCN), but that does not mean there’s no hope for these animals.


Mountain gorilla infant. Photo courtesy of the Dian Fossey Gorilla Fund.

According to the results of a census published in May 2018, the mountain gorilla population in East Africa’s Virunga Mountains numbered 604 as of June 2016, up from 480 in 2010. The population hit an all-time low of 242 individuals in 1981.

The mountain gorilla (Gorilla beringei beringei) is a subspecies of the eastern gorilla with two distinct sub-populations: one in the Virunga Mountains and another in Uganda’s Bwindi Impenetrable National Park. A census conducted in 2011 found approximately 400 gorillas living in Bwindi Impenetrable National Park, meaning that the total number of mountain gorillas is now believed to be more than 1,000 individuals.

According to the Dian Fossey Gorilla Fund International, the recent census of the Virunga sub-population confirms that the mountain gorilla is “the only wild ape population whose numbers are known to be increasing.” The group also noted that research has shown that the increasing mountain gorilla population is due largely to “intensive daily protection” provided by national park authorities and other conservationists.

Dr. Tara Stoinski, President, CEO, and Chief Scientific Officer of the Fossey Fund, which was a partner organization involved in the census, said that the results represent “a huge success for conservation at a time when such success stories are increasingly rare.” Stoinski added that “All those working to protect mountain gorillas — the governments of Rwanda, Uganda and [Democratic Republic of the] Congo; conservation organizations; and local communities — have a lot to be proud of.”

Members of one of the survey teams that performed the census of mountain gorillas. They found 604 individuals are living in the Virunga Mountains, bringing the total mountain gorilla population above 1,000. Photo courtesy of the Dian Fossey Gorilla Fund.

The mountain gorilla is listed as Critically Endangered on the IUCN Red List, though the IUCN’s last assessment of the subspecies’ threat status was completed in 2008.

Mountain gorillas’ conservation status is still quite perilous, however, as they continue to face numerous threats due to their limited habitat and increasing pressures from encroaching human populations. Wire and rope snares set for game like antelope, in particular, pose a significant danger. The survey teams who performed the census said they discovered more than 380 snares, which they dismantled — though they were too late in at least one case in which they came upon a snare that already contained a dead gorilla.

The long-running conflict in eastern Democratic Republic of the Congo (DRC) often spills over into Virunga National Park, as well, taking a further toll on wildlife. Earlier this year, five rangers and a driver were killed in an ambush by suspected members of an armed militia in what’s been called the deadliest attack in Virunga National Park’s history.

Mountain gorilla silverback male. Photo courtesy of the Dian Fossey Gorilla Fund.

The IUCN released a report in January showing that the park, which was founded in 1925 as a means of protecting the mountain gorillas living in the forests of the Virunga Mountains, is one of the most threatened natural World Heritage Sites in the world.

“The continued growth of the mountain gorilla population is a fervent reminder of the importance of continued conservation strategies,” Virunga National Park’s director and chief warden, Emmanuel de Merode, said in a statement. “In light of hardships the Park has suffered, this news is incredibly important to us.”

The census of the Virunga sub-population was led by protected area authorities with the governments of the DRC, Rwanda, and Uganda, the three range states of the mountain gorilla, and supported by the International Gorilla Conservation Programme, a coalition of Fauna & Flora International and WWF.

Members of a survey team with mountain gorillas. Photo courtesy of the Dian Fossey Gorilla Fund.

Conservationists were quick to celebrate the increasing mountain gorilla population as a much-needed instance of good news, even if they remain wary of the many persistent and looming threats the subspecies must still contend with.

“This is fabulous news for mountain gorillas and shows what we can do for wildlife when NGOs, governments and their communities work together,” Margaret Kinnaird, Wildlife Practice Leader at WWF, said in a statement.

“However, the high number of snares encountered and the numerous other threats they face including climate change indicate that the battle is far from won. The three gorilla range countries and their partners must continue to work together to safeguard the Virunga Massif — not only for the protection of these incredible creatures but also for the welfare of the local people with whom they share the landscape. The mountain gorilla story can be a model for how to restore and maintain our earth’s precious biodiversity.”

Sir David Attenborough even weighed in, saying in a statement: “When I first visited the mountain gorillas in 1979, the situation was dire; the number of these remarkable animals was dreadfully small. It is incredibly heartening therefore to see how the efforts of so many different groups — communities, governments, NGOs — have paid off. The threats to mountain gorillas haven’t disappeared entirely, of course, so now the challenge must be to ensure that these achievements are sustained long into the future.”

The Leakey Foundation has funded research on gorillas since our beginning in 1968. You can help!


• Robbins, M., Gray, M., Kümpel, N., Lanjouw, A., Maisels, F., Mugisha, A., Spelman, L. & Williamson, L. 2008. Gorilla beringei ssp. beringei. The IUCN Red List of Threatened Species 2008: e.T39999A10292321. doi:10.2305/IUCN.UK.2008.RLTS.T39999A10292321.en. Downloaded on 05 June 2018.

This article was written by Mike Gaworecki for and was republished with a Creative Commons license. Click here to read the original article.

California Academy of Sciences Welcomes New Anthropology Curator

Dr. Todd Braje focuses on coastal ecosystems. Photo © 2018 California Academy of Sciences

The Leakey Foundation is excited to welcome Dr. Todd Braje as the new Irvine Chair of Anthropology at the California Academy of Sciences’ Institute for Biodiversity Science and Sustainability. This position was previously held by Leakey Foundation grantee Dr. Zeray Alemseged (2008-2017) and Leakey Foundation grantee and Scientific Executive Committee Member Dr. Nina Jablonski (1995-1998). The Leakey Foundation looks forward to continuing our long tradition of collaborating with the California Academy of Sciences to produce fascinating and informative human evolution lectures.

Using archaeological sites as his guide, Braje interprets millennia of human interaction with marine and terrestrial ecosystems to establish historical baselines for coastal resource managers. In his new role, Braje will draw from the Academy’s collections to understand how coastal ecosystems—particularly those in California—are changing, how humans have shaped these environments, and how past interactions can inform future management.

“I find the Academy’s sustainability mission incredibly inspiring,” says Braje, most recently a professor at San Diego State University. “Although I study deep history, my goal is to carve a better path forward based on our understanding of past human-environment interactions.”

“Todd brings an invaluable perspective to the Academy,” says Dr. Shannon Bennett, Academy Chief of Science. “As an anthropologist, his ability to blend an historical perspective of human interactions with the environment is critical to how we understand our collective impact on the planet. Todd has a unique role that will touch many different collections, not just anthropology. He is painting a detailed picture of how human relationships with coastal resources are changing, particularly in California, and what we can do differently to ensure these environments persist into the future.”

Chasing an eroding heritage

Born and raised in Indiana, Braje eventually found his way to the sea. After graduating with a degree in anthropology, he followed a Peace Corps assignment to the Kingdom of Tonga, a Polynesian archipelago in the South Pacific. Here he drew inspiration from outside the walls of traditional academia, exploring Tonga’s vibrant communities by bike and foot.

“In Tonga, my homestay family taught me about their culture by taking me to ancestral sites. Ultimately this is what inspired me to pursue coastal archaeology and island historical ecology,” says Braje, who later earned master’s and doctorate degrees in anthropology.

But with each passing year, Braje’s chosen career path presents a growing challenge. “Every time I go to the field I see our human heritage eroding into the ocean, and very little being done to stop it,” he says. “Rising sea levels, accelerated by human-caused climate change, are exacerbating erosion and undercutting coastal sites. In one moment, we can have an entire archaeological site calve off into the ocean before anyone has dated or sampled it.”

As sites crumble and history washes out to sea, answers for how to manage the future are also vanishing. “We’re losing a critical toolkit in our fight to build future sustainable systems. Deep history tells us what the world once looked like before the dramatic impacts of modern day. Without this perspective, it’s difficult to know where we need to focus. Most of human history,” he adds, “is not written down.”

Braje recognizes the importance of his role: As a steward of coastal history, he collaborates with indigenous communities, ecologists, policymakers, and other local stakeholders to keep history alive by documenting oral traditions and studying archeological sites.

Tracing the kelp highway

Though rising sea levels have inundated many once-dry regions along the coast, Braje isn’t deterred—he has simply moved his research underwater. He and his colleagues are scouring the ancient coastlines of central Oregon and the Channel Islands for submerged archaeological sites, conducting shipboard sonar surveys and probing the seabed for sediment cores. “It’s the world’s biggest dartboard,” he says, “but cores extracted from the seabed can hold important historical artifacts from coastal populations that came before us.”

Also submerged are clues to understanding early maritime migrations—how, when, and where the first people arrived in the Americas. In a dramatic shift from just a couple decades ago when most archaeologists believed people walked into the New World, many now believe that modern humans first arrived from Northeast Asia by boat, journeying along a great “kelp highway” that provided an abundance of resources for the intrepid explorers. Braje is retracing their trek, looking for potential “pit stops” along the way that would indicate where populations fished, foraged, and hunted.

Fishing for a baseline

Despite anthropology’s historical roots, Braje’s work has critical modern applications for coastal resource management. His study of former fisheries has lent important insights to Channel Islands policymakers, and his burgeoning projects in both the Solomon and Coronados Islands are designed to do the same.

In the Channel Islands—a Southern California archipelago—humans have fished abalone for more than ten thousand years. Although abalone populations have experienced rapid declines in recent years, some species like red abalone are showing signs of recovery. By examining ancient shell middens (archaeological trash piles) and other deep historical evidence, Braje can trace millennia-long patterns of when and where abalone thrived along the Channel Islands and were available to human fishers. His findings provide a critical perspective that will help policymakers decide when red abalone is recovered and viable for fishing again.

Now in the Bay Area, Braje will expand his research to the unique islands and coastal ecosystems of the San Francisco Bay and beyond.

“The San Francisco Bay has been utterly transformed by a combination of natural and cultural forces over the last 10,000 years. If we want to protect our precious bay and coastline and still build sustainable systems, we must first understand the unique human-environment dynamics that have shaped our home,” says Braje. “My goal is to explore best practices for coastal stewardship by understanding how those who came before us looked after and shaped this special place.”

Guenon Monkeys Cross Species Boundary

“Jimmy” is a hybrid male monkey in Dr. Kate Detwiler’s study group in Gombe National Park. Photo credit: Manano Mpongo/ Gombe Hybrid Monkey Project

By Gisele Galoustian

Leakey Foundation grantee Kate Detwiler from Florida Atlantic University is the first to document that two genetically distinct species of guenon monkeys inhabiting Gombe National Park in Tanzania, Africa, have been successfully mating and producing hybrid offspring for hundreds maybe even thousands of years. Her secret weapon? Poop.

Prior studies and conventional wisdom have suggested that the physical characteristics of guenon monkeys with a variety of dazzling colors and very distinct facial features like bushy beards and huge nose spots are a function of keeping them from interbreeding. The idea is that their mate choices and the signals they use to select a mate are species specific and that they share common traits linked to their species.

So if their faces don’t match, they shouldn’t be mating, right? Wrong, according to evidence from a novel study published in the International Journal of Primatology.

For the study, Detwiler, who first studied these monkeys in Gombe National Park in 1994, examined the extent and pattern of genetic transfer or gene flow from “red-tailed” monkeys (Cercopithecus ascanius) to “blue” monkeys (Cercopithecus mitis) due to hybridization.

These two species are the only forest guenons that colonized the narrow riverine forests along Lake Tanganyika that characterize Gombe National Park. They co-exist in the same forests as Jane Goodall’s chimpanzees and baboons. Detwiler identifies hybrid monkeys by their combined markings from both parental species. She estimates that about 15 percent of this population is made up of hybrids, which is very unusual.

Kate Detwiler, Ph.D., lead author and an assistant professor in the Department of Anthropology in FAU’s Dorothy F. Schmidt College of Arts and Letters, who first studied these monkeys in 1994, collects fecal samples from monkeys in Gombe National Park. Photo credit: Amelia Villaseñor

Using mitochondrial DNA, extracted non-invasively from the feces of 144 red-tailed monkeys, blue monkeys, and hybrids, Detwiler is the first to show the movement of genetic material from one guenon species to another in an active hybrid zone. After examining the fecal samples, she found that all of the monkeys – hybrids, red-tails and blues have red-tailed mitochondrial DNA – all traced back to female red-tailed monkeys.

For this lineage of monkeys, it is the first time that science shows that not only is the DNA there, but so are the hybrids. Detwiler used mitochondrial DNA because it is more abundant than nuclear DNA in fecal samples, and only comes from the mother – indicating the maternal species in the hybridizing pair.

“There’s a lot of promiscuity taking place in Gombe National Park. Red-tails are mating with blues, blues are mating with red-tails, blues are mating with blues, red-tails are mating with red-tails, and hybrids are mating with everyone,” said Detwiler. “But we’re just not seeing any negative consequences from these two very different species repeatedly mating and producing offspring on an ongoing basis. If the differences in their facial features are so important and signal that they shouldn’t be mating, then why is this happening and why do I keep finding hybrid infants?”

A key finding from the study shows that the blue monkeys in Gombe National Park emerged out of the hybrid population, tracing their origins back to hybridization events between resident red-tails and blues most likely from outside the park. For her control groups, Detwiler collected and examined feces from blue monkeys from a park to the north and a park to the south where hybrids do not exist. These monkeys only had blue monkey mitochondrial DNA.

Detwiler speculates that red-tailed monkeys got to Gombe National Park first and thrived in the environment. Male blue monkeys outside the park had to find new homes after they were kicked out of their groups, which happens when they reach sexual maturity. Sex-driven, they ventured out into the landscape to find appropriate mates – female blue monkeys. Instead, they found the red-tailed females. Apparently, some female red-tailed monkeys were attracted to novel males with different faces and welcomed the sexual advances from these male blue monkeys.

“I keep coming back to the idea that if they are only supposed to mate with their own kind, then why did these red-tailed monkeys mate with the blue monkeys, especially if they had males of their own species around,” said Detwiler. “The female red-tailed monkeys present as willing partners and they are not coerced or forced into copulation with blue monkeys.”

Today, Gombe is an isolated forest habitat. Because they are very social and have had to share close quarters for decades or even centuries, Detwiler believes that they have socially learned that if you grow up in a hybrid group it is okay to mate with everyone.

“The Gombe hybrid population is extremely valuable because it can be used as a model system to better understand what hybridization looks like and how genetic material moves between species,” said Detwiler. “We have this amazing laboratory in nature to help us answer many questions about hybridization and how species boundaries are maintained. This research is very timely because hybridization often occurs in response to environmental changes, as we are seeing with climate change and modified landscapes — it is nature’s way to respond.”

This work was funded by a faculty research seed grant from FAU’s Division of Research, FAU’s Department of Anthropology, and doctoral dissertation research grants from the NSF (0424444), The Leakey Foundation, and the Wenner-Gren Foundation.

Video credit: Maneno Mpongo and Elizabeth Tapanes/Gombe Hybrid Monkey Project