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One of the best ways to study human evolution is by comparing us with nonhuman species that, evolutionarily speaking, are closely related to us. That closeness can help scientists narrow down precisely what makes us human, but that scope is so narrow it can also be extremely hard to define. To address this complication, researchers from Stanford University have developed a new technique for comparing genetic differences.

An image, from previous research, of human cortical spheroids derived in the lab of Sergiu Pașca, associate professor of psychiatry and behavioral sciences. (Image credit: Timothy Archibald)

Through two separate sets of experiments with this technique, the researchers discovered new genetic differences between humans and chimpanzees. They found a significant disparity in the expression of the gene SSTR2 – which modulates the activity of neurons in the cerebral cortex and has been linked, in humans, to certain neuropsychiatric diseases such as Alzheimer’s dementia and schizophrenia – and the gene EVC2 , which is related to facial shape. The results were published March 17 in Nature and Nature Genetics , respectively.

“It’s important to study human evolution, not only to understand where we came from, but also why humans get so many diseases that aren’t seen in other species,” said Rachel Agoglia, a recent Stanford genetics graduate student who is lead author of the Nature paper.

The Nature paper details the new technique, which involves fusing human and chimpanzee skin cells that had been modified to act like stem cells – highly malleable cells that can be prodded to transform into a variety of other cell types (albeit not a full organism).

“These cells serve a very important specific purpose in this type of study by allowing us to precisely compare human and chimpanzee genes and their activities side-by-side,” said Hunter Fraser, associate professor of biology at Stanford’s  School of Humanities and Sciences . Fraser is senior author of the Nature Genetics paper and co-senior author of the Nature paper with Sergiu Pașca, associate professor of psychiatry and behavioral sciences in the Stanford  School of Medicine .

Close comparisons

The Fraser lab is particularly interested in how the genetics of humans and other primates compare at the level of cis-regulatory elements, which affect the expression of nearby genes (located on the same DNA molecule, or chromosome). The alternative – called trans-regulatory factors – can regulate the expression of distant genes on other chromosomes elsewhere in the genome. Due to their broad effects, trans-regulatory factors (such as proteins) are less likely to differ among closely related species than cis-regulatory elements.

But even when scientists have access to similar cells from humans and chimpanzees, there is a risk of confounding factors. For example, differences in the timing of development between species is a significant hurdle in studying brain development, explained Pașca. This is because human brains and chimpanzee brains develop at very different rates and there is no exact way to directly compare them. By housing human and chimpanzee DNA within the same cellular nucleus, scientists can exclude most confounding factors.

For the initial experiments using these cells, Agoglia coaxed the cells into forming so-called cortical spheroids or organoids – a bundle of brain cells that closely mimics a developing mammalian cerebral cortex. The Pașca lab has been at the forefront of developing brain organoids and assembloids for the purpose of researching how the human brain is assembled and how this process goes awry in disease.

“The human brain is essentially inaccessible at the molecular and cellular level for most of its development, so we introduced cortical spheroids to help us gain access to these important processes,” said Pașca, who is also the Bonnie Uytengsu and Family Director of Stanford Brain Organogenesis.

As the 3D clusters of brain cells develop and mature in a dish, their genetic activity mimics what happens in early neurodevelopment in each species. Because the human and chimpanzee DNA are bound together in the same cellular environment, they are exposed to the same conditions and mature in parallel. Therefore, any observed differences in the genetic activity of the two can reasonably be attributed to actual genetic differences between our two species.

Through studying brain organoids derived from the fused cells that were grown for 200 days, the researchers found thousands of genes that showed cis-regulatory differences between species. They decided to further investigate one of these genes – SSTR2 – which was more strongly expressed in human neurons and functions as a receptor for a neurotransmitter called somatostatin. In subsequent comparisons between human and chimpanzee cells, the researchers confirmed this elevated protein expression of SSTR2 in human cortical cells. Further, when the researchers exposed the chimpanzee cells and human cells to a small molecule drug that binds to SSTR2 , they found that human neurons responded much more to the drug than the chimpanzee cells.

This suggests a way by which the activity of human neurons in cortical circuits can be modified by neurotransmitters. Interestingly, this neuromodulatory activity may also be related to disease since SSTR2 has been shown to be involved in brain disease.

“Evolution of the primate brain may have involved adding sophisticated neuromodulatory features to neural circuits, which under certain conditions can be perturbed and increase susceptibility to neuropsychiatric disease,” said Pașca.

Fraser said these results are essentially “a proof of concept that the activity we’re seeing in these fused cells is actually relevant for cellular physiology.”

Investigating extreme differences

For the experiments published in Nature Genetics , the team coaxed their fused cells into cranial neural crest cells, which give rise to bones and cartilage in the skull and face, and determine facial appearance.

“We were interested in these types of cells because facial differences are considered some of the most extreme anatomical differences between humans and chimps – and these differences actually affect other aspects of our behavior and evolution, like feeding, our senses, brain expansion and speech,” said David Gokhman, a postdoctoral scholar in the Fraser lab and lead author of the Nature Genetics paper. “Also, the most common congenital diseases in humans are related to facial structure.”

In the fused cells, the researchers identified a gene expression pathway that is much more active in the chimpanzee genes of the cells than in the human genes – with one specific gene, called EVC2 , appearing to be six times more active in chimpanzees. Existing research has shown that people who have inactive EVC2 genes have flatter faces than others, suggesting that this gene could explain why humans have flatter faces than other primates.

What’s more, the researchers determined that 25 observable facial features associated with inactive EVC2 are noticeably different between humans and chimpanzees – and 23 of those are different in the direction the researchers would have predicted, given lower EVC2 activity in humans. In follow-up experiments, where the researchers reduced the activity of EVC2 in mice, the rodents, too, developed flatter faces.

Another tool in the toolbox

This new experimental platform is not intended to replace existing cell comparison studies, but the researchers hope it will support many new findings about human evolution, and evolution in general.

“Human development and the human genome have been very well studied,” said Fraser. “My lab is very interested in human evolution, but, because we can build on such a wealth of knowledge, this work can also reveal new insights into the process of evolution more broadly.”

Looking forward, the Fraser lab is working on differentiating the fused cells into other cell types, such as muscle cells, other types of neurons, skin cells and cartilage to expand their studies of uniquely human traits. The Pașca lab, meanwhile, is interested in investigating genetic dissimilarities related to astrocytes – large, multi-functional cells in the central nervous system often overlooked by scientists in favor of the flashier neurons.

“While people often think about how neurons have evolved, we should not underestimate how astrocytes have changed during evolution. The size difference alone, between human astrocytes and astrocytes in other primates, is massive,” said Pașca. “My mentor, the late Ben Barres, called astrocytes ‘the basis of humanity’ and we absolutely think he was onto something.”

Additional Stanford co-authors for the Nature paper are former research assistant Danqiong Sun, postdoctoral scholar Fikri Birey, senior research scientist Se-Jin Yoon, postdoctoral scholar Yuki Miura and former research associate Karen Sabatini.

This work was funded by a Stanford Bio-X Interdisciplinary Initiatives Seed Grant, the National Institutes of Health, the Department of Defense, the Stanford Center for Computational, Evolutionary and Human Genomics, the Stanford Medicine’s Dean’s Fellowship, MCHRI, the American Epilepsy Society, the Stanford Wu Tsai Neurosciences Institute’s Big Idea Grants on Brain Rejuvenation and Human Brain Organogenesis, the Kwan Research Fund, the New York Stem Cell Robertson Investigator Award, and the Chan Zuckerberg Ben Barres Investigator Award.

Additional Stanford co-authors for the Nature Genetics paper are graduate student Maia Kinnebrew; former undergraduate Wei Gordon; former technician Danqiong Sun; postdoctoral research fellows Vivek Bajpai and Sahin Naqvi; Dmitri Petrov, the Michelle and Kevin Douglas Professor in the School of Humanities and Sciences; Joanna Wysocka, the Lorry Lokey Professor and professor of developmental biology; and Rajat Rohatgi, associate professor of biochemistry and of medicine. Researchers from University of California, San Francisco; University of Michigan, Ann Arbor; Yerkes National Primate Research Center; Emory University School of Medicine; and University of Pennsylvania are also co-authors.

This work was funded by the Human Frontier, Rothschild and Zuckerman fellowships, and the National Institutes of Health.

Fraser is a member of  Stanford Bio-X , the Maternal & Child Health Research Institute (MCHRI) , and the Stanford Cancer Institute . Pașca is a member of Stanford Bio-X, MCHRI and the Wu Tsai Neurosciences Institute , and a faculty fellow of Stanford ChEM-H .

To read all stories about Stanford science, subscribe to the biweekly  Stanford Science Digest .

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Taylor Kubota, Stanford News Service: (650) 724-7707;  [email protected]

Journal of Human Evolution

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The Journal of Human Evolution , known to all paleoanthropologists as JHE, is the leading technical outlet for articles and news notes in human and nonhuman primate evolution, Paleolithic archaeology, and geochronological, taphonomic, and faunal studies related to those topics. It was founded in 1972 by Brunetto Chiarelli of the Anthropological Institute at the University of Florence, Italy. He produced 14 volumes through the end of 1985, with a wide view of “human evolution” including most of physical anthropology and some allied disciplines. A laxity of the reviewing process among other factors led Academic Press, which actually owned the journal (as does its successor Elsevier), to seek new editors.

Peter Andrews (London) and Eric Delson (New York) reorganized JHE, adding a new Board of Associate Editors with the express plan that they would coordinate review of all standard submissions (other than brief notes and book reviews), combining 3 external reviews with...

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Delson, E., Begun, D.R. (2020). Journal of Human Evolution. In: Smith, C. (eds) Encyclopedia of Global Archaeology. Springer, Cham. https://doi.org/10.1007/978-3-030-30018-0_754

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Evotourism ®

A Smithsonian magazine special report

AT THE SMITHSONIAN

Seven new things we learned about human evolution in 2021.

Paleoanthropologists Briana Pobiner and Ryan McRae reveal some of the year’s best findings in human origins studies

Briana Pobiner and Ryan McRae

This year—2021—has been a year of progress in overcoming the effects of the Covid-19 pandemic on human evolution research. With some research projects around the world back up and running, we wanted to highlight new and exciting discoveries from 13 different countries on five different continents. Human evolution is the study of what links us all together, and we hope you enjoy these stories we picked to show the geographic and cultural diversity of human evolution research, as well as the different types of evidence for human evolution, including fossils, archaeology, genetics, and even footprints!

New Paranthropus robustus fossils from South Africa show microevolution within a single species.

The human fossil record, like any fossil record, is full of gaps and incomplete specimens that make our understanding of complex evolutionary trends difficult. Identifying species and the process by which new species emerge from fossils falls in the realm of macroevolution , or evolution over broad time scales. These trends and changes tend to be more pronounced and easier to identify in the fossil record; think about how different a Tyrannosaurus rex and a saber-toothed cat are from each other. Human evolution only took place over the course of 5 to 8 million years, a much shorter span compared to the roughly 200 million years since dinosaurs and mammals shared a common ancestor. Because of this, smaller-scale evolutionary changes within a single species or lineage over time, called microevolution , is often difficult to detect.

Fossils of one early human species, Paranthropus robustus , are known from multiple cave sites in South Africa. Like other Paranthropus species, P. robustus is defined by large, broad cheeks, massive molars and premolars, and a skull highly adapted for intense chewing. Fossils of P. robustus from Swartkrans cave, just 20 miles west of Johannesburg, are dated to around 1.8 million years ago and show a distinct sagittal crest, or ridge of bone along the top of the skull, with their jaws indicating a more efficient bite force. Newly discovered fossils of P. robustus from Drimolen cave , about 25 miles north of Johannesburg, described by Jesse Martin from La Trobe University and colleagues in January, are at least 200,000 years older (2.04-1.95 million years old) and have a differently positioned sagittal crest and a less efficient bite force, among other small differences. Despite numerous disparities between fossils at the two sites, they much more closely resemble each other than any other known species of hominin. Because of this, researchers kept them as the same species from two different time points in a single lineage . The differences between fossils at the two sites highlight microevolution within this Paranthropus lineage .

Fossil children from Kenya, France, and South Africa tell us how ancient and modern human burial practices changed over time.

Most of the human fossil record includes the remains of adult individuals; that’s likely because larger and thicker adult bones, and bones of larger individuals, are more likely to survive the burial, fossilization, and discovery processes. The fossil record also gets much richer after the practice of intentional human burial began, starting at least 100,000 years ago .

In November, María Martinón-Torres from CENIEH (National Research Center on Human Evolution) in Spain, Nicole Boivin and Michael Petraglia from the Max Planck Institute for the Science of Human History in Germany, and other colleagues announced the oldest known human burial in Africa —a two-and-a-half to three-year-old child from the site of Panga ya Saidi in Kenya. The child, nicknamed “Mtoto” which means “child" in Kiswahili, was deliberately buried in a tightly flexed position about 78,000 years ago, according to luminescence dating. The way the child’s head was positioned indicates possible burial with a perishable support, like a pillow. In December, a team led by University of Colorado, Denver’s Jaime Hodgkins reported the oldest known burial of a female modern human infant in Europe . She was buried in Arma Veirana Cave in Italy 10,000 years ago with an eagle-owl talon, four shell pendants, and more than 60 shell beads with patterns of wear indicating that adults had clearly worn them for a long time beforehand. This evidence indicates her treatment as a full person by the Mesolithic hunter-gatherer group she belonged to. After extracted DNA determined that she was a girl, the team nicknamed her “Neve” which means “snow” in Italian. Aside from our own species, Neanderthals are also known to sometimes purposefully bury their dead . In December, a team led by Antoine Balzeau from the CNRS (the French National Centre for Scientific Research) and Muséum National d’Histoire Naturelle in France and Asier Gómez-Olivencia from the University of the Basque Country in Spain provided both new and re-studied information on the archaeological context of the La Ferrassie 8 Neanderthal skeleton, a two-year-old buried in France about 41,000 years ago. They conclude that this child, who is one of the most recently directly dated Neanderthals (by Carbon-14) and whose partial skeleton was originally excavated in 1970 and 1973, was purposefully buried . There have also been suggestions that a third species, Homo naledi , known from South Africa between about 335,000 and 236,000 years ago, purposefully buried their dead—though without any ritual context. In November, a team led by University of the Witwatersrand’s Lee Berger published two papers with details of skull and tooth fragments of a four to six-year-old Homo naledi child fossil , nicknamed “Leti” after the Setswana word “letimela” meaning “the lost one.” Given the location of the child’s skull found in a very narrow, remote and inaccessible part of the Rising Star cave system, about a half mile from Swartkrans, this first partial skull of a child of Homo naledi yet recovered might support the idea that this species also deliberately disposed of their dead.

The first Europeans had recent Neanderthal relatives, according to genetic evidence from Czechia and Bulgaria.

Modern humans, Homo sapiens , evolved in Africa and eventually made it to every corner of the world. That is not news. However, we are still understanding how and when the earliest human migrations occurred. We also know that our ancestors interacted with other species of humans at the time, including Neanderthals , based on genetic evidence of Neanderthal DNA in modern humans alive today—an average of 1.9 percent in Europeans.

Remains of some of the earliest humans in Europe were described this year by multiple teams, except they were not fully human. All three of the earliest Homo sapiens in Europe exhibit evidence of Neanderthal interbreeding (admixture) in their recent genealogical past. In April, Kay Prüfer and a team from the Max Planck Institute for the Science of Human History described a human skull from Zlatý kůň, Czechia, dating to around 45,000 years old . This skull contains roughly 3.2 percent Neanderthal DNA in the highly variable regions of the genome, comparable with other humans from around that time. Interestingly, some of these regions indicating Neanderthal admixture were not the same as modern humans, and this individual is not directly ancestral to any population of modern humans, meaning they belonged to a population that has no living descendants. Also in April, Mateja Hajdinjak and a team from the Max Planck Institute for Evolutionary Anthropology described three similar genomes from individuals found in Bacho Kiro Cave, Bulgaria, dating between 46,000 and 42,000 years old . These individuals carry 3.8, 3.4, and 3.0 percent Neanderthal DNA, more than the modern human average. Based on the distribution of these sequences, the team concluded that the three individuals each had a Neanderthal ancestor only six or seven generations back. This is roughly the equivalent length of time from the turn of the twentieth century to today. Interestingly, these three genomes represent two distinct populations of humans that occupied the Bulgarian cave—one of which is directly ancestral to east Asian populations and Indigenous Americans, the other of which is directly ancestral to later western Europeans. These findings suggest that there is continuity of human occupation of Eurasia from the earliest known individuals to present day and that mixing with Neanderthals was likely common, even among different Homo sapiens populations.

A warty pig from Indonesia, a kangaroo from Australia, and a conch shell instrument from France all represent different forms of ancient art.

Currently, the world’s oldest representational or figurative art is a cave painting of a Sulawesi warty pig found in Leang Tedongnge, Indonesia, that was dated to at least 45,500 years ago using Uranium series dating—and reported in January by a team led by Adam Brumm and Maxime Aubert from Griffith University. In February, a team led by Damien Finch from the University of Melbourne in Australia worked with the Balanggarra Aboriginal Corporation, which represents the Traditional Owners of the land in the Kimberly region of Australia, to radiocarbon date mud wasp nests from rock shelters in this area. While there is fossil evidence of modern humans in Australia dating back to at least 50,000 years ago , this team determined that the oldest known Australian Aboriginal figurative rock paintings date back to between around 17,000 and 13,000 years ago . The naturalistic rock paintings mainly depict animals and some plants; the oldest example is of a about 6.5 footlong kangaroo painting on the ceiling of a rock shelter dated to around 17,300 years ago. Right around that time, about 18,000 years ago, an ancient human in France cut off the top of a conch shell and trimmed its jagged outer lip smooth so it could be used as the world’s oldest wind instrument . A team led by Carole Fritz and Gilles Tostello from the Université de Toulouse in France reported in February that they re-examined this shell, discovered in Marsoulas Cave in 1931, using CT scanning. In addition to the modifications described above, they found red fingerprint-sized and shaped dots on the internal surface of the shell, made with ochre pigment also used to create art on the walls of the cave. They also found traces of a wax or resin around the broken opening, which they interpreted as traces of an adhesive used to attach a mouthpiece as found in other conch shell instruments.

Fossil finds from China and Israel complicate the landscape of human diversity in the late Pleistocene.

This year a new species was named from fossil material found in northeast China: Homo longi . A team from Hebei University in China including Qiang Ji, Xijun Ni, Qingfeng Shao and colleagues described this new species dating to at least 146,000 years old. The story behind the discovery of this cranium is fascinating! It was hidden in a well from the Japanese occupying forces in the town of Harbin for 80 years and only recently rediscovered. Due to this history, the dating and provenience of the cranium are difficult to ascertain, but the morphology suggests a mosaic of primitive-like features as seen in Homo heidelbergensis , and other more derived features as seen in Homo sapiens and Neanderthals . Although the cranium closely resembles some other east Asian finds such as the Dali cranium , the team named a new species based on the unique suite of features. This newly named species may represent a distinct new lineage, or may potentially be the first cranial evidence of an enigmatic group of recent human relatives—the Denisovans . Adding to the increasingly complex picture of late Pleistocene Homo are finds from Nesher Ramla in Israel dating to 120,000 to 130,000 years old , described in June by Tel Aviv University’s Israel Hershkovitz and colleagues. Like the Homo longi cranium, the parietal bone, mandible and teeth recovered from Nesher Ramla exhibit a mix of primitive and derived features. The parietal and mandible have stronger affiliations with archaic Homo , such as Homo erectus , while all three parts have features linking them to Neanderthals. Declining to name a new species , the team instead suggests that these finds may represent a link between earlier fossils with “Neanderthal-like features” from Qesem Cave and other sites around 400,000 years ago to later occupation by full Neanderthals closer to 70,000 years ago. Regardless of what these finds may come to represent in the form of new species, they tell us that modern-like traits did not evolve simultaneously, and that the landscape of human interaction in the late Pleistocene was more complex than we realize.

The ghosts of modern humans past were found in DNA in dirt from Denisova Cave in Russia.

Denisova Cave in Russia, which has yielded fossil evidence of Denisovans and Neanderthals (and even remains of a 13-year-old girl who was a hybrid with a Neanderthal mother and Denisovan father), is a paleoanthropological gift that keeps on giving! In June, a team led by Elena Zavala and Matthias Meyer from the Max Planck Institute for Evolutionary Anthropology in Germany and Zenobia Jacobs and Richard Roberts from the University of Wollongong in Australia analyzed DNA from 728 sediment samples from Denisova Cave —the largest analysis ever of sediment DNA from a single excavation site. They found ancient DNA from Denisovans and Neanderthals… and modern humans, whose fossils have not been found there, but who were suspected to have lived there based on Upper Paleolithic jewelry typically made by ancient modern humans found in 45,000-year-old layers there. The study also provided more details about the timing and environmental conditions of occupation of the cave by these three hominin species: first Denisovans were there, between 250,000 and 170,000 years ago; then Neanderthals arrived at the end of this time period (during a colder period) and joined the Denisovans, except between 130,000 and 100,000 years ago (during a warmer period) when only Neanderthal DNA was detected. The Denisovans who came back to the cave after 100,000 years ago have different mitochondrial DNA, suggesting they were from a different population. Finally, modern humans arrived at Denisova Cave by 45,000 years ago. Both fossil and genetic evidence point to a landscape of multiple interacting human species in the late Pleistocene, and it seems like Denisova Cave was the place to be!

Fossilized footprints bring to light new interpretations of behavior and migration in Tanzania, the United States and Spain.

Usually when we think of fossils, we think of the mineralized remnants of bone that represent the skeletons of long since passed organisms. Yet trace fossils, such as fossilized footprints, give us direct evidence of organisms at a specific place in a specific time. The Laetoli footprints , for example, represent the earliest undoubted bipedal hominin, Australopithecus afarensis (Lucy’s species) at 3.6 million years ago. In December, a team led by Ellison McNutt from Ohio University announced that their reanalysis of some of the footprints from Site A at Laetoli were not left by a bear, as had been hypothesized, but by a bipedal hominin. Furthermore, because they are so different from the well-known footprints from Site G, they represent a different bipedal species walking within 1 kilometer (0.6 miles) of each other within the span of a few days! Recently uncovered and dated footprints in White Sands National Park , New Mexico, described in September by a team led by Matthew Bennett of Bournemouth University, place modern humans in the area between 23,000 to 21,000 years ago. Hypotheses as to how Indigenous Americans migrated into North America vary in terms of method (ice-free land corridor versus coastal route) as well as timing. Regardless of the means by which people traveled to North America, migration was highly unlikely, if not impossible, during the last glacial maximum (LGM), roughly 26,000 to 20,000 years ago. These footprints place modern humans south of the ice sheet during this period, meaning that they most likely migrated prior to the LGM . This significantly expands the duration of human occupation past the 13,000 years ago supported by Clovis culture and the roughly 20,000 years ago supported by other evidence. Furthermore, it means that humans and megafauna, like giant ground sloths and wooly mammoths, coexisted for longer than previously thought, potentially lending credit to the theory that their extinction was not caused by humans. Also interesting is that most of these footprints were likely made by children and teenagers, potentially pointing to division of labor within a community. Speaking of footprints left by ancient children, a team led by Eduardo Mayoral from Universidad de Huelva reported 87 Neanderthal footprints from the seaside site of Matalascañas in southwestern Spain in March. Dated at about 106,000 years ago, these are now the oldest Neanderthal footprints in Europe, and possibly in the world. The researchers conclude that of the 36 Neanderthals that left these footprints, 11 were children; the group may have been hunting for birds and small animals, fishing, searching for shellfish… or just frolicking on the seashore. Aw.

A version of this article  was originally published  on the PLOS SciComm blog.

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Briana Pobiner | READ MORE

Briana Pobiner is a paleoanthropologist with the National Museum of Natural History’s Human Origins Program . She lead's the program's education and outreach efforts. 

Ryan McRae | READ MORE

Dr. Ryan McRae is a paleoanthropologist studying the hominin fossil record on a macroscopic scale. He currently works for the National Museum of Natural History’s Human Origins Program as a contractor focusing on research, education, and outreach, and is an adjunct assistant professor of anatomy at the George Washington University School of Medicine and Health Sciences.

• ORIGINAL SCIENTIFIC ARTICLE
• Open access
• Published: 29 July 2010

Human Origins Studies: A Historical Perspective

• Tom Gundling 1  

Evolution: Education and Outreach volume  3 ,  pages 314–321 ( 2010 ) Cite this article

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Research into the deep history of the human species is a relatively young science which can be divided into two broad periods. The first spans the century between the publication of Darwin’s Origin and the end of World War II. This period is characterized by the recovery of the first non-modern human fossils and subsequent attempts at reconstructing family trees as visual representations of the transition from ape to human. The second period, from 1945 to the present, is marked by a dramatic upsurge in the quantity of research, with a concomitant increase in specialization. During this time, emphasis shifted from classification of fossil humans to paleoecology in which hominids were seen as parts of complex evolving ecosystems. This shift is in no small part due to the incorporation of neo-Darwinian synthetic theory. Finally, technological innovation and changes in social context are considered as influences on human origins studies.

Introduction

Considering the grand sweep of history, the realization that human beings gradually evolved from some non-human ancestor represents a very recent insight. Even so, the goal of this one brief essay cannot be to provide an in-depth description and analysis of every significant development within the field of paleoanthropology, but rather to identify broad patterns and highlight a collection of “events” that are most germane in shaping current understanding of our evolutionary origin. These events naturally include the accretion of fossil material, the raw data which is the direct, if mute, testimony of the past. These fossil discoveries are situated among technological breakthroughs, theoretical shifts, and changes in the sociocultural context in which human origins studies were conducted. It is only through such a contextualized historical approach that we can truly grasp our current understanding of human origins. Foibles of the past remind us to be critical in assessing newly produced knowledge, yet simultaneously we can genuinely appreciate the enormous strides that have been made.

In addition to selective coverage, a second caveat is that this review will focus on research by scientists writing in English. Non-modern hominids Footnote 1 are a cosmopolitan bunch, having been discovered throughout Africa, Asia, and Europe, and there is a significant literature in other languages. In an effort to ameliorate both of these shortcomings, numerous secondary references are included in the bibliography, providing more in-depth information on specific topics. For example, some texts approach the history of paleoanthropology by detailing a single time period (Bowler 1986 ), early human species (Walker and Shipman 1996 ), or researcher (Morell 1995 ), and there are quite a few that consider the subject more comprehensively (Leakey and Goodall 1969 ; Reader 1988 ; Lewin 1997 ; Tattersall 2008 ). In addition, there are a handful of encyclopedia format tomes (Jones et al. 1994 ; Spencer 1997 ; Delson et al. 2000 ), textbooks (Conroy 2005 ; Cela-Conde and Ayala 2007 ; Klein 2009 ), and “coffee table” popular volumes (Stringer and Andrews 2005 ; Johanson and Edgar 2006 ) that in part address the history of human origins studies. Moreover, these texts contain abundant references to the primary literature if that level of scrutiny is desired.

In seeking to provide a useful heuristic framework for the purposes of this particular essay, human origins studies can be broken down into two very broad periods. The first is roughly the century between 1850 and 1950 when research, often conducted by individuals with training outside of anthropology, focused on taxonomy and phylogeny. In other words, although scientists were cognizant that climate change (e.g., northern hemisphere glaciations) would have directly impacted the evolution of early humans, they were mainly interested in collecting “missing links,” naming them, and creating family trees. The second period, from 1950 to the present, is characterized by the relatively rapid development of paleoanthropology as it is currently practiced. Here the emphasis is only partly on the hominids themselves, with ecological context being of equal importance.

The Emergence of Human Origins Studies

This review begins with two mid-nineteenth century developments which are often conflated, but were initially distinct. The first is the acceptance of a temporal association of human material culture (stone tools), with extinct Ice Age mammals (Van Riper 1993 ; Sommer 2007 ). This was significant in that it opened up a considerable prehistory for the human species, well beyond estimates derived from literal scriptural interpretation. However, while acknowledging a lengthy antiquity for the human species, there was, at the time, no reason to suspect that the makers of the stone tools were not fully modern humans in a biological sense. The second major development was the publication of Darwin’s On the Origin of Species in 1859 (Darwin 1859 ). Darwin is rightfully credited with being the most influential, although by no means the first individual to broach the subject of descent with modification, or transmutation theory, as he put it (for an overview of pre-evolutionary ideas related to human origins, see Greene 1959 , Bowler 2003 ). Darwin’s central thesis was that all living species shared a common ancestry, with “endless forms most beautiful” having diverged via natural selection, and although he only briefly mentioned his own species the inference was clear. These two events dovetailed into the now quotidian, but then controversial, notion that humans had evolved over a vast expanse of time (Grayson 1983 ).

While Darwin was initially reticent to discuss human evolution in any detail, his colleague Thomas H. Huxley harbored no such reluctance when he published Man’s Place in Nature: Essays in 1863 (Huxley 1900 ). Darwin freely admitted that the veracity of his audacious proposal would have to withstand paleontological scrutiny and that his theory would collapse in the absence of transitional fossil forms. Huxley’s advantage, beyond his more outspoken personality, was that he actually had a fossil human to describe. The first Neandertal recognized by science was discovered in 1856; however, its description only appeared in English three years later, just as Darwin was going to press (Trinkaus and Shipman 1993 ). Huxley provided a detailed description of the eponymous cranium coupled with carefully composed line drawings (Huxley 1900 ). However, while the importance of the Neandertals in providing empirical evidence documenting an ancient and morphologically distinct human form cannot be discounted, these people hardly bridged the gap separating humans and the great apes. Although a few dissenting voices denied the close evolutionary relationship among humans and the “man-like” apes, and consequently an ape phase of human ancestry, most scientists accepted the overwhelming morphological and embryological evidence in support of just such a relationship. This acceptance was in no small part due to Huxley’s meticulous comparison of gorilla and human anatomy in which he concluded that the gorilla and its close relation, the chimpanzee, represented the nearest approach to humanity in nature.

If Neandertals were more or less human, then more distant, primitive “missing links” remained to be discovered. Just such fossils were recovered on the island of Java in the 1890s by Dutch physician Eugene Dubois, who had traveled to Indonesia as part of the army but with the express purpose of finding the remains of primitive humans (Shipman 2001 ). Java Man consisted of a skull cap, a femur, and a few isolated teeth which taken in combination suggested an early human with a much smaller cranial capacity relative to Neandertals or Homo sapiens (roughly 1,000 vs. 1,500 cubic centimeters), although the femur appeared modern. Dubois did not receive the universal accolades and acceptance he coveted, but his fossils bolstered the conventional wisdom at the time that humans first evolved somewhere in Asia.

During the early twentieth century, the early hominid fossil record grew significantly, if not exponentially, and evolution was widely accepted in scientific circles even while large segments of the lay public remained skeptical. Certainly, there were disagreements over whether natural selection was a sufficient evolutionary mechanism in itself (Bowler 1983 ), but the basic premise of biological change through time was affirmed. The recovery of additional Neandertal remains in Europe refuted lingering claims of pathology regarding the original Neander Valley specimen and solidified the interpretation that the latter was representative of a population of archaic humans occupying Ice Age Europe. Some Neandertal remains were interpreted as not only indicating intentional internment but also associated funerary ritual. The European fossil record was extended significantly with the recovery of a robust lower jaw from Mauer, Germany discovered in 1907.

In 1912 in England, heretofore devoid of non-modern hominid remains despite the prominence of several British scholars in human origins studies, the announcement of hominid fossils from Piltdown was warmly received locally, if with some incredulity abroad. Piltdown was significant since it reified the “brain first” hypothesis, in which primitive humans evolved a large brain before other key human traits evolved. Although a favorite of intelligent design creationism advocates, Piltdown is actually a beautiful example of the scientific method at work, whereby new evidence eventually calls into question prior interpretation, and in this case recognition of intentional fraud (Spencer 1990 ). It was, after all, a new relative dating method measuring the fluorine content of fossils that in 1953 exposed the non-contemporaneity of the jaw and skull. In any case, in the first decades of the twentieth century, a fairly simple human family tree was beginning to emerge (see McCown and Kennedy 1972 and especially Delisle 2007 for exceptions). Relatively small-brained Pithecanthropus led to the more capacious Neandertals and Piltdown, who in turn evolved into modern H. sapiens . Yet the truly ape-like human ancestors remained elusive.

Africa as the Cradle of Humanity

In 1921 a skull bearing superficial resemblance to European Neandertals was recovered as part of mining operations at a place called Broken Hill in Northern Rhodesia (now Zambia). Rhodesian Man marks the recovery of the first in a very long line of non-modern hominids from the African continent. A mere four years later, University of Witwatersrand anatomist Raymond Dart, Australian by birth and having been trained in England, published a brief paper describing the fossil skull of a juvenile “ape” discovered in a limestone quarry near Taung, South Africa. Dart identified certain features of the face, the teeth, the cranium, and the brain of Australopithecus africanus that foreshadowed those of H. sapiens and made the startling claim that what was essentially a bipedal ape signaled the beginning of the human lineage separate from the African great apes.

Initially, with only the one individual, and a juvenile at that, Dart found little support. His most ardent advocate, Scottish physician and paleontologist Robert Broom discovered additional fragmentary remains of the australopithecines, as they were then called, in other South African caves in the 1930s, but these were initially insufficient to sway opinion (Dart 1959 ). This was perhaps due to the near-simultaneous discovery of significant hominid remains from Zhoukoutien (Dragon Bone Hill) in China which quickly eclipsed whatever controversy the diminutive skull from Taung elicited, and despite Broom’s ongoing efforts. As was the case with Java Man, the more complete Chinese fossils fulfilled the expectations of many scientists who anticipated that earliest human ancestors evolved to the East. Comparative analysis of the Javanese and Chinese fossils revealed a great deal of similarity, and all of the fossils were ultimately subsumed in the species Homo erectus .

The Neo-Darwinian Synthesis and the New Physical Anthropology

For several disparate reasons, the decades following the end of World War II (WWII) rather quickly led to a science of paleoanthropology that is recognizably modern. One significant factor relevant in the U.S. if not everywhere, was the dramatic upsurge in enrollment at colleges and universities. The G.I. Bill and subsequent effects of Civil Rights legislation that greatly increased access to higher education meant that millions more students went to college and hence the expansion of existing campuses and programs and in some cases the appearance of entirely new colleges and universities Footnote 2 . As a result, greater numbers of faculty were required who could teach courses and supervise research in diverse academic programs, which in turn led to an attendant rise in the numbers of graduate students themselves who went on to secure positions at institutions of higher learning. Consequently, many disciplines experienced significant increases in research activity, including physical anthropology, and it is worth noting that this was the first generation of researchers whose formal training was in physical anthropology, not in some allied field such as anatomy or medicine. The dramatic rise in practitioners not only increased the knowledge base in terms of simple quantity, but specialization within the field also began to emerge.

A second crucial development that transformed human evolutionary studies was theoretical in nature. Changing ideas regarding the process of evolution had been fermenting and roiling in biology circles for several decades before they infiltrated the study of human origins. In essence, a consensus was reached among biologists ( sensu lato ) that Darwinian natural selection acting on variation arising from random mutation was a sufficient mechanism to explain evolutionary change. For anthropologists, although questions of taxonomy and phylogeny remained important, the intellectual fallout of the so-called neo-Darwinian synthesis led to the “New Physical Anthropology” in which early hominid fossils, rather than representative of some platonic archetype, were interpreted as unique members of variable populations. Focusing on evolution as a process effecting change in populations over time, in contrast to the comparatively myopic sorting of the resulting pattern , arguably represents the most significant theoretical shift in thinking about evolution since Darwin.

Given the comparative de-emphasis on iconic types, the bloated alpha taxonomy of the past was reduced to a mere handful of hominid species displaying previously under-appreciated within species variability. This great reduction in hominid names and consequent simplification of hominid family trees has led some modern scholars to lament what they see as a return to the bad old days of teleology and orthogenesis. Yet there can be little doubt that the “splitting” taxonomic philosophy of the past where almost every new specimen received a new species or quite frequently a new genus name was in dire need of revision.

Just as species types came under scrutiny, so did the concept of evolutionary grades which had up to this point made clear distinctions between the categories of ape and human. While this may have provided some welcome taxonomic clarity, it was artificial in that it ignored the evolutionary reality that at some point members of the human lineage were very ape-like. This realization, obvious in retrospect, led to the widespread acceptance of the South African australopithecines as human ancestors, and the important corollary that bipedalism preceded other distinctive human attributes (Gundling 2005 ).

In addition to increased research activity and theoretical shifts, by the early 1960s technological innovations for the first time permitted the creation of a reliable absolute timescale of human evolution. Comparative protein analysis demonstrated that the African apes were most similar genetically to H. sapiens , inferring their recent common ancestry to the exclusion of other apes and monkeys. Molecular clocks based upon mutation rates and calibrated by the fossil record suggested that this common ancestor lived as little as a few million years ago, although recent estimates put this ancestor at seven to five million years ago. Consequently, known early and middle Miocene ape species became suspect as purported human ancestors, since they preceded the split between the hominid and great ape lineages. Most notably this eventually led to the downfall of Ramapithecus , a Miocene ape genus once widely hailed as a very ancient and very primitive hominid Lewin ( 1997 ).

While molecular studies of living species effectively imposed a theoretical maximum on the age of the hominid lineage, the temporal framework of human origins was further clarified with the introduction of the new potassium argon (K-Ar) method of absolute dating. Louis and Mary Leakey had been scouring the fossil-bearing sediments in and around eastern Africa’s Great Rift Valley for decades when Mary discovered the skull of a robust australopithecine at Olduvai Gorge in 1959. Significantly, Zinjanthropus , the genus coined for the new skull, was discovered within sediments near the base of the Pleistocene Epoch. Volcanic minerals from associated strata were dated to approximately 1.75 million years ago using the K–Ar method, nearly double the age estimated from using other more crude means. This greatly expanded time range certainly bolstered claims for the australopithecines as human ancestors rather than extinct collateral cousins to the “true” human lineage, yet to be discovered.

As an aside, Louis Leakey’s interest in understanding the human past was not limited to the collection of fossils. Sherwood Washburn, a main architect of the new physical anthropology, along with Irven DeVore, conducted pioneering studies of savanna baboons, large-bodied, terrestrial, and highly social primates that served as living proxies for modeling early hominid behavioral ecology (Washburn and DeVore 1961 ). Leakey, on the other hand, took a more phylogenetically based approach and hired scholars to conduct research into the behavior of the great apes as a potential new data source informing hypotheses of early hominid behavior. Jane Goodall was the first, studying chimpanzee behavior at Gombe in Tanzania, then came Dianne Fossey who undertook a longitudinal study of mountain gorillas in Rwanda, and finally Birute Galdikas traveled to Indonesia to conduct field studies of the orangutan (see Kinzey 1987 and De Waal 2001 for more recent primate studies that explicitly address questions of human behavioral evolution).

The emergence of paleoanthropology as a truly multidisciplinary endeavor, concerned with a more holistic picture of our evolutionary past, was a logical extension of the post-WWII new physical anthropology which eschewed simple classification and promoted variable populations as the unit of study. Naturally, these hominid populations did not exist in a vacuum but were components of complex, evolving ecosystems. Hence, field work began to emphasize the collection of greater contextual data in an effort to reconstruct biological and physical environments in which these human ancestors existed and evolved. One of the first field projects to adopt this new approach was an international expedition centered around the Omo River Valley in southern Ethiopia, beginning in 1967. Remarkably, of the 50 papers collected in the resulting volume, only five primarily focus on the hominid remains themselves (Coppens et al. 1976 ).

Early Human Diet and Subsistence

One major aspect of early hominid ecology that occupied researchers engaged in such multidisciplinary efforts was subsistence, which has understandably been of great interest to paleoanthropologists, particularly after 1950 as scientists endeavored to contextualize the fossil remains of distant ancestors. What early humans ate, how food was acquired and processed, even how it was distributed among members of a social group, became viable questions. Throughout the 1950s and 1960s it was widely assumed that the social, cognitive, and technological skills associated with big-game hunting drove the evolution of the human species; in fact the allure of “Man the Hunter” is longstanding in Western thought (Cartmill 1993 ). Raymond Dart, as part of his second foray into human origins studies, proposed that Australopithecus had already developed a hunting strategy facilitated by a technology comprised of durable animal parts that he referred to as the osteodontokeratic (bone, tooth, horn) culture. This concept was enthusiastically embraced by writer Robert Ardrey, who published a series of four popular novels documenting the success of these “killer apes” in the context of a changing environment (e.g., Ardrey 1976 ). Research scientists were only slightly less enthusiastic in championing such ideas (Lee and DeVore 1968 ) which remain popular, if more nuanced today (Wrangham and Peterson 1996 ).

Mirroring changes in the broader society, by the early 1970s some anthropologists challenged the “Man the Hunter” hypothesis and developed an alternative that focused on the central role of women in child rearing and gathering of food resources (Dahlberg 1981 ). These studies used ethnographic data from extant food-foraging societies, the rarity of which injected a sense of urgency on the part of anthropologists. Not long after the “Women the Gatherer” model appeared as a second wave feminist rejoinder to the previously unquestioned authority of “Man the Hunter,” another group of researchers also began to question the big-game hunting scenario. Archeologists, geologists, and paleontologists began working on “site formation processes” to get a better understanding of how assemblages of fragmented animal bones and stone tools came to be commingled. Over the next few decades, often with recourse to modern ecosystems as analogs, one of the main conclusions drawn from the new science called taphonomy (=laws of burial) was the potential importance of scavenging. The association of “bones and stones” was no longer assumed to be the signature of hominid big-game hunting but instead interpreted as meals containing essential fat and protein scavenged by early humans. Perhaps even more disconcerting, some sites were reinterpreted as the remains of carnivore kills occasionally including early humans themselves (Brain 1981 ; Hart and Sussman 2008 ).

Here’s Lucy

If Mary and Louis Leakey’s discoveries at Olduvai put the Great Rift Valley on the map, during the 1970s eastern Africa was validated as the center of early hominid studies. The Leakey’s son Richard established himself on the east side of Lake Turkana in northern Kenya, where his expeditions uncovered a prolific cache of early hominid fossils, some of which corroborated the occasionally controversial claims made by his parents a decade earlier. Sediments around the lake yielded hominid fossils of robust australopithecines, early members of genus Homo , and an early African variant of Asian H. erectus , these days referred to as Homo ergaster (Leakey and Lewin 1978 ). The latter includes a mostly complete skeleton, KNM-WT15000, which has become iconic for the species (Walker and Shipman 1996 ).

Arguably the most significant fossil discovery of the 1970s was another partial skeleton, AL-288, from Hadar, Ethiopia, better known as Lucy (Johanson and Edey 1981 ). Here was a single individual represented by numerous skeletal elements, and although her morphology was generally similar to the “gracile” australopithecines of South Africa, she was even more primitive in some respects. Consequently her discoverers coined a new species name, Australopithecus afarensis that included not only the Hadar specimens but fossils collected by Mary Leakey’s expedition at Laetoli in Tanzania. The latter is renowned for its famous footprint trail preserved in solidified volcanic ash, imparting convincing evidence for bipedalism at 3.6 million years ago. Hadar is also replete with datable volcanic sediments, and Lucy’s status as the most primitive hominid was reinforced by firm radiometric dates which placed the fossils at greater than 3.0 million years ago, at the time astonishingly ancient.

One other significant event from the 1970s bears mentioning. Although the American Journal of Physical Anthropology was first published in 1918, it is perhaps surprising that a journal explicitly dedicated to the study of human evolution did not appear in the U.S. until 1972. Since then the Journal of Human Evolution has been the premier academic forum for publications related to human evolution, and in 1992, the Paleoanthropology Society was established, which organizes its own conference and publishes an online journal.

Modern Human Origins

The question of modern human origins has been debated for centuries, long predating paleoanthropology as a scientific discipline. One of the central issues, which became particularly evident as Renaissance and Enlightenment Europeans began to travel the globe on a regular basis, was how to explain the physical diversity of human populations. Two broad perspectives emerged, one which viewed all people as having a single origin and another which believed that supposedly distinct races had separate origins. The pre-Darwinian debates between so-called monogenists and polygenists were recast with the advent of an evolutionary paradigm in the mid-nineteenth century. Within this new theoretical context, monogenists believed that all living humans evolved from a common ancestor that was already H. sapiens , while the polygenists believed that the races had deeper roots and had descended from different non-modern ancestors (e.g., H. erectus or in a few instances different ape species). A major step towards resolving this debate came in 1987 with an analysis of living human mitochondrial DNA diversity which concluded that H. sapiens had a recent African origin. The discovery of essentially modern human fossils at the 160,000-year-old site of Herto, Ehtiopia, provides paleontological support for a recent African origin, and many subsequent genetic studies have supported this basic conclusion. However, the possibility of some gene flow between migrating early modern humans and local archaic populations remains plausible (compare Stringer and McKie 1996 and Wolpoff and Caspari 1998 , also see Relethford 2003 for a geneticist’s perspective).

Conclusion: Twenty-First Century Paleoanthropology

New fossil discoveries, technological innovations, theoretical advances, and social transformations will continue to inform knowledge of our deep past. Recovery of hominid fossils, some from previously unknown time periods and geographic locations, continues at a brisk rate. Many of the most significant recent discoveries are beginning to fill in the crucial African late Miocene time period during which our lineage ramified from that leading to the chimpanzee (Gibbons 2006 ). Of particular note, one of these fossils was discovered in Chad, quite a distance from established sites in the Great Rift Valley, challenging the long standing hypothesis that hominids evolved in the savanna grasslands of eastern Africa while the African ape ancestors remained sequestered in their tropical rainforest refugium. Moreover, botanical, faunal, and geological evidence associated with very early fossil hominids in Ethiopia and Kenya intimate a forested environment, a discovery that clearly constrains hypotheses explaining the success of the bipedal adaptation.

Other significant fossil discoveries from the early Pleistocene site of Dmanisi in the Republic of Georgia have energized discussion of the initial expansion of early humans beyond the tropics of Africa (Wong 2006 ). Not only are these fossils considerably older than prior known Eurasian specimens, but they are morphologically primitive, especially in terms of stature and cranial capacity, and are associated with very simple (“mode 1”) lithic technology. These early migrants hardly manifest the tall striding bipeds equipped with comparatively advanced Acheulian bifacial tools so often depicted in earlier “out of Africa” scenarios Footnote 3 , which are at least in part based on the iconic WT15000 skeleton mentioned earlier.

Perhaps the most surprising discovery of the last decade is the diminutive 18,000-year-old skeleton from the Indonesian island of Flores, which has sparked a spirited, occasionally acrimonious debate between those advocates of a replacement model of modern human origins and those inclined towards regional continuity (Morwood and van Oosterzee 2007 ). The former, comprised of the team who made the discovery and their allies, interpret the remains as those of a surprisingly primitive hominid akin to early Homo , and perhaps the first documented example of the effects of island dwarfing on an early human population. Other scholars believe the remains to be those of a pathological modern human, whose illness resulted in a cascade of skeletal and dental anomalies. Ongoing research on Flores and other nearby locations will undoubtedly resolve this debate.

New discoveries are not limited to the paleontological record but also include behavioral information gleaned from archaeology. Symbolic expression in the form of language, art (including music), and religion is undoubtedly one of the most distinctive human traits. Evidence for such behavior has proved elusive beyond the seeming cultural explosion perceived in the Upper Paleolithic of Europe beginning around 35,000 years before present. However, archeological evidence for at least some of these behaviors has recently been coaxed out of several sites in sub-Saharan Africa. Advanced utilitarian objects such as blades and harpoons have been recovered well back into the Middle Stone Age and use of ochre and shells for body adornment has been found at sites approaching 100 kiloannum (Balter 2009 ).

Recent advances also include a plethora of technological innovations that have allowed anthropologists to hone traditional inquiries in the areas of dating (e.g., single crystal, laser fusion, argon–argon dating), systematic analysis (e.g., geometric morphometrics), and paleoenvironmental reconstruction (e.g., stable isotope analysis). The badly distorted remains of the spectacular 4.4 megaanum skeleton of Ardipithecus ramidus from Aramis, Ethiopia was restored in part using digital imaging technology (Gibbons 2009 ). Additionally, new technology is facilitating, perhaps even driving, novel questions such as those related to the emergence of the unique human life history pattern.

While fossils provide real-time evidence for human evolution, signals from our ancient past are also encoded into our modern DNA. The groundbreaking work of the 1960s effectively demonstrated our close affinity with the African great apes, and today’s genomic analyses comparing humans and chimpanzees are beginning to reveal differences in much finer detail than heretofore possible. Already several areas within the human genome have been identified as having undergone intense selection; these regions may be related to the evolution of the especially dexterous human thumb, reduction of muscles of mastication in the wake of the ability to cook food, the greatly enlarged neo-cortex, and our ability for spoken language.

In addition to modern DNA analyses, ancient DNA analysis has informed the “Neandertal problem” providing preliminary evidence in support of the replacement hypothesis, at least in Europe, whereby modern humans arriving there equipped with Upper Paleolithic technology drove the indigenous Neandertals to extinction. Even more recent genomic analyses, however, suggest that a small but detectable degree of interbreeding occurred when expanding modern human populations emerging from the African tropics encountered Neandertal populations in the Middle East around 120,000 years before present (Gibbons 2010 ).

In conclusion, our understanding of human origins, like all scientific knowledge, is the result of an ongoing, iterative process. Over the last few decades, the accelerating pace of fossil discoveries and the incorporation of innovative technologies have corroborated and enhanced much of what we already suspected to be true, although there have been a few surprises. No doubt this pattern will continue into the foreseeable future as we slowly, yet inexorably, piece together the circumstances by which our lineage became human.

Hominidae (=hominid) is the biological group (clade) to which humans and their extinct ancestors belong. For many current scholars, this group is distinguished at some lower taxonomic level, usually the tribe Hominini (=hominin). In this study, I maintain the traditional use of Hominidae simply to be consistent with the historical literature. For the same reason, I use the subfamily designation Australopithecinae (=australopithecine) for all of the African “bipedal apes.” This group is certainly paraphyletic, to use the modern jargon, and as a result an increasing number of scholars prefer to use the less formal term australopith to lump together the various African species.

According to the U.S. Census Bureau, there were about 2.2 million students enrolled in U.S. colleges in 1950. That number doubled by 1963 to just less than 4.4 million and doubled again to over 9 million by 1972.

I usually avoid this term despite its heavy usage within the scientific community. I believe that the “Out of Africa” trope perpetuates an anti-Africa bias which seems to suggest that early humans, on several occasions, left Africa wholesale as if there was something inherently undesirable about the place. I suppose that “hominid extra-tropical range expansion” doesn’t have the same ring, but it is more accurate.

Ardrey R. The hunting hypothesis. New York: Atheneum; 1976.

Google Scholar  

Balter M. On the origin of art and symbolism. Science. 2009;323:709–11.

Article   CAS   PubMed   Google Scholar  

Bowler PJ. The eclipse of darwinism. Anti-darwinian evolution theories in the decades around 1900. Baltimore: The Johns Hopkins University Press; 1983.

Bowler PJ. Theories of human evolution: a century of debate 1844–1944. Baltimore: The Johns Hopkins University Press; 1986.

Bowler PJ. Evolution: the history of an idea. 3rd ed. Berkeley: University of California Press; 2003.

Brain CK. The hunters or the hunted? An introduction to African cave taphonomy. Chicago: University of Chicago Press; 1981.

Cartmill M. A view to a death in the morning: hunting and nature through history. Cambridge: Harvard University Press; 1993.

Cela-Conde CJ, Ayala FJ. Human evolution: trails from the past. Oxford: Oxford University Press; 2007.

Conroy GC. Reconstructing human origins. 2nd ed. New York: W.W. Norton; 2005.

Coppens Y, Howell FC, Isaac GL, Leakey REF. Earliest Man and environments in the lake rudolf basin: stratigraphy, paleoecology and evolution. University of Chicago Press; 1976.

Dahlberg F, editor. Woman the gatherer. New Haven: Yale University Press; 1981.

Dart RA. Adventures with the missing link. Britain: Hamish Hamilton, Ltd.; 1959.

Darwin CR. On the origin of species by means of natural selection, or the preservation of favoured races in the struggle for life. London: John Murray; 1859.

Book   Google Scholar  

De Waal FB, editor. Tree of origin: what primate behavior can tell us about human social evolution. Cambridge: Harvard University Press; 2001.

Delisle RG. Debating humankind’s place in nature 1860–2000: the nature of paleoanthropology. Upper Saddle River: Pearson Education, Inc.; 2007.

Delson E, Tattersall I, Van Couvering JA, Brooks AS. Encyclopedia of human evolution and prehistory. 2nd ed. New York: Garland Publishing, Inc.; 2000.

Gibbons A. The first human: the race to discover our earliest ancestors. New York: Doubleday; 2006.

Gibbons A. Breakthrough of the year: Ardipithecus ramidus . Science. 2009;326:1598–9.

Gibbons A. Close encounters of the prehistoric kind. Science. 2010;328:680–4.

Grayson DK. The establishment of human antiquity. New York: Academic; 1983.

Greene JC. The death of Adam: evolution and its impact on western thought. Ames: Iowa State University Press; 1959.

Gundling T. First in line: tracing our ape ancestry. New Haven: Yale University Press; 2005.

Hart D, Sussman RW. Man the hunted: primates, predators, and human evolution. Expandedth ed. Boulder: Westview Press; 2008.

Huxley TH. Man’s place in nature: essays. New York: D. Appleton & Co.; 1900 [1863]

Johanson D, Edey M. Lucy: the beginnings of humankind. New York: Simon and Schuster; 1981.

Johanson DJ, Edgar B. From Lucy to language. Revised, updated, and expanded. New York: Simon & Schuster; 2006.

Jones S, Martin R, Pilbeam D, editors. The cambridge encyclopedia of human evolution. Cambridge: Cambridge University Press; 1994.

Kinzey WG, editor. The evolution of human behavior: primate models. Albany: SUNY Press; 1987.

Klein RG. The human career: human biological and cultural origins. 3rd ed. Chicago: University of Chicago Press; 2009.

Leakey LSB, Goodall VM. Unveiling man’s origins. Cambridge: Schenkman Publ. Co.; 1969.

Leakey RE, Lewin R. People of the lake: mankind and its beginnings. New York: Doubleday; 1978.

Lee RB, DeVore I, editors. Man the hunter. Chicago: Aldine Publishing Co.; 1968.

Lewin RL. Bones of contention: controversies in the search for human origins. 2nd ed. Chicago: University of Chicago Press; 1997.

McCown TD, Kennedy KAR, editors. Climbing man’s family tree: a collection of major writings on human phylogeny, 1699 to 1971. Englewood Cliffs: Prentice Hall; 1972.

Morell V. Ancestral passions: the Leakey family and the quest for humankind’s beginnings. New York: Simon & Schuster; 1995.

Morwood MJ, van Oosterzee P. A new human: the startling discovery and the strange story of the “Hobbits” of Flores, Indonesia. New York: Harper Collins; 2007.

Reader J. Missing Links: the hunt for earliest man. 2nd ed. London: Pelican Books; 1988.

Relethford JH. Reflections of our past: how human history is revealed in our genes. Boulder: Westview Press; 2003.

Shipman P. The man who found the missing link: Eugene Dubois and his lifelong quest to prove Darwin right. New York: Simon and Schuster; 2001.

Sommer M. Bones and ochre: the curious afterlife of the red lady of paviland. Cambridge: Harvard University Press; 2007.

Spencer F. Piltdown: a scientific forgery. London: British Museum (Natural History); 1990.

Spencer F, editor. History of physical anthropology: an encyclopedia. New York: Garland Publishing, Inc.; 1997.

Stringer C, Andrews P. The complete world of human evolution. London: Thames and Hudson; 2005.

Stringer C, McKie R. African exodus: the origins of modern humanity. New York: Henry Holt; 1996.

Tattersall I. The fossil trail: how we know what we think we know about human evolution. 2nd ed. Oxford: Oxford University Press; 2008.

Trinkaus E, Shipman P. The Neandertals: changing the image of mankind. New York: Alfred A. Knopf; 1993.

Van Riper AB. Men among mammoths: victorian science and the discovery of human prehistory. Chicago: Chicago University Press; 1993.

Walker A, Shipman P. The wisdom of the bones: in search of human origins. New York: Alfred A. Knopf; 1996.

Washburn SL, DeVore I. The social life of baboons. Sci. Am. June; 1961

Wolpoff M, Caspari R. Race and human evolution. Boulder: Westview Press; 1998 [1997]

Wong K. Stranger in a new land. Sci Am. 2006;16(2):38–47.

Article   Google Scholar  

Wrangham R, Peterson D. Demonic males: apes and the origins of human violence. New York: Houghton Mifflin; 1996.

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RETRACTED ARTICLE: A brief history of human evolution: challenging Darwin’s claim

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There is a consensus among evolutionists today that man first appeared in Africa approximately four million years ago. Others counter this theory saying, “... when shall we speak of man as man”? The timeline they give is approximately one million years and to fully understand one million years is still a difficult task.

However, another even better way to understand time and man is to study it in terms of generations. So, keeping in mind that primitive people married and had children early, twenty years will make an average generation. According to this there would be 50,000 generations in a million years. Keeping this in mind if we calculate generations we find that 250 generations back take us to the time when written history began. While, another 250 generations back would take us to the time (10,000 years ago), when cultivation began, and man started settled life. Now we are left with 49,500 generations of men, plus a time span of 990,000 years. Keeping these statistics in mind let us ask the question once more, when should we speak of man as man?

Therefore, this paper attempts not only to understand the timeframe “when we can really call Man? – Man” in light of the so-called history of human evolution but also to understand that if the specie roaming the earth for a million years was truly man’s ancestor, as is claimed by Charles Darwin. Then what took man’s ancestor so long to show signs of development that we only witness in the last 12000 years.

Moreover, while keeping man’s progress under consideration of the last 12000 years, it will further shed light on why there are serious reservations about Charles Darwin theory of human evolution. As many scientists, evolutionists, archeologist and different religious scriptures strongly claim that man came to the earth fully developed and did not evolve from a lesser specie.

Introduction

Prehistory simply means the time before written history began. More than 99% of man’s story is prehistory. The consensus by the historians is that man is about 1 million years old, but he did not write anything, until 5000 years ago. Although, prehistoric man did not leave us any written records he unintentionally left us information on his way of life, which is interpreted by different kinds of scientists. These scientists are specialist in physical anthropologists, human paleontologists, archaeologist and other fields. It is their job to find out what happened before written history began (Braidwood 1964a , b ). Some historians believe that it is their duty to bring together various facts given by these scientists and put them together in such a way as to reach objective conclusions. E. H. Carr has two ideas on what an objective historian is: one is that he “... has the capacity to rise above the limited vision of his own situation in society and in history – a capacity which... is partly dependent on his capacity to recognize the extent of his involvement in that situation, to recognize, that is to say, the impossibility of total objectivity”, and his other idea is “...simply that he gets his facts right, but rather that he chooses the right facts, or, in other words, that he applies the right standard of significance” (Marwick 1970 ).

Man has been blessed with the best attributes and qualities among all living species. It is one, specie that can control all other species with its intelligence and wisdom. It is the one specie that has the ability to reason and think rationally. C. M. Bowra says, “based on the humanistic faith that man is worth studying for his own sake”, that “Human beings are the element in our environment which is of most consequence to every child of man” (Grant 1965 , 1952 ). Therefore we should not be a surprised when we see many theories and counter theories with regard to the dates of the origin, the evolution or the different phases of this development.

An enormous span of human history needs to be covered in order to have answers to these questions. Robert J. Braidwood, in the preface to his book, Prehistoric Men , states,

“New discoveries and new techniques for the interpretation of the evidence of mankind’s past appear almost daily. The newer finds and techniques necessitate reconsideration of older evidence. Slowly but surely we move toward fuller understandings of those beings whose history holds the greatest fascination for all of mankind-men themselves.”

Whatever, material I have put together in this paper and whatever hypothesis, assumptions or conclusions, I shall make will be based on the work done up until this time. What the future discoveries behold is still unknown and mysterious to all of us.

There are many different ways and views about how to understand prehistory and history. Pre-historians have given two main ways to accomplish this task. One is either to go down the ladder of time, whereas the other is to climb up the ladder. I will follow the first path for this paper.

There is a general consensus among evolutionists today that man first appeared in Africa approximately 4 million years ago. Others counter this theory saying, “... when shall we speak of man as man”? The timeline they give is approximately 1 million years. Even to fully understand 1 million years is very difficult task. For example, if we compare this whole time period to 1 day, it would be something like this:

“The present time is midnight, and Jesus was born just two minutes and fifty seconds ago. Earliest history began about seven minutes ago. Everything before 11:53 P.M. was in prehistoric time” (Braidwood 1964a , 1967b : 10–11).

Another, even better way to understand time is to study it in terms of generations. So, keeping in mind that primitive people married and had children early, 20 years will make an average generation. According to this there would be 50,000 generations in a million years. Keeping this in mind if we calculate generations we find that 250 generations back take us to the time when written history began,

“...David was king of Israel less than 150 generations ago, Julius Caesar was alive just 100 generations ago, Columbus was there 25 generations ago and the United States is just 10 generations old. So, the current scenario is that there were 49,750 generations of men before written history began!” (Braidwood 1964a , 1967b )

And another 250 generations back would take us to the time (10,000 years ago), when cultivation began and man started settled life. Now we are left with 49,500 generations of men, plus a time span of 990,000 years. Keeping these statistics in mind let us ask the question once more, when should we speak of man as man? Can the human mind accept this fact that for such a long period of time and for so many generations, man did not make any effort to change either his surrounding or his life style? Why was he only a hunter and a gatherer for thousands and thousands of years? Why did it take him such a long time to change himself and his way of life? Or would it not be wrong to say here that it was not really man who was roaming the earth at that time, but some other species, with fewer qualities than what man has been blessed with. Some human evolutionists have tried to call these species man’s ancestors and they believe that man eventually evolved out of these species, whereas others have refuted this concept. According to them, man did not evolve out of any other species: man was born a man with the best of qualities among all living species.

Results and discussion

Let us briefly review human evolution, in order to understand this concept and to try to find answers to questions that are still confusing us today. Let us begin with the evolution of animals. Zoologists have classified the members of the animal kingdom according to their differences and similarities. We humans fall under this kingdom because we move and eat with our mouth; we are vertebrates because of our backbone, and we are mammals because we are warm-blooded and we breast-feed our offspring. We are primates because we have grasping hands, flexible limbs, and a highly developed sense of vision. We are also members of the family Hominoidea , the taxonomic group which includes both humans and apes, because of the absence of a tail, swinging arms, and the shape of our teeth. The term hominoid refers to all present and past apes and humans, while hominid refers specifically to present and past humans (Price and Feinman 1993 , 1997a , 1997b , c , d , e , f ). Hominids are known as walking creatures with comparatively large brains; humans today are the sole living representation of this group. According to some evolutionists, fossil bones and genetic studies indicate that the hominids shared an ancestor with the great African ape. It is at this point that the record becomes more complex, when the study of primate evolution turns into the study of hominid evolution. According to Richard Klein of Stanford University, determining the genus and species of the fossil bones of early humans is a very difficult task. The fragmentary pieces of the early fossil finds represent only a few hundred separate individuals. Determining the age of the fossils is also very difficult. He says paleo-anthropology is more like a court of law than a physics laboratory, where we reassess and even redraw the whole family tree on finding a new fragment (Price and Feinman 1993 , 1997a , 1997b , c , d , e , f ).

Although we humans differ considerably from apes, genetically we are closest to them of all hominoids. As far as our genetic composition is concerned we share 98.4% of our genetic material with chimpanzees, while gorillas are only 2.3% different from us genetically. These statistics show great similarities and suggests that the last ancestor shared by the great ape and human lines was probably a chimpanzee-like creature. Thus, changes in genetic regulatory mechanism play an important role in the evolution of different lineages.

Geography had quite an important influence in shaping the development of humanity. It was in the middle of the Tertiary period (5–25 million of years ago) when the climate was much warmer and wetter than what it is today, and tropical forests grew across much of Africa, Europe and Asia that an increase in the variety of mammals occurred. Many species of apes lived in these forests, including one that is considered by some to be the ancestor of modern humans.

Then sometime around 5 million years ago, towards the end of the Tertiary time, the global temperatures began to cool, ice caps formed at the poles, and the climate grew drier. As a result, the area of the tropical forests grew smaller, giving way to expanses of open woodland and grasslands. These developments did not occur all at once but evolved slowly. In East Africa the hominoids groups were trapped in shrinking patches of forest. Before this they had lived in the trees and moved on four feet when travelling around the forest bed. Now in order to cross wide stretches of open land quickly, some hominoids began walking on two feet, like modern humans. It was under these conditions that the hominids split from this ape or the hominoid group. It is believed that this whole process of hominization began in Africa, which is the only continent where fossils of early hominids dating back to four - 5 million years are found today. The distinction was made due to their new mode of locomotion, known as “bipedalism” (walking on two feet rather than on four). These early hominids are called australopithecines: specialist believe in addition to walking upright with two feet, they had comparatively larger brains and they depended on tools for their survival (Bahn 2002a , b , c , d , e ).

While studying the fossils we also need to study how these beings modified objects and landscapes, thus creating an archaeological record. It is important to study both, as it helps us to understand how human beings evolved into what they are today. Some of the best evidence comes from sites at Hadar, Swartkrans, Olduvai, Laetoli and Koobi Fora. Some new fossil finds from Ethiopia, Kenya and Tanzania have pushed back the age of the earliest known hominids and have modified our understanding of their appearance and behavior (Price and Feinman 1993 , 1997a , 1997b , c , d , e , f ).

The oldest known Australopithecus species is A. Anamensis , which dates approximately 4 million years ago. The remaining early hominid fossils have been assigned to the species A. Afarensis , which includes the famous Lucy from Hadar (see Fig.  1 - A. Afarensis, Lucy from Hadar, Ethiopia), Ethiopia. Other direct evidence of hominid bipedalism is a fossilized trail of footprints some 3.6 million years old, found in eastern Africa at Laetoli, Tanzania (see Fig.  2 - Replica of the fossilized trail of footprints some 3.6 million years old, found in eastern Africa at Laetoli, Tanzania are exhibited in the National Museum of Nature and Science, Tokyo, Japan). Another evolutionary trend that occurred during this time was the change in dental pattern. Fossils of hominids dating back to three to 2 million years from southern and eastern African sites have small front teeth and large cheek teeth. Some form of gracile australopithecine is thought to have evolved into the first members of the genus homo about 2 million years ago, known as Homo habilis (which literally means handyman). Anthropologists continue to disagree about what caused this complete transition from ape to humans. Whereas many paleontologists believe that more than one species belonging to the genus homo may have co-existed in eastern Africa during the early Pleistocene, along with the robust australopithecines (species with massive teeth and jaws). It is also during this time that we find the oldest undisputed stone tools from Ethiopia, classified under the Oldowan tradition. By 1.8 million years ago the early homos had either disappeared or they evolved into H. erectus (upright man), the first member of the genus homo to spread out of Africa into Asia and Europe (see Fig.  3 - H. erectus (upright man), the first member of the genus homo to spread out of Africa into Asia and Europe.). They were taller than modern humans, but were very much like us in other respects, though their brains were still much smaller. It is assumed that they were capable of using fire and speech to a certain extent, and went, to far areas like Central Asia, Southwest Asia, South Asia, Europe and China. The Acheulean stone-tool tradition is associated with them. Archaeologists traditionally assign the Oldowan and Acheulean traditions to a single period known as the Lower Paleolithic in Europe and the Early Stone Age in Africa. But in recent years archaeologists have concluded that it is quite misleading to associate any Stone Age traditions uniquely to a single hominid species. Traditions can vary due to the availability of particular resources in different areas. Keeping all the above details in mind, paleoanthropologists consider a logical link between H. erectus , the more primitive hominids and our own species. They believe that the Acheulean stone – tool making ability was a determinant factor in the migration of early hominids out of Africa into new environments. The earliest handaxes found from outside of Africa are in Ubeidiya, Israel. While the punctuationists and gradualists continue to debate about the fate of H. erectus and the origin of the H. sapiens , up to this time indicates that our own species issued from H. erectus (Schultz and Lavenda 1998a , b , c , d ).

A. Afarensis , Lucy from Hadar, Ethiopia ( https://en.wikipedia.org/wiki/Lucy_(Australopithecus )

Replica of the fossilized trail of footprints some 3.6 million years old, found in eastern Africa at Laetoli, Tanzania are exhibited in the National Museum of Nature and Science, Tokyo, Japan

H. erectus (upright man), the first member of the genus homo to spread out of Africa into Asia and Europe. ( http://animals.wikia.com/wiki/Homo_erectus )

Towards the end of the Tertiary Period the global climate had started to cool down and this continued into the next Quaternary Period, also known as the Ice Age (see Fig.  4 - Map of the Last Ice Age.). In spite of its name the climate was not cold all the time: there were frequent warm intervals interglacial periods, separate from the cold, dry glacial periods. As these changes were quite rapid, the animals and plants sometimes found it difficult and sometimes impossible to adapt and to survive under the new climatic changes. H. erectus responded to these changes by developing a bigger brain. This meant that greater intelligence was available now for problem solving. One million years ago the brain of H. erectus was approximately three - quarters the size of the modern human brain (Haywood 1997 , 1989 n.d. ,). But recent studies have shown that stone tools found in association with animal bones were not used for slaughtering animals but for cleaning the skins and cutting up the meat. Plants probably formed a large part of their diet. No ornaments or art work is found and neither is there any evidence of them burying their dead (Bahn 2002a , b , c , d , e ). So the question remains, was H. erectus man’s ancestor?

Map of the Last Ice Age. ( http://www.kerbtier.de/Pages/Themenseiten/enPhylogenie.html )

It was sometime between 500,000 and 200,000 years ago that the fossils of H. erectus start to disappear from the fossil records and were replaced by fossils that show a mosaic of features found in H. erectus and H. sapiens (literally wise men). Though these fossils differ considerably from one another they collectively are known as archaic H. sapiens . They are poorly dated and paleoanthropologists find them difficult to classify and to relate specifically to H. sapiens . There are two major interpretations of the evidence of these fossil records. Punctuationists favor the “replacement model”: according to them H. erectus was a single, long- lived, geographically dispersed species, and had only one sub-population most probably located in Africa around 200,000 to 130,000 years ago that underwent evolutionary changes there to produce H. sapiens . Their descendants then later migrated to other regions. Whereas, the gradualists favor the “regional continuity model”. According to it H. erectus eventually evolved into H. sapiens gradually throughout the entire regions, retaining regionally distinct physical characteristics (Schultz and Lavenda 1998a , b , c , d ). However, archaeologists have found no record to back the claim that either H. erectus or archaic H. sapiens was truly our ancestor. There is no evidence of personal ornamentation - jewelry, beads, or any kinds of art form, exists, nor are there paintings, sculptures or engravings that show that use of more than basic instincts. The ability to think and reason is still missing (Ingold 1994a , b , c ). Therefore, whether they were really our ancestor, might be questioned.

In Europe a specie known as Neanderthals flourished between 130,000 and 35,000 years ago, they are considered by some to be the descendant of archaic H. sapiens . A large collection of fossils remains, tells us that they were shorter and more robust than modern H. sapiens with bulbous noses that helped them to survive cold conditions by reducing heat loss. But during the 1980s, new evidence revealed that Neanderthals appeared in Europe and western Asia at the same time anatomically modern H. sapiens appeared in Africa. These dates and the new evidence have made paleoanthropologists revise their traditional understanding of the relationship between Neanderthals and modern peoples. The archaic H. sapiens could no longer be considered the ancestors of the Neanderthals as data found in Israel suggests that the Neanderthals and the moderns lived side by side in south-western Asia for at least 45,000 years without losing their anatomical distinctiveness. Even more puzzling is the fact that both were using tools made in the same way. Mousterian experts disagree whether Neanderthals created a religion and whether hunting was important to them, but there is compelling fossil evidence from many sites and regions that they buried their dead and looked after their sick and old. Moreover, it seems probable, given the scant evidence for any form of art or ornamentation, that they did not make use of symbols, which is a critical element in the development of human language. Another thing that concerns us today is whether the Neanderthals and the moderns interbred, and whether the modern human populations today contain any Neanderthal genes. This situation creates a dispute among historians today (Schultz and Lavenda 1998a , b , c , d ). There are many questions that come to mind. Why did the Neanderthals look after their sick and old? Why did they start to bury their dead? Why were they using tools? Some believe that all these actions can be related to basic instincts. Tool use is not a distinctive characteristic of humans but; animals too may use or even make simple tools; however, using tools to make other tools does distinguish humans from animals. For example, the sea otter wields a rock to break open the shell of an abalone. And the anthropologist Jane Goodall has observed chimpanzees using a variety of tools in their daily life: thrashing about with branches for display, using clubs and missiles for defense, selecting a twig and stripping its bark to probe the nests of termites and attract them to the stick, then to be eaten by the wise chimp. West African chimps even use stone and wooden hammers to crack and open nutshells (Price and Feinman 1993 , 1997a , 1997b , c , d , e , f ).

So, the question becomes was the Neanderthal man’s ancestor? Because when it came to something like creating symbols for speech they were unable to do so, since it required more brain capacity, reasoning and intelligence that they unfortunately lacked. The question whether the Neanderthals and modern humans interbred was recently addressed by paleoanthropologists who claim, that there was no interbreeding between the two. Mitochondrial DNA studies suggest that all humans living today are part of a relatively homogeneous population that originated in Africa within the last few hundred thousand years. In 1997, genetic researchers extracted and decoded a mitochondrial DNA fragment for the original Neander Valley specimen. The analysis revealed significant differences in its DNA from all living humans, suggesting that there was an ancient split between the two lineages, perhaps more than 500,000 years ago. Although, Neanderthals did not disappear from Western Europe until 30,000 years ago, possibly later, it is a common belief that modern H. sapiens may have forced them into extinction (Bahn 2002a , b , c , d , e ).

Now let’s enter the last and the most important phase of human evolution. This phase is further divided into two phases. During the first phase, there is a general consensus among paleoanthropologists today that modern human ( H. sapiens ) evolved in Africa sometime between 100,000 to 150,000 years ago and spread around the globe. Recent studies in genetic evolution also support the view that Africa was the home of the original human population. However, debate continues about the nature of their dispersal. Most believe that a spreading wave of modern humans replaced existing populations of archaic H. sapiens entirely. This process of dispersal was complex and involved multiple movements of people and genes. In the caves of Qafzeh and Skhul in Israel remains of modern humans similar to found in Ethiopia and Tanzania, (150,000–100,000 years ago), and in South Africa (100,000–90,000 years ago) have been found. This is the first evidence that we have of modern humans (if they were) outside of Africa. Though, the fossils of these modern humans are still associated with the archaic stone tool traditions (like those of the Neanderthals ), and like the ones also associated with the race of modern Africans (Bahn 2002a , b , c , d , e ).

According to paleoanthropologists a second phase began roughly around 40,000 years ago, when “a behavioral revolution” took place: whether it was the continuity of the race or whether the race of the: “so called out of Africa H. sapiens ” was totally replaced by the current human race are two questions that are under continuing discussion. But no one can disregard or refute the dramatic changes of many both in anatomy and in behavior, that have taken place over the last 40,000 years when compared with the previous million years. Recent evidence from molecular biology has added support to this picture of rapid and recent change, resulting in the current humans that are not only genetically but also behaviorally and anatomically modern. Evidence also points to an African center for the origin of modern humans. As they moved out of Africa they very soon replaced the variety of other Homo geneses roaming the world (Ingold 1994a , b , c ).

Modern humans besides having many biological differences from other homo species are according to some “closer to the angles”. We possess many attributes that differentiate us from other species. Our large brain and intelligence enables us to think rationally and make decisions rather than to follow basic instincts like other species. We as humans have moved from purely instinctual behavior to reason and thought. We, in a given situation may flee from a fire, but we can also turn back into the same fire to save someone else (Price and Feinman 1993 , 1997a , 1997b , c , d , e , f ). It is during this phase, that we see fully developed linguistic and modern technological skills which appear to have developed in the modern man. There is general disagreement as to when this really happened, as the evidence found in this regard is both uncertain and open to doubt. Some believe it was 100,000 years ago, whereas others say it was as recent as 50,000 to 40,000 years ago. However, according to the archaeological record, it is only after 50,000 years that we find abundant examples of art and advanced technology. The first uncontested ritual behavior evidence that we have are the ostrich eggshells beads found from Enkapune ya Muto (Kenya) dating to 46,000 years ago. Beside these archaeologists also witness the appearance of ornaments, engravings, sculptures, and other form of symbolism, which unmistakably confirm the presence of modern human language. But the fossils of this time remain ambiguous, because they lack any anatomical evidence for linguistic abilities. Despite lack of evidence found in the fossils, the archaeological record speaks louder than words. The manipulation of these symbols (ornaments, engravings, etc.) are linked with the fundamental improvements that occurred in technical abilities, which undoubtedly played an important role in the global spread of the modern human species. Their ability to invent new technologies and cope with different environments helped them to colonize the globe at a rapid speed (Bahn 2002a , b , c , d , e ).

So, it would not be wrong to presume at this stage, that it was the abilities to talk, think, reason and communicate that differentiated modern humans from all other creatures before him. Clifford Geertz of Princeton University has described humans as,

“...toolmaking, talking, symbolizing animals: Only they laugh; only they know when they will die; only they disdain to mate with family members; only they contrive those visions of other worlds called art. They have not just mentality but consciousness, not just needs but values, not just fears but conscience, not just a past but a history. Only they have culture.”

According to the famous anthropologist Leslie White, culture is our “extrasomatic” means of survival, it is the nonbiological, nongenetic behavior and sociability that have carried us through the millennia and spread us into diverse environments across the planet. So, in short, culture is a group of ideas and actions that are learned and transmitted from one generation to the next generation. Human culture embodies our experiences and behaviors which are summarized in our language and are transferred to us through our parents and peers. It is as impossible to have human identity without social contact as it is to have biological existence without parents. There is a famous story that Tarzan of the comic book and movie was an ape before he met Jane. It is only culture that enables us to find our place on earth, to create Gods, to anticipate death, to travel to the worlds beyond, and last but not the least to study archaeology, in order to find answers about our past (Price and Feinman 1993 , 1997a , 1997b , c , d , e , f ). It was this cultural development, which was both very rapid and at an alarming speed, that until today evolutionists, biological scientist, paleoanthropologist and many more have been unable to understand. For example, if we can believe the evolutionist man’s ancestor first appeared on earth 4 million years ago and then slowly evolved into Modern Human only around 40,000 years ago. While keeping this in mind we witness, that after 40,000 to 10,000 years age man not only developed new technologies, but he also modified his environment. And only 10,000 years have taken him from bows and arrows to thermonuclear weapons, and the production of the latter has taken only twenty more years (The New Encyclopedia Britannica n.d. ).

Another important fact that seems to refute the claim of the evolutionists today is that, there are no signs of any intermediate forms found in the fossil records. Charles Darwin, who is known as the father of the theory of evolution, as state in his book, The Origins of Species claims,

“If my theory be true, numberless intermediate varieties, linking most closely all of the species of the same group together must assuredly have existed... Consequently, evidence of their former existence could be found only amongst fossil remains.”(Darwin 1964 )

The fossil records today show few intermediate forms; on the other hand, we see fully-formed living species seem to emerge suddenly without any evolutionary transitional form between them. This lack of factual evidence is enough to back their claim that all living species are created separately, and that life appeared on earth all of a sudden and fully-formed. Derek V. Ager, a famous British evolutionist admits this fact by saying;

“The point emerges that if we examine the fossil record in detail, whether at the level of Orders or of Species, we find – over and over again – not gradual evolution, but the sudden explosion of one group at the expense of another.”(Ager 1976 )

The fact that all living species were created separately, suddenly and fully-formed without any evolutionary ancestor is yet again backed by evolutionist biologist Douglas Futuyma, who claimed,

“Creation and evolution, between them, exhaust the possible explanations for the origin of living things. Organisms either appeared on the earth fully developed or they did not. If they did not, they must have developed from pre-existing species by some process of modification. If they did appear in a fully developed state, they must indeed have been created by some omnipotent intelligence.”(Futuyma 1983 )

Fossil records today back this claim that all living species emerged fully developed and in a perfect state on earth.

Let’s counter Charles Darwin’s claim about the ‘origin of man’, that he evolved from some ape-like creatures. The evolutionists who back him claim, that during the 5 million years of man’s evolution, man evolved from one stage of species to another. How these different stages evolved one after the other have already been discussed in detail. So, in short, according to evolutionists who give counter arguments claiming that, the first stage that is, Australopithecus also known as South African ape is nothing but an old ape that has become extinct. Extensive research was carried out by anatomists from both England and America, namely, Lord Solly Zuckerman and Prof. Charles Oxnard, have showed that they belonged to an ordinary ape species that became extinct and bore no resemblance to humans (Zuckerman 1970a , b ; Oxnard 1970 ).

The next stage of evolution is Homo , which is further divided into Homo habilis , Homo erectus and Homo sapiens . Each of these is considered to be one another’s ancestor. However recent fossil findings by the paleoanthropologist have revealed that Australopithecus , Homo habilis , and Homo erectus lived in different parts of the world at the same time (Walker 1980 ; Kesol 1970 ; Leakey 1971 ). And certain groups of Homo erectus lived until modern times. Homo sapiens and Neanderthals have also co-existed at the same time and also in the same region (Kluger 1996 ). The invalidity of this claim becomes more obvious when paleontologist fail to find any evolutionary trends in these so-called ancestors of man. Stephen Jay Gould a paleontologist from Harvard University explains this deadlock:

“What has become of our ladder if there are three coexisting lineages of hominids ( A. africanus , the robust australopithecines , and H. habilies ), none clearly derived from another? Moreover, none of the three display any evolutionary trends during their tenure on earth.”(Gould 1976 )

This claim was also backed by Lord Solly Zuckerman, who studied Australopithecus fossils for fifteen years, finally concluded that there is, in fact, no such family tree branching out from ape-like creatures to man.

He also formed a ‘spectrum of science’, in which sciences ranging from those he considered scientific to those he considered unscientific were put together. According to him, the most scientific, that is, depending on the concrete data-fields are chemistry and physics. After these are the biological sciences and then the social sciences. The most unscientific sciences which are at the far end of the spectrum include the extra-sensory perception, telepathy, sixth sense and finally human evolution. He explains this formation;

“We then move right off the register of objective truth into those fields of presumed biological science, like extrasensory perception or the interpretation of man’s fossil history, where to the faithful [evolutionist] anything is possible – and where the ardent believer [in evolution] is sometimes able to believe several contradictory things at the same time.”(Zuckerman 1970a , b )

Keeping all the arguments and counter arguments in mind with respect to the theory of man’s evolution, I shall conclude by quoting a few sentences from Harun Yahya’s book, ‘Fascism: The Bloody Ideology of Darwinism’ ,

“...the theory of evolution is a claim evidently at variance with scientific findings. The theory’s claim on the origin of life is inconsistent with science, the evolutionary mechanisms it proposes have no evolutionary power, and fossils demonstrate that the intermediate forms required by the theory never existed. So, it certainly follows that the theory of evolution should be pushed aside as an unscientific idea.”(Yahya 2002a , b , c )

Richard C. Lewontin who is a well-known geneticist and an evolutionist from Harvard University claims that he is first and foremost a materialist and then a scientist. He confesses;

“It is not that the methods and institutions of science somehow compel us to accept a material explanation of the phenomenal world, but, on the contrary, that we are forced by our a priori adherence to material causes to create an apparatus of investigation and a set of concepts that produce material explanations, no matter how counter-intuitive, no matter how mystifying to the uninitiated. Moreover, that materialism is absolute, so we cannot allow a Divine Foot in the door.”(Lewontin 1997 )

So, in short, the evolutionists who give materialist answers to the hundreds of questions that arise in the conscious thinking mind of the modern man today, are not only further creating confusions but have in a way failed to satisfy the logical and rational human mind. How can one believe that an unconscious matter can create life? How can one believe that matter created thousands and thousands of living things and living species with their own distinct attributes, qualities, and characteristics when scientist until today are not sure even how a small thing such as a simple cell can be formed? They know that it is formed when proteins come together, but how they come together, in what ratios and form a cell is a process that they have failed to understand (The Usborne Internet Linked Encyclopedia of World History, s.v., “human cell” 2000 ). For many years now engineers from around the world have been trying to make a three-dimensional television that can match the quality of the human eye. Yes, they have being successful in making a three-dimensional television screen, but you cannot watch it without putting on special glasses; moreover, it only creates artificial three dimension. Similarly ears, engineers have failed to produce a device that can ensure the same quality and clarity of sound that the human ear perceives. Another thing which is even more important than seeing and hearing abilities is the ‘consciousness’ that man has been blessed with (Yahya 2002a , b , c ). It is this consciousness that creates the major difference between man and all other living species. It is this that takes man one step ahead of all others. It is this ability that makes us flee from a fire, but we can go back in the same fire to save someone. It is this ability that helps us to understand and comprehend, that despite of the best of qualities given to us in this world, there are certain things that are still beyond our reach, control and comprehension. Even we humans have limitations, and this concept was well taken and understood even by early man since antiquity. He also knew that he had no control over the elements and there was some “Divine Force” somewhere, which had everything under its control. Hence it would not be wrong to presume here, that it was at this point in time around approximately 50,000 to 40,000 years ago, that the modern man entered the scene, and all the other species predating him were not actually ‘man’, or his ancestors. Hence, man was born a man with the best of qualities and a consciousness to understand the ‘Divine’ which has helped him not only to conquer but also to rule the world.

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Ager, A. Derek. 1976. The nature of the fossil record. Proceedings of the British Geological Association 87: 133.

Google Scholar  

Bahn, G. Paul, ed. 2002a. The atlas of world archaeology . Oxfordshire: Andromeda. Publisher is Checkmark Books.

Bahn, Paul G., ed. 2002b. The Atlas of World Archaeology , 16. Oxfordshire: Andromeda.

Bahn, Paul G., ed. 2002c. The Atlas of World Archaeology , 22. Oxfordshire: Andromeda.

Bahn, Paul G., ed. 2002d. The Atlas of World Archaeology , 27. Oxfordshire: Andromeda.

Bahn, Paul G., ed. 2002e. The Atlas of World Archaeology , 28–29. Oxfordshire: Andromeda.

Braidwood, J. Robert. 1967. Prehistoric men . Toledo: Scott Foresman and Company.

Braidwood, Robert J. 1964a. Prehistoric men , 1–9. Chicago: Chicago History Museum.

Braidwood, Robert J. 1964b. Prehistoric men , 10–11. Chicago: Chicago History Museum.

Darwin, Charles. 1964. The origin of species: a facsimile of the first edition , 184. Boston: Harvard University Press.

Futuyma, J. Douglas. 1983. Science on trail , 197. New York: Pantheon Books.

Gould, J.S. 1976. Natural history . Vol. 85, 30. New York: American Museum of Natural History.

Grant, Micheal. 1952. Ancient history . London: First Harper TorchBook Edition.

Grant, Micheal. 1965. Ancient history , 5. New York: Harper and Row.

Haywood, John. 1997. Atlas of world history . Oxford: Sandcastle Books.

Haywood, John. 1989. n.d. The Atlas of Past Times , 12–13.

Ingold, Tim, ed. 1994a. Companion encyclopaedia of anthropology . New York: Routledge.

Ingold, Tim, ed. 1994b. Companion encyclopedia of anthropology , 87–88. New York: Routledge.

Ingold, Tim, ed. 1994c. Companion encyclopedia of anthropology , 88–92. New York: Routledge.

Kesol, J.A. 1970. Physical anthropology . 1st ed, 221. New York.

Kluger, Jeffrey. 1996. Not So Extinct After All: The Primitive Homo Erectus May Have Survived Long Enough To Coexist With Modern Humans. Time .

Leakey, M.D. 1971. Olduvai Gorge . Vol. 3, 272. Cambridge: Cambridge University Press.

Lewontin, Richard. 1997. The demon-haunted world , 28. The New York Review of Books.

Marwick, Arthur. 1970. The nature of history , 192–193. London: Palgrave Macmillan.

Oxnard, E. Charles. 1970. The place of australopithecines in human evolution: grounds of doubt. Nature 258 (New York: 389.

Article   Google Scholar  

Price, Douglas T., and Gary M. Feinman. 1993. Images of the Past . 2nd ed. California: Mayfield Publishing Company.

Price, T. Douglas, and Gary M. Feinman. 1997a. Images of the Past , 10. Houston: Mayfield Publishing.

Price, T. Douglas, and Gary M. Feinman. 1997b. Images of the Past , 16. Houston: Mayfield Publishing.

Price, T. Douglas, and Gary M. Feinman. 1997c. Images of the Past , 12. Houston: Mayfield.

Price, T. Douglas, and Gary M. Feinman. 1997d. Images of the Past , 39–40. Houston: Mayfield.

Price, T. Douglas, and Gary M. Feinman. 1997e. Images of the Past , 39. Houston: Mayfield.

Price, T. Douglas, and Gary M. Feinman. 1997f. Images of the Past , 41. Houston: Mayfield.

Schultz, A. Emily, and Robert H. Lavenda. 1998a. Anthropology- a perspective on the human condition . Belmont: Wadsworth.

Schultz, Emily A., and Robert H. Lavenda. 1998b. Anthropology- a perspective on the human condition , 128–152. California: Mayfield.

Schultz, Emily A., and Robert H. Lavenda. 1998c. Anthropology- a perspective on the human condition , 175. California: Mayfield.

Schultz, Emily A., and Robert H. Lavenda. 1998d. Anthropology- a perspective on the human condition , 176. California: Mayfield.

n.d. The New Encyclopedia Britannica . 15th ed, 995.

Jane Bingham, Fiona Chandler, Sam Taplin. 2000. The Usborne Internet-linked Encyclopedia of World History. In World History. Usborne. ISBN 0746041683, 9780746041680.

Walker, Alan. 1980. Science . Vol. 207, 1103.

Yahya, Harun. 2002a. Fascism: the bloody ideology of Darwinism . Turkey: Kultur.

Yahya, Harun. 2002b. Fascism: The Bloody Ideology of Darwinism , 234. Ankara: Global Publishing.

Yahya, Harun. 2002c. Fascism: The Bloody Ideology of Darwinism , 232–234. Turkey: Global Publishing.

Zuckerman, Solly. 1970a. Beyond the ivory tower , 75–94. New York: Toplinger Publications.

Zuckerman, Solly. 1970b. Beyond The Ivory Tower , 19. New York: Toplinger Publications.

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In unicellular organisms, sexual reproduction typically begins with the fusion of two cells (plasmogamy) followed by the fusion of their two haploid nuclei (karyogamy) and finally meiosis. Most work on the evolution of sexual reproduction focuses on the benefits of the genetic recombination that takes place during meiosis. However, the selection pressures that may have driven the early evolution of binary cell fusion, which sets the stage for the evolution of karyogamy by bringing nuclei together in the same cell, have seen less attention. In this paper we develop a model for the coevolution of cell size and binary cell fusion rate. The model assumes that larger cells experience a survival advantage from their larger cytoplasmic volume. We find that under favourable environmental conditions, populations can evolve to produce larger cells that undergo obligate binary cell fission. However, under challenging environmental conditions, populations can evolve to subsequently produce smaller cells under binary cell fission that nevertheless retain a survival advantage by fusing with other cells. The model thus parsimoniously recaptures the empirical observation that sexual reproduction is typically triggered by adverse environmental conditions in many unicellular eukaryotes and draws conceptual links to the literature on the evolution of multicellularity.

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A study on the evolution of forest landscape patterns in the fuxin region of china combining sc-unet and spatial pattern perspectives.

1. Introduction

2. study region and data, 2.1. overview of the study region, 2.2. research information, 3. research methodology, 3.1. spatial and channel reconstruction convolution, 3.2. sc-unet forest extraction model, 3.3. a morphology-based approach to analyzing forest spatial patterns, 3.4. multivariate weighted results, 4. experiments and analysis, 4.1. experimental data, 4.2. model training, 4.3. experimental results, 5. discussion, 5.1. spatial and temporal changes of forest land in the fuxin region, 5.2. evolution of the spatio-temporal patterns of forest landscape categories in the fuxin region based on mspa, 5.3. limitations and future research, 6. conclusions, author contributions, institutional review board statement, informed consent statement, data availability statement, acknowledgments, conflicts of interest.

• Peng, X.; He, G.; She, W.; Zhang, X.; Wang, G.; Yin, R.; Long, T. A Comparison of Random Forest Algorithm-Based Forest Extraction with GF-1 WFV, Landsat 8 and Sentinel-2 Images. Remote Sens. 2022 , 14 , 5296. [ Google Scholar ] [ CrossRef ]
• Zhu, H.; Zhang, B.; Song, W.; Xie, Q.; Chang, X.; Zhao, R. Forest Canopy Height Estimation by Integrating Structural Equation Modeling and Multiple Weighted Regression. Forests 2024 , 15 , 369. [ Google Scholar ] [ CrossRef ]
• Zhang, B.; Zhu, H.; Xu, W.; Xu, S.; Chang, X.; Song, W.; Zhu, J. A Fourier–Legendre Polynomial Forest Height Inversion Model Based on a Single-Baseline Configuration. Forests 2024 , 15 , 49. [ Google Scholar ] [ CrossRef ]
• De Frenne, P.; Lenoir, J.; Luoto, M.; Scheffers, B.; Zellweger, F.; Aalto, J.; Ashcroft, M.; Christiansen, D.; Decocq, G.; De Pauw, K.; et al. Forest microclimates and climate change: Importance, drivers and future research agenda. Glob. Chang. Biol. 2021 , 27 , 2279–2297. [ Google Scholar ] [ CrossRef ]
• Ripple, W.J.; Bradshaw, G.A.; Spies, T.A. Measuring forest landscape patterns in the Cascade Range of Oregon, USA. Biol. Conserv. 1991 , 57 , 73–88. [ Google Scholar ] [ CrossRef ]
• Wang, J.; Yang, X. A hierarchical approach to forest landscape pattern characterization. Environ. Manag. 2012 , 49 , 64–81. [ Google Scholar ] [ CrossRef ]
• Tariq, A.; Jiango, Y.; Li, Q.; Gao, J.; Lu, L.; Soufan, W.; Almutairi, K.; Habib-ur-Rahman, M. Modelling, mapping and monitoring of forest cover changes, using support vector machine, kernel logistic regression and naive bayes tree models with optical remote sensing data. Heliyon 2023 , 9 , e13212. [ Google Scholar ] [ CrossRef ]
• Cross, M.D.; Scambos, T.; Pacifici, F.; Marshall, W.E. Determining effective meter-scale image data and spectral vegetation indices for tropical forest tree species differentiation. IEEE J. Sel. Top. Appl. Earth Obs. Remote Sens. 2019 , 12 , 2934–2943. [ Google Scholar ] [ CrossRef ]
• Zhang, X.; Zhang, Y.; Liu, L.; Zhang, J.; Gao, J. Remote sensing monitoring of the subalpine coniferous forests and quantitative analysis of the characteristics of succession in east mountain area of Tibetan Plateau—A case study with Zamtange county. Agric. Sci. Technol. 2011 , 12 , 926–930. [ Google Scholar ]
• Franklin, S.E.; Hall, R.J.; Moskal, L.M.; Maudie, A.J.; Lavigne, M.B. Incorporating texture into classification of forest species composition from airborne multispectral images. Int. J. Remote Sens. 2000 , 21 , 61–79. [ Google Scholar ] [ CrossRef ]
• Cheng, K.; Wang, J. Forest type classification based on integrated spectral-spatial-temporal features and random forest algorithm—A case study in the Qinling mountains. Forests 2019 , 10 , 559. [ Google Scholar ] [ CrossRef ]
• Xie, G.; Niculescu, S. Mapping and monitoring of land cover/land use (LCLU) changes in the Crozon peninsula (Brittany, France) from 2007 to 2018 by machine learning algorithms (support vector machine, random forest, and convolutional neural network) and by post-classification comparison (PCC). Remote Sens. 2021 , 13 , 3899. [ Google Scholar ] [ CrossRef ]
• Haq, M.A.; Rahaman, G.; Baral, P.; Ghosh, A. Deep learning based supervised image classification using UAV images for forest areas classification. J. Indian Soc. Remote Sens. 2021 , 49 , 601–606. [ Google Scholar ] [ CrossRef ]
• Chai, J.; Zeng, H.; Li, A.; Ngai, E.W.T. Deep learning in computer vision: A critical review of emerging techniques and application scenarios. Mach. Learn. Appl. 2021 , 6 , 100134. [ Google Scholar ] [ CrossRef ]
• Dhanya, V.G.; Subeesh, A.; Kushwaha, N.L.; Vishwakarma, D.K.; Kumar, T.N.; Ritika, G.; Singh, A.N. Deep learning based computer vision approaches for smart agricultural applications. Artif. Intell. Agric. 2022 , 6 , 211–229. [ Google Scholar ] [ CrossRef ]
• Moshayedi, A.J.; Roy, A.S.; Kolahdooz, A.; Shuxin, Y. Deep learning application pros and cons over algorithm deep learning application pros and cons over algorithm. EAI Endorsed Trans. AI Robot. 2022 , 1 , 7. [ Google Scholar ]
• Ghasemi, Y.; Jeong, H.; Choi, S.H.; Park, K.B.; Lee, J.Y. Deep learning-based object detection in augmented reality: A systematic review. Comput. Ind. 2022 , 139 , 103661. [ Google Scholar ] [ CrossRef ]
• Chang, X.; Zhang, B.; Zhu, H.; Song, W.; Ren, D.; Dai, J. A Spatial and Temporal Evolution Analysis of Desert Land Changes in Inner Mongolia by Combining a Structural Equation Model and Deep Learning. Remote Sens. 2023 , 15 , 3617. [ Google Scholar ] [ CrossRef ]
• Dong, S.; Wang, P.; Abbas, K. A survey on deep learning and its applications. Comput. Sci. Rev. 2021 , 40 , 100379. [ Google Scholar ] [ CrossRef ]
• Long, J.; Shelhamer, E.; Darrell, T. Fully convolutional networks for semantic segmentation. In Proceedings of the IEEE Conference on Computer Vision and Pattern Recognition, Boston, MA, USA, 7–12 June 2015; pp. 3431–3440. [ Google Scholar ]
• Wang, J.; Song, L.; Li, Z.; Sun, H.; Sun, J.; Zheng, N. End-to-end object detection with fully convolutional network. In Proceedings of the IEEE/CVF Conference on Computer Vision and Pattern Recognition, Nashville, TN, USA, 20–25 June 2021; pp. 15849–15858. [ Google Scholar ]
• Wu, S.; Shi, J.; Chen, Z. HG-FCN: Hierarchical grid fully convolutional network for fast VVC intra coding. IEEE Trans. Circuits Syst. Video Technol. 2022 , 32 , 5638–5649. [ Google Scholar ] [ CrossRef ]
• Wang, S.; Liu, C.; Zhang, Y. Fully convolution network architecture for steel-beam crack detection in fast-stitching images. Mech. Syst. Signal Process. 2022 , 165 , 108377. [ Google Scholar ] [ CrossRef ]
• Zhu, H.; Zhang, B.; Chang, X.; Song, W.; Dai, J.; Li, J. A Study of Sandy Land Changes in the Chifeng Region from 1990 to 2020 Based on Dynamic Convolution. Sustainability 2023 , 15 , 12931. [ Google Scholar ] [ CrossRef ]
• Ronneberger, O.; Fischer, P.; Brox, T. U-net: Convolutional networks for biomedical image segmentation. In Proceedings of the Medical Image Computing and Computer-Assisted Intervention–MICCAI 2015: 18th International Conference, Munich, Germany, 5–9 October 2015; proceedings, part III 18. Springer International Publishing: Berlin/Heidelberg, Germany, 2015; pp. 234–241. [ Google Scholar ]
• Su, Z.; Li, W.; Ma, Z.; Gao, R. An improved U-Net method for the semantic segmentation of remote sensing images. Appl. Intell. 2022 , 52 , 3276–3288. [ Google Scholar ] [ CrossRef ]
• John, D.; Zhang, C. An attention-based U-Net for detecting deforestation within satellite sensor imagery. Int. J. Appl. Earth Obs. Geoinf. 2022 , 107 , 102685. [ Google Scholar ] [ CrossRef ]
• Meena, S.R.; Soares, L.P.; Grohmann, C.H.; Westen, C.; Bhuyan, K.; Singh, R.P.; Floris, M.; Catani, F. Landslide detection in the Himalayas using machine learning algorithms and U-Net. Landslides 2022 , 19 , 1209–1229. [ Google Scholar ] [ CrossRef ]
• Alsabhan, W.; Alotaiby, T.; Dudin, B. Detecting buildings and nonbuildings from satellite images using U-Net. Comput. Intell. Neurosci. 2022 , 2022 , 4831223. [ Google Scholar ] [ CrossRef ] [ PubMed ]
• Wang, H.; Miao, F. Building extraction from remote sensing images using deep residual U-Net. Eur. J. Remote Sens. 2022 , 55 , 71–85. [ Google Scholar ] [ CrossRef ]
• Yan, C.; Fan, X.; Fan, J.; Wang, N. Improved U-Net remote sensing classification algorithm based on Multi-Feature Fusion Perception. Remote Sens. 2022 , 14 , 1118. [ Google Scholar ] [ CrossRef ]
• Zhang, R.; Zhu, D. Study of land cover classification based on knowledge rules using high-resolution remote sensing images. Expert Syst. Appl. 2011 , 38 , 3647–3652. [ Google Scholar ] [ CrossRef ]
• Chen, B.; Xia, M.; Huang, J. Mfanet: A multi-level feature aggregation network for semantic segmentation of land cover. Remote Sens. 2021 , 13 , 731. [ Google Scholar ] [ CrossRef ]
• Navin, M.S.; Agilandeeswari, L. Comprehensive review on land use/land cover change classification in remote sensing. J. Spectr. Imaging 2020 , 9 , a8. [ Google Scholar ] [ CrossRef ]
• Kupidura, P. The Comparison of Different Methods of Texture Analysis for Their Efficacy for Land Use Classification in Satellite Imagery. Remote Sens. 2019 , 11 , 1233. [ Google Scholar ] [ CrossRef ]
• Liu, M.; Shi, J.; Li, Z.; Li, C.; Zhu, J.; Liu, S. Towards better analysis of deep convolutional neural networks. IEEE Trans. Vis. Comput. Graph. 2016 , 23 , 91–100. [ Google Scholar ] [ CrossRef ]
• Xu, R.; Samat, A.; Zhu, E.; Li, E.; Li, W. Unsupervised Domain Adaptation with Contrastive Learning-Based Discriminative Feature Augmentation for RS Image Classification. Remote Sens. 2024 , 16 , 1974. [ Google Scholar ] [ CrossRef ]
• Lu, Y.; Li, H.; Zhang, C.; Zhang, S. Object-Based Semi-Supervised Spatial Attention Residual UNet for Urban High-Resolution Remote Sensing Image Classification. Remote Sens. 2024 , 16 , 1444. [ Google Scholar ] [ CrossRef ]
• Han, Y.; Guo, J.; Yang, H.; Guan, R.; Zhang, T. SSMA-YOLO: A Lightweight YOLO Model with Enhanced Feature Extraction and Fusion Capabilities for Drone-Aerial Ship Image Detection. Drones 2024 , 8 , 145. [ Google Scholar ] [ CrossRef ]
• Wu, Q.; Feng, D.; Cao, C.; Zeng, X.; Feng, Z.; Wu, J.; Huang, Z. Improved Mask R-CNN for Aircraft Detection in Remote Sensing Images. Sensors 2021 , 21 , 2618. [ Google Scholar ] [ CrossRef ] [ PubMed ]
• Li, S.; Ganguly, S.; Dungan, J.L.; Wang, W.; Nemani, R.R. Sentinel-2 MSI radiometric characterization and cross-calibration with Landsat-8 OLI. Adv. Remote Sens. 2017 , 6 , 147. [ Google Scholar ] [ CrossRef ]
• Gašparović, M.; Jogun, T. The effect of fusing Sentinel-2 bands on land-cover classification. Int. J. Remote Sens. 2018 , 39 , 822–841. [ Google Scholar ] [ CrossRef ]
• Lamquin, N.; Woolliams, E.; Bruniquel, V.; Gascon, F.; Gorroño, J.; Govaerts, Y.; Leroy, V.; Lonjou, V.; Alhammoud, B.; Barsi, J.A.; et al. An inter-comparison exercise of Sentinel-2 radiometric validations assessed by independent expert groups. Remote Sens. Environ. 2019 , 233 , 111369. [ Google Scholar ] [ CrossRef ]
• Su, W.; Zhang, M.; Jiang, K.; Zhu, D.; Huang, J.; Wang, P. Atmospheric correction method for Sentinel-2 satellite imagery. Acta Opt. Sin. 2018 , 38 , 0128001. [ Google Scholar ] [ CrossRef ]
• Yin, F.; Lewis, P.E.; Gómez-Dans, J.L. Bayesian atmospheric correction over land: Sentinel-2/MSI and Landsat 8/OLI. Geosci. Model Dev. 2022 , 15 , 7933–7976. [ Google Scholar ] [ CrossRef ]
• Li, J.F.; Wen, Y.; He, L.H. SCConv: Spatial and channel reconstruction convolution for feature redundancy. In Proceedings of the 2023 IEEE/CVF Conference on Computer Vision and Pattern Recognition, Vancouver, BC, Canada, 17–24 June 2023; IEEE Press: New York, NY, USA, 2023; pp. 6153–6162. [ Google Scholar ]
• Chen, C.F.; Oh, J.; Fan, Q.; Pistoia, M. SC-Conv: Sparse-complementary convolution for efficient model utilization on CNNs. In Proceedings of the 2018 IEEE International Symposium on Multimedia (ISM), Taichung, Taiwan, 10–12 December 2018; pp. 97–100. [ Google Scholar ]
• Wu, Y.; He, K. Group Normalization. In Proceedings of the European Conference on Computer Vision (ECCV), Munich, Germany, 8–14 September 2018; Volume 3, pp. 3–19. [ Google Scholar ]
• Li, X.; Wang, W.; Hu, X.; Yang, J. Selective kernel networks. In Proceedings of the 2019 IEEE/CVF Conference on Computer Vision and Pattern Recognition, Long Beach, CA, USA, 15–20 June 2019; IEEE Press: New York, NY, USA, 2019; pp. 510–519. [ Google Scholar ]
• Ogundokun, R.O.; Maskeliunas, R.; Misra, S.; Damaševičius, R. Improved CNN based on batch normalization and Adam optimizer. In Proceedings of the International Conference on Computational Science and Its Applications, Malaga, Spain, 4–7 July 2022; Springer International Publishing: Cham, Switzerland, 2022; pp. 593–604. [ Google Scholar ]
• Lange, S.; Helfrich, K.; Ye, Q. Batch normalization preconditioning for neural network training. J. Mach. Learn. Res. 2022 , 23 , 1–41. [ Google Scholar ]
• Yu, X.; Zheng, Z.; Meng, L.; Li, L. Scene Recognition of Remotely Sensed Images Based on Bayes Adjoint Batch Normalization ; Geomatics and Information Science of Wuhan University: Wuhan, China, 2023. [ Google Scholar ]
• Yang, X.; Zhu, Y.; Guo, Y.; Zhou, D. An image super-resolution network based on multi-scale convolution fusion. Vis. Comput. 2022 , 38 , 4307–4317. [ Google Scholar ] [ CrossRef ]
• Hu, H.; Chen, Y.; Xu, J.; Borse, S.; Cai, H.; Porikli, F.; Wang, X. Learning implicit feature alignment function for semantic segmentation. In Proceedings of the European Conference on Computer Vision, Tel Aviv, Israel, 23–27 October 2022; Springer Nature: Cham, Switzerland, 2022; pp. 487–505. [ Google Scholar ]
• Ren, S.; Zhao, N.; Wen, Q.; Han, G.; He, S. Unifying Global-Local Representations in Salient Object Detection with Transformers. IEEE Trans. Emerg. Top. Comput. Intell. 2024 , 8 , 2870–2879. [ Google Scholar ] [ CrossRef ]
• Soille, P.; Vogt, P. Morphological segmentation of binary patterns. Pattern Recognit. Lett. 2009 , 30 , 456–459. [ Google Scholar ] [ CrossRef ]
• Vogt, P.; Ritters, K. GuidosToolbox: Universal digital image object analysis. Eur. J. Remote Sens. 2017 , 50 , 352–361. [ Google Scholar ] [ CrossRef ]
• Xiong, C.; Wu, Z.; Zeng, Z.Y.; Gong, J.Z.; Li, J.T. Spatiotemporal evolution of forest landscape pattern in Guangdong-HongKong-Macao Greater Bay Area based on “Spatial Morphology-Fragmentation-Aggregation”. Acta Ecol. Sin. 2023 , 43 , 3032–3044. [ Google Scholar ]
• Pan, Y.; Jiao, S.; Hu, J.; Guo, Q.; Yang, Y. An Ecological Resilience Assessment of a Resource-Based City Based on Morphological Spatial Pattern Analysis. Sustainability 2024 , 16 , 6476. [ Google Scholar ] [ CrossRef ]
• Zhu, H.; Zhang, B.; Song, W.; Dai, J.; Lan, X.; Chang, X. Power-Weighted Prediction of Photovoltaic Power Generation in the Context of Structural Equation Modeling. Sustainability 2023 , 15 , 10808. [ Google Scholar ] [ CrossRef ]
• Hu, L.; Xu, N.; Liang, J.; Li, Z.; Chen, L.; Zhao, F. Advancing the Mapping of Mangrove Forests at National-Scale Using Sentinel-1 and Sentinel-2 Time-Series Data with Google Earth Engine: A Case Study in China. Remote Sens. 2020 , 12 , 3120. [ Google Scholar ] [ CrossRef ]
• Yang, L.; Driscol, J.; Sarigai, S.; Wu, Q.; Chen, H.; Lippitt, C.D. Google Earth Engine and Artificial Intelligence (AI): A Comprehensive Review. Remote Sens. 2022 , 14 , 3253. [ Google Scholar ] [ CrossRef ]
• Li, L.; Zhu, L.; Xu, N.; Liang, Y.; Zhang, Z.; Liu, J.; Li, X. Climate Change and Diurnal Warming: Impacts on the Growth of Different Vegetation Types in the North–South Transition Zone of China. Land 2022 , 12 , 13. [ Google Scholar ] [ CrossRef ]

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Wang, F.; Yang, F.; Wang, Z. A Study on the Evolution of Forest Landscape Patterns in the Fuxin Region of China Combining SC-UNet and Spatial Pattern Perspectives. Sustainability 2024 , 16 , 7067. https://doi.org/10.3390/su16167067

Wang F, Yang F, Wang Z. A Study on the Evolution of Forest Landscape Patterns in the Fuxin Region of China Combining SC-UNet and Spatial Pattern Perspectives. Sustainability . 2024; 16(16):7067. https://doi.org/10.3390/su16167067

Wang, Feiyue, Fan Yang, and Zixue Wang. 2024. "A Study on the Evolution of Forest Landscape Patterns in the Fuxin Region of China Combining SC-UNet and Spatial Pattern Perspectives" Sustainability 16, no. 16: 7067. https://doi.org/10.3390/su16167067

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• Published: 06 August 2024

Early evolution of small body size in Homo floresiensis

• Yousuke Kaifu   ORCID: orcid.org/0000-0003-0483-104X 1 ,
• Iwan Kurniawan   ORCID: orcid.org/0000-0002-7816-7471 2 ,
• Soichiro Mizushima 3 ,
• Junmei Sawada 4 ,
• Michael Lague 5 ,
• Ruly Setiawan 2 ,
• Indra Sutisna 6 ,
• Unggul P. Wibowo 6 ,
• Gen Suwa   ORCID: orcid.org/0000-0003-3880-2458 1 ,
• Reiko T. Kono   ORCID: orcid.org/0000-0001-8488-6301 7 ,
• Tomohiko Sasaki 8 ,
• Adam Brumm   ORCID: orcid.org/0000-0002-2276-3258 9 &
• Gerrit D. van den Bergh   ORCID: orcid.org/0000-0003-1507-3336 10  

Nature Communications volume  15 , Article number:  6381 ( 2024 ) Cite this article

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• Archaeology
• Biological anthropology
• Palaeontology

Recent discoveries of Homo floresiensis and H. luzonensis raise questions regarding how extreme body size reduction occurred in some extinct Homo species in insular environments. Previous investigations at Mata Menge, Flores Island, Indonesia, suggested that the early Middle Pleistocene ancestors of H. floresiensis had even smaller jaws and teeth. Here, we report additional hominin fossils from the same deposits at Mata Menge. An adult humerus is estimated to be 9 − 16% shorter and thinner than the type specimen of H. floresiensis dated to ~60,000 years ago, and is smaller than any other Plio-Pleistocene adult hominin humeri hitherto reported. The newly recovered teeth are both exceptionally small; one of them bears closer morphological similarities to early Javanese H. erectus . The H. floresiensis lineage most likely evolved from early Asian H. erectus and was a long-lasting lineage on Flores with markedly diminutive body size since at least ~700,000 years ago.

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Introduction.

The So’a Basin in central Flores, Indonesia, is a key region for elucidating the origin and evolution of H. floresiensis , a diminutive hominin species known from the Late Pleistocene at Liang Bua, a limestone cave in western Flores 1 , 2 . As with another small-bodied Homo discovered in Luzon 3 , the evolutionary history of this insular hominin species has been the subject of protracted debate 4 . Previous field studies of the Early to Middle Pleistocene (Calabrian–Chibanian) sequence of the So’a Basin have recovered fossil remains of endemic fauna (dwarfed Stegodon , Komodo monitor, giant rat, birds, crocodiles and tortoises) 5 , 6 , technologically simple stone artefacts (the oldest of which date back to at least 1.02 ± 0.02 million years ago [Ma]) 7 , 8 , and, importantly, a fragmentary mandible and six isolated teeth of a small-sized hominin 9 . These hominin fossils were excavated from a sandstone layer of fluvial origin (Layer II) of the upper fossil-bearing interval at the Mata Menge site, which is dated to between 0.65 and 0.773 Ma 5 , 6 . These fossils exhibit general morphological affinities to the type series of H. floresiensis from Liang Bua (0.1–0.06 Ma) 10 and to early H. erectus from Java (1.1–0.8 Ma) 11 , but lack the unique molar specializations characterizing the former and were substantially smaller than the latter 9 .

Overall, the Mata Menge fossils suggest that they represent an ancestral segment of the Liang Bua H. floresiensis lineage, and that the Flores hominins are dwarfed descendants of large-bodied early Asian H. erectus 9 . Some cladistic/phylogenetic analyzes, however, support a direct evolutionary link between H. floresiensis and smaller-bodied basal Homo such as H. habilis or even Australopithecus 12 , 13 , 14 . It is important to resolve this controversy in order to elucidate the pattern and timing of body size evolution in the Flores hominins.

Notably, the Mata Menge mandible and teeth are slightly smaller than the type specimens of H. floresiensis from Liang Bua. This suggests that drastic dentognathic reduction had occurred on Flores by the early Middle Pleistocene epoch, more than 600,000 years before the earliest fossil evidence for H. floresiensis at Liang Bua. Until now, however, the lack of postcranial elements in the Mata Menge assemblage had limited our understanding of body size evolution on Flores.

In this paper, we report the discovery and morphology of a hominin postcranial fossil from Mata Menge, an extremely small distal humeral shaft (SOA-MM9) (Fig.  1 ). This specimen and two small-sized teeth (SOA-MM10 and SOA-MM11) were recovered as additions to the existing hominin assemblage from Layer II at this site (Table  1 ). Our histomorphic examination confirms the adult status of the humerus. We also show that shaft morphology is more similar to small-bodied Homo (e.g., LB1 and H. naledi ) than to Australopithecus (e.g., A.L. 288-1), and a molar crown (SOA-MM11) bears closer shape similarities to early Javanese H. erectus than to early African Homo . The increased Mata Menge fossil sample supports its classification to an early representative of H. floresiensis , which probably experienced drastic body size reduction from large-bodied Asian H. erectus sometime between ~1.0 and 0.7 Ma.

a – f SOA-MM9 humerus in anterior, lateral, posterior, medial, proximal, and distal views. g Micro-CT section of SOA-MM9 at the distal end indicated in ( c ). h and i LB1 humerus in anterior and medial views. Note the hollowed area on the posterior surface of the distal end (filled triangles in f and g ), which serves as an osteometric landmark (hOF point). Abbreviations: ant = anterior, post = posterior, lat = lateral, med = medial, DT = deltoid tuberocity, NF = nutrient foramen, HS = location for histological section. j SOA-MM10 right d c . From left to right, occlusal, labial, and lingual views. k SOA-MM11 left M 3 . From left to right, occlusal, buccal, and lingual views.

Context and geological age

All hominin fossils originated from the top of a ribbon-shaped, indurated pebbly sandstone layer (Layer II), which was deposited in a small stream channel on a volcaniclastic alluvial fan 5  ~ 0.7 Ma ago. This age estimate is based on the identification of the Brunhes-Matuyama boundary 15 dated at 0.773 Ma by palaeomagnetic measurements combined with a series of fission track dates on zircons in tuffaceous strata stratigraphically 16.5 m below Layer II 5 , 6 . A minimum age of 0.65 ± 0.02 Ma for Layer II is provided by a 40 Ar/ 39 Ar date on single hornblende crystals from an airfall tephra (PGT-2) occurring stratigraphically at 14 m above Layer II.

Layer II, with a maximum thickness of 50 cm, overlies a reddish paleosol (Layer III) with an undulating erosional contact. A series of massive, tuffaceous clay-rich mudflow layers (Layers Ia-f) sealed off Layers II and III subsequently (Fig.  2 ; Supplementary Note  1 ). The humerus fragment SOA-MM9 was retrieved in several pieces within one week of opening Excavation 32 A in 2013, but was recognized as such only in 2015 after reconstruction in the laboratory. The specimen was damaged in the process of excavating it from the extremely compact sandstone of Layer II. A maxillary deciduous canine (d c : SOA-MM10) was excavated in 2015 at ~5 cm below the boundary between Layers I and II, while a mandibular third molar (M 3 : SOA-MM11) was excavated in 2016 at ~15 cm below the top of Layer II. All hominin fossils are concentrated in the upper part of Layer II, while fossils of other fauna tend to be distributed more evenly in this unit. There is evidence for fluvial transportation of the fossils prior to burial, with many (but not all) specimens fractured (apart from excavation damage), weathered and/or rounded to some extent 16 . However, the three hominin fossils described here show minimal/no evidence of abrasion. Taphonomic and sedimentological observations suggest that the hominin fossils were deposited during a moderate to low-energy flow regime in the stream, following a relatively brief period on the surface during which the remains were disarticulated (Supplementary Note  1 ). Shortly after incorporation of the fossils in the stream bed the entire stream valley was filled with a 6.5 m thick sequence of mudflows. Succeeding field excavations in 2017–2019 and 2023 have yielded no more hominin fossils from this site.

a Digital Elevation Map (DEM) of Flores showing the location of the So’a Basin and the cave Liang Bua. b DEM of the So’a Basin showing the location of the Mata Menge excavations. c Photo of the west baulk of the southern excavation sector (Sector 32C) in the upper fossil-bearing interval at Mata Menge. Layer III is a reddish sandy paleosol, overlain with an erosional contact by a sandy fluvial layer (Layer II). Layers II and III are covered by a series of clay rich ashy mudflow units (Layers Ia-f). Deciduous canine SOA-MM10 was recovered at 5 cm below the top of Layer II (indicated with the blue dashed rectangle; the blue rectangle is enlarged in e). Also note the large Stegodon bones resting on top of Layer II and covered by the mudflow units. d Photo of the northwest corner of the excavation in sector 32 A, taken on 7 November 2013, four weeks after the retrieval of the hominin humerus fragment SOA-MM9. The fossil was excavated from the top of Layer II, with the approximate position indicated with the dashed blue oval. e Detail of the contact between Layer I and Layer II at the spot of the deciduous canine SOA-MM10. f SOA-MM10 still partly embedded in the sandstone of Layer II. g Mata Menge excavation grid (UTM Zone 51 L) showing the 1 ×1 m quadrants excavated towards the end of the 2016 field season in gray. The positions of the hominin fossils described in this paper are indicated with green dots, those described previously 9 with gray dots. Light shading represents the step trench excavated in 2010, which first revealed the presence of the Mata Menge upper fossil-bearing interval bone bed. h SOA-MM11 surrounded by its sandstone matrix when excavated in 2016. The maps ( a and b ) created with GeoMapApp ( www.geomapapp.org ) / CC BY / CC BY (Ref. 67 )”.

Developmental age of the humerus (SOA-MM9)

This specimen is an undistorted, distal half of the right humeral shaft that measures 88 mm in maximum preserved length (Fig.  1 ; Supplementary Note  2 ). Despite its small size, cortical bone histomorphology of SOA-MM9 clearly indicates its adult status. We examined its development stage based on age-associated increase of osteons and related structures, a method widely utilized for age estimates of extant and fossil hominins 17 , 18 , 19 , 20 , 21 .

Histological sections were examined for cortical samples taken at the mid-posterior shafts of SOA-MM9 (‘HS’ in Fig.  1 ) and from a modern human sample (N = 20, see Supplementary Data  1 ). To allow for regional variation in osteon formation within each bone area 21 , data were collected from two additional (nearby) sites in the midshaft for all the modern human ( H. sapiens ) specimens (Fig.  3a ). In two parameters indicative of bone maturity, Osteon Population Density (OPD) 21 and Haversian Canal Index (HCI), SOA-MM9 was found to exhibit distinctly greater values (OPD = 16.5, HCI = 0.85) than in any of the modern human subadult humeri (0.0–8.9 and 0.0–0.63, respectively) (Fig.  3b , Supplementary Data  1 ). The values for the Mata Menge humerus are also greater than the means of our modern human adult samples (13.6 and 0.78, respectively), indicating that the SOA-MM9 individual was well within adulthood at time of death. Although the external cortical surfaces of SOA-MM9 exhibit microscopic damages that might have reduced one of the marginal osteons to half (~100 microns) (Fig.  3c ), such post-depositional alterations would have limited impact on our age estimation. Even if we assume surface abrasion of 200 microns, the OPD value for SOA-MM9 would drop only slightly to ~15.8. Furthermore, the dominance of secondary osteons in the outer cortex (Fig.  3c ) indicates that subperiosteal bone growth during the growth period had already been terminated in this individual 22 .

a Three sites in a cortical section of the mid-posterior humeral shaft used for the histomorphological analyzes. ‘Middle field’ is a circle drawn in the outer one-third of the midsagittal line, and the ‘lateral field’ and ‘medial field’ are corresponding circles 3 mm apart from there. b Age-related histomorphometric values for SOA-MM9 and modern humans. The data from the three sites are plotted for all the modern human specimens, whereas the data for SOA-MM9 is from the lateral field only. The dotted lines are the means of the modern human adult subsample. See Supplementary Data  1 for the original data. c Cortical section of the posterior midshaft of SOA-MM9 observed by an ultra-high-definition microscope (VHX-7000, Keyence). The original image (left) and the same image colored for intact secondary osteons (red), fragmentary osteons (orange and yellow), and the intact inner surface (blue) (right). Note the dominance of secondary osteons around the exterior surface (downside of the images). Only one section (shown here) was produced to minimize the damage to the original specimen. d Lateral supracondylar ridge of SOA-MM9 and its CT section (the arrows). Note the weak but distinct development of the ridge as a slight eversion.

No evidence of pathology was found in SOA-MM9. Cortical bone thinning and woven bone would be pathognomic of some metabolic disorders 17 , but these features are not evident in SOA-MM9. The relative cortical bone thickness of SOA-MM9 (0.07: the ratio of cortical bone thickness relative to the humeral shaft circumference) (Supplementary Data  1 ) is almost identical to the mean for the modern human adult sample (0.069). Patients with osteogenesis imperfecta, which may lead to short stature, exhibit subnormal OPD values 19 , a tendency that is in opposition to the SOA-MM9 condition. Additionally, the weak but distinct development of the lateral supracondylar ridge of SOA-MM9 (Fig.  3d ) indicates normal development of the extensor carpi radialis longus muscle.

Humeral size

In all available dimensions of shaft diameter/circumference and length, SOA-MM9 is smaller than LB1 ( H. floresiensis ) and any other adult individuals of small-bodied fossil hominins ( Australopithecus and H. naledi : Supplementary Data  2 ). Its minimum circumference (46 mm) is less than U.W. 101-283 (47.5 mm), BOU-12/1 (52 mm), and the smallest humeri in our prehistoric modern human sample (46.5 mm, N = 1050, see Supplementary Data  2 ). Centroid size at the ~19% level cross-section is also the smallest compared to any sampled adult specimens of Australopithecus , Paranthropus , and Homo including H. naledi and Liang Bua H. floresiensis (Fig.  4 , Supplementary Data  3 ). The distal shaft length measured between the NF (nutrient foramen) and hOF (superior margin of the hollow leading to the olecranon fossa) points of SOA-MM9 (58 mm) is distinctly shorter than the other hominin fossils, including LB1 (64 mm) (Supplementary Data  2 ), although the vertical position of NF is variable in human humeri 23 .

Symbols: circle = australopith; square = Homo ; black = adult (possibly including some late adolescents); white = subadult. The specimens in each morphological group are listed in Supplementary Data  3 along with their size values. A standard box plot (the median at horizontal line and the ‘whiskers’ representing minimum/maximum values) is shown for a sample (N = 21) of modern humans. Transverse dashed line for Group 2 connects specimens belonging to Au. sediba and Au. sp. indet . Vertical dashed lines represent the smallest (proximal section) and largest (distal section) values obtained for SOA-MM9, which is smaller than all adult Plio-Pleistocene fossil hominin humeri (including LB1) and similar in size to specimens of H. naledi . Note that fully adult status (proximal epiphyseal fusion or developed muscle markings on the shaft) cannot be confirmed for some specimens such as the smallest individuals of Group 1b (SKX 10924) and Group 4 (SK 2598 and SK 24600). See Supplementary Data  3 for other notes.

The fragmentary nature of SOA-MM9 precludes a precise reconstruction of its original length, but it can be estimated as follows. First, the preserved proximo-posterior end of SOA-MM9 is very close to the 50% level because this portion exhibits the following suite of features characteristic of hominin humeral midshafts: 1) The radial sulcus (spiral groove) is present not on the lateral surface but on the posterolateral aspect, seen as a flattened area in CT slice nos. 1900 and 2000 (‘RS’ in Fig.  5 ); 2) in lateral view, the anterior margin exhibits a slight concavity around no. 1900 (‘AM’ in Fig.  5 , see the surface rendered image on the left side), indicating that this part, which is ~13 mm below the preserved proximo-posterior end, leads to the deltoid tuberosity proximally. The distal margin of the deltoid tuberosity is situated on the antero-lateral surface at the 48.4% level in our modern human sample mean (N = 366, range: 43−53%), 51% in LB1 24 , and 48% in KNM-WT 15000 25 ; 3) NF is present 21 mm distal to the preserved proximo-posterior end of SOA-MM9. The projected distance along the shaft from the 50% level and the lower margin of the NF is 23 mm in LB1, 1 mm in KNM-WT15000, 15 mm in MH2 ( Au. sediba ) 26 , and 21.2 mm in our modern human sample mean (N = 366, SD = 10.0 mm, range = –2 to 59 mm). Each of the above three characters shows substantial variation, but their simultaneous expression at the preserved proximal shaft strongly suggests that SOA-MM9’s proximo-dorsal end was very close to the original 50% level. This positioning is consistent with other shaft morphologies exhibited by SOA-MM9 (Supplementary Note  3 , Fig.  5 , Supplementary Figs.  1 and 2 , and Supplementary Data  4 ).

Left: Surface rendered images. Clockwise from top left: anterior, posterior, right lateral and left lateral views. Right: CT sections and reconstructed cortical bones at the slice level indicated by the numerals (250 − 2183). The slice nos. 607, 573 and 639 are estimated 19% levels (the best estimate and probable range: see “Methods”). The slice thickness of these CT sections is 0.04 mm, so that the difference of 100 corresponds to 4 mm. Cortical bone reconstruction was made with reference to the intact bones in the same or adjacent slices. The preserved cortical bones are in pink, the extrapolated portions are in blue, and the regions ‘transplanted and trimmed’ from nearby slices are in other colors. No. 900 was reconstructed after a minor positional correction of the small bone indicated by the star, which is slightly dislocated inward in the current reconstruction.

Next, distally, the CT section no. 250 of SOA-MM9 (Fig.  5 ), which is sliced at the hOF point, corresponds to the 12.5–14% level (or 11.5–15% more broadly) of maximum length (as explained below). Because of the observed allometric relationships that shorter modern human humeri tend to have relatively large distal epiphyses (Supplementary Note  3 ), we referred to the following two samples to draw upon the above figures. One is the short-statured prehistoric Holocene population from Tanegashima Island, Japan (N = 13, maximum humeral length: 245–292 mm), and the other is a subset of short humeri from the prehistoric Jomon population from Japan (N = 10, maximum humeral length: 240–250 mm). The means of the hOF levels in these samples were 12.5% and 14%, respectively, with their ranges 11.5 − 13.5% and 13 − 15%, respectively. The equivalent values in short hominin fossil humeri are 13% in both LB1 and A.L. 288-1.

Based on the above evaluations of SOA-MM9 humeral shaft preservation (proximal, 50% level; distal, 12.5 − 14% or 11.5 − 15% level), the original maximum humeral length of SOA-MM9 is estimated to be 211–220 mm or 206–226 mm, respectively. Alternatively, if we apply the mean ratios between the NF-hOF length and the maximum humeral length in our modern human male and female samples (0.29 to 0.30, Supplementary Data  2 ), the estimated maximum length of SOA-MM9 is 194–200 mm, but we surmise that this is less reliable given the weak correlation between the two measurements ( r  = 0.37).

Comparative humeral morphology

SOA-MM9 lacks characteristic features of Australopithecus distal humeri, such as a prominent flange-like lateral supracondylar ridge, a projecting medial supracondylar crest, and marked curvature in the sagittal plane, although expression of these traits tends to be weak in comparatively gracile specimens of this genus 26 , 27 , 28 , 29 . With respect to cross-sectional shape (distal shaft 19% level), SOA-MM9 is similar to small-bodied Homo ( H. naledi and H. floresiensis ) in having a mediolaterally narrow profile that is unusual among the comparative groups (Fig.  6 ). It is different, however, from small-bodied Australopithecus individuals such as A.L. 288-1, whose humerus is only slightly longer than estimated for SOA-MM9 (Fig.  6 ). Procrustes distances support the cross-section shape variation summarized by the PCA results. Whereas the distances of SOA-MM9 to group mean shapes (Supplementary Fig.  4a ) or individual specimens (Supplementary Fig.  4b ) of H. naledi and H. floresiensis do not exceed the degree of within-species variation represented by our modern human sample, the same distances to the other fossil taxa are much greater.

a Principal component analysis (PCA) of distal diaphyseal shape among 40 fossil hominin humeri. Convex hulls define the fossil groups (Groups 1-3 = australopith, Group 4 =  H. habilis , Group 5 =  H. erectus s.l ., Group 6 =  H. naledi : see Supplementary Data  3 for more details) and the dotted line connects LB1 to the average shape of the three sampled sections of SOA-MM9. The two diaphyseal outlines depict shape variation exclusively along PC1. SOA-MM9 is extreme along PC1 and is most similar in overall shape (based on Procrustes distance) to LB1 and to specimens of H. naledi (Group 6). See Supplementary Fig.  3 for a two-dimensional presentation of this result. b Cross-sectional outlines of the distal humeral diaphysis of SOA-MM9 and LB1 in comparison to group averages for modern humans and the fossil hominin morphological groups. Shape configurations are shown scaled by anteroposterior width (with anterior towards the top and lateral to the right).

Maxillary deciduous canine (d c : SOA-MM10)

This right tooth preserves a complete crown and a broken, short segment of the root (Fig.  1j ). The crown is extremely small, situated well below the reported range of H. sapiens (Fig.  7a , and Supplementary Table  1 ), in a similar way to the previously reported Mata Menge d c s (Fig.  7b ). The specimen has a primitive, relatively low distal shoulder that resembles Australopithecus and Sangiran H. erectus homologs, although PCAs based on four or five linear measurements indicate that this morphology is marginally within the large variation seen in H. sapiens (Supplementary Fig.  5 ). Occlusal wear exposes a small dentine patch on the cusp tip and a thin line of dentine on the elongated distal incisal margin. The latter suggests the presence of a primitive, tall mesial cusp configuration on its occluding deciduous first molar (dm 1 ), as known for an early Javanese H. erectus dm 1 from Sangiran, S7–67 30 .

Horizontal crown dimensions of maxillary deciduous canines (d c : a ) and mandibular third molars (M 3 : c ), as well as the previously reported mandibular deciduous canines (d c : b ). The large crosses in a and b indicate 2 SD ranges for the smallest-toothed modern population as for d c and d c (India 66 ). d Plots of PC scores derived from the normalized Elliptic Fourier Analysis (EFA) of M 3 crown contour. Shape differences along PC axes are shown on left teeth for two standard deviations from the origin. Symbols: ‘a’ (orange) = Au. afarensis , ‘a’ (magenta) = Au. africanus , ‘h’ (blue) = H. habilis sensu lato , ‘D’ (orange) = Dmanisi Homo , ‘z’ (violet) = Zhoukoudian H. erectus , ‘S’ (green) = early H. erectus (Sangiran Lower), ‘s’ (green) = early H. erectus (Sangiran Upper), ‘L’ (red) = Liang Bua, ‘S’ (red) = Soa (Mata Menge). ‘x’ = H. sapiens (Japan) for ( a ) and ( b ) H. sapiens (Japan) for ( c ). Specimen numbers are indicated for selected samples in each symbol. Source data are provided as a Source Data file.

Mandibular third molar (M 3 : SOA-MM11)

The preserved left crown has reduced distal cusp areas, distally protruding hypoconulid, and no distal interproximal facet (Fig.  1k ). Wear has flattened much of the occlusal surface, except for the metaconid that remains relatively high. Crown diameters are comparable to those of the smaller Liang Bua individual (LB6/1) and are marginally within the large variation exhibited by our global H. sapiens sample (Fig.  7c ). It has five principal cusps arranged in a ‘+’ pattern and is different from the mandibular third molars of Liang Bua H. floresiensis that exhibit a derived, four-cusped morphology (LB1, LB6/1) 31 , 32 . Occlusal crown contour examined by normalized Elliptic Fourier Analysis shows that it clusters firmly with Sangiran H. erectus and marginally with H. ergaster , having a mesiodistally short crown. It is outside the range of variation exhibited by H. habilis sensu lato , which is primarily characterized by a mesiodistally elongated and distally tapered crown (Fig.  7d , Fig.  8 , Supplementary Note  4 , Supplementary Fig.  6 ) with a tendency for better developed hypoconulids and accessory cusps 33 , 34 , 35 .

LB1 and LB6/1: Liang Bua H. floresiensis , Sangiran 22: early Javanese H. erectus , OH13: H. habilis .

Comparison of size with the Liang Bua H. floresiensis

In all available measurements of the mandibular body, postcanine teeth (P 3 , M 1/2 and M 3 ) and distal humerus, the Mata Menge fossils reported here or previously 9 are smaller than the Liang Bua H. floresiensis remains (LB1 and LB6/1) by 1–21% (Table  2 ). Stature estimates, based on humeral length of SOA-MM9 (211−220 mm) and LB1 (243 mm), are 103–108 cm and 121 cm, respectively, using the human pygmy model 36 ; or 93–96 cm and 102 cm, respectively, using the ape model 37 .

All the ten hominin remains so far discovered from Mata Menge were excavated from a narrow area (about 7 m × 20 m) within the upper part of Layer II (Fig.  2 ). We previously reported that one mandible fragment (SOA-MM4) and six isolated teeth (SOA-MM1, 2, 5, 6, 7 and 8) from this collection represent at least one adult and two children 9 (Table  1 ). The limited wear on the mesial incisal margin of the new right d c (SOA-MM10) does not match the extensive wear on the distal incisal margin of the previously reported right d c (SOA-MM8), but the degree of wear does not preclude the possibility that SOA-MM10 and SOA-MM7 (left d c ) are from the same child. While SOA-MM7 was recovered from sieving, this specimen and SOA-MM10 were found within 6 m horizontal distance of each other. The new permanent molar (SOA-MM11), a moderately worn left M 3 , is obviously a different individual from SOA-MM1, a lightly worn left M 1 (or M 2 ) belonging to an adolescent (or a young adult if this was a M 2 ). Therefore, the current Mata Menge hominin assemblage includes at least four individuals including one adult, one adolescent/young adult, and two children (Table  1 ).

The observation that all four (or more) individuals are extremely diminutive supports the argument that small body size was not an idiosyncratic (individual) character but a population feature of the early Middle Pleistocene hominins of Flores. The markedly small deciduous teeth from at least two individuals, which are almost outside the large variation range of modern humans (Fig.  7 ), also indicate that the Mata Menge hominins had diminutive dental size at birth. Additionally, the strikingly small adult humerus (SOA-MM9) reported here demonstrates that this character was not limited to the dentognathic elements but also extended to upper arm size. On this note, it is worth highlighting that the two or more Mata Menge adult/adolescent individuals are consistently smaller than the two adults of Liang Bua H. floresiensis (Table  1 ). This strongly suggests that by ~0.7 Ma, hominins on Flores were already as small as, or perhaps slightly smaller than, the Late Pleistocene H. floresiensis (Supplementary Note  5 ).

Based on the previously recovered dentognathic sample, it was suggested that the Mata Menge fossils could be reasonably assigned to H. floresiensis 9 . Now that a new arm bone and additional dental remains belonging to this assemblage display strong affinities with the Liang Bua remains, we can more confidently classify these early Middle Pleistocene hominins into H. floresiensis . Notable minor differences between the two widely separated chronological forms include the lack of molar morphological specializations (see below) and possibly the smaller body and dental sizes in the earlier Mata Menge hominin.

This study also contributes to the debate over the origin and evolution of H. floresiensis . It was previously reported that the Mata Menge hominins now assigned to H. floresiensis were more similar to early Javanese H. erectus than to Australopithecus and H. habilis sensu lato in mandibular body form and M 1 (or M 2 ) shape 9 , a finding that runs contrary to hypotheses that assume a direct evolutionary link between H. floresiensis and pre- H. erectus hominins such as H. habilis 12 , 13 , 14 . The present study indicates that the shape similarity between the Mata Menge fossils and early Javanese H. erectus also applies to M 3 , and that the Mata Menge molars lack the unique specialization seen in the Liang Bua H. floresiensis homologies (i.e., four-cusped, mesiodistally shortened and somewhat distorted molar crowns 9 , 32 , 37 ). Therefore, archaic H. floresiensis at Mata Menge probably represents the dwarfed lineage of early Javanese H. erectus at a stage prior to unique molar specializations. Alternatively, if H. habilis s.l . was ancestral to Mata Menge/Liang Bua H. floresiensis , the latter would need to have experienced substantial molar size reduction of ~65–60% in mesiodistal and buccolingual crown diameters (from H. habilis means), and this accompanied by form changes comparable to the early Javanese H. erectus condition. Because no such allometric relationships are evident between molar crown size and form within H. habilis (Supplementary Note  6 ), the hypothesis that H. floresiensis is a direct lineal descendant of H. habilis s.l . is not supported. In contrast, molar size reduced from the Lower to Upper Sangiran dental assemblages (Fig.  7c ) without significant form changes (Fig.  7d ), confirming that such local evolution could occur. Additionally, although the humeral shaft morphology of SOA-MM9 does not indicate an affinity with either H. erectus or H. habilis , its cross-sectional shape is most similar to that of dwarfed taxa of Homo ( H. floresiensis and H. naledi ) and unlike that of small-bodied Australopithecus individuals.

Coupled with the recently revised arrival date for H. erectus on Java ( ~ 1.1 Ma, or at most younger than 1.3–1.5 Ma) 11 and hominins on Flores (1.0–1.27 Ma) 6 , as well as the reported craniometric and odontometric analyzes which almost unanimously support strong affinities of H. floresiensis with H. erectus (particularly early H. erectus from Java) 37 , 38 , 39 , 40 , 41 , 42 , the following evolutionary model emerges. The earliest Flores hominins appeared on this Wallacean island ~1.0–1.27 Ma, probably unintentionally (i.e., through accidental ‘rafting’, perhaps on tsunami debris), and possibly as part of the initial colonization of the Sunda Shelf region by early H. erectus . The Flores hominins experienced substantial body size reduction soon after this event (within ~300,000 years), despite the presence of large-bodied predators such as ~3 meter-long Komodo monitors and crocodiles from the earliest paleontological record ( ~ 1.4 Ma) onwards 6 . This implies that giant reptilians did not represent a serious predation threat for early H. floresiensis or its progenitors. This early evolutionary event was followed by long-term stability in hominin body size, possibly also in cultural adaptations (e.g., stone technology 6 , 7 , 8 ), and minor morphological specialization in the dentition. How the small brain size reported for the ~60,000 years old LB1 1 , 43 evolved still remains unknown. At present, however, the available fossil data imply that small body size had been a functional adaptation for these insular hominins during and slightly beyond the Middle Pleistocene and indeed potentially up until the arrival of H. sapiens on Flores around 50,000 years ago; an event that, we suspect, precipitated the demise of H. floresiensis 10 .

Permission to undertake excavations at Mata Menge was granted by the Indonesian State Ministry of Research and Technology (RISTEK permits 300/SIP/FRP/SM/VIII/2013 and 2183/FRP/SM/X/2015), the provincial government of East Nusa Tenggara in Kupang, and the Ngada District Administration. CT scan of SOA-MM9 was conducted in Tokyo with a permission issued in 2015 from the Geological Agency, Bandung.

CT scan and measurements

A micro-CT scan of SOA-MM9 was taken by using TXS320-ACTIS (Tesco Co.) at the National Museum of Nature and Science, Tokyo, with the following scanning parameters: 205 kV and 0.2 mA with a 0.5 mm thick copper plate prefilter, a 1024 × 1024 matrix, 0.04 mm pixel size and 0.04 mm slice interval (0.043 mm slice thickness). Micro-CT scans (voxel size = 0.156 mm) of 88 adult prehistoric Japanese (Holocene hunter-gatherer-fishers from the Jomon period) were also obtained for the length estimation of SOA-MM9 (see below). Linear measurements were taken using a spreading digital caliper (to the nearest 0.1 mm), an osteometric board (to the nearest 1.0 or 0.5 mm) and measuring tape (to the nearest 0.5 mm). Cross sectional properties were calculated based on CT scans at 15–50% vertical level of the shaft, using the software CT-Rugle (ver. 1.2, Medic Engineering Inc., Japan) and ImageJ (ver. 1.53f51, National Institutes of Health, USA).

Humeral analyzes

Comparative samples.

To characterize its humeral morphology as a specialized insular hominin group, SOA-MM9 was compared with a wide variety of Pliocene and Pleistocene Afro-Asian hominin humeri ( Ardipithecus , Australopithecus , Paranthropus , African early Homo , Dmanisi Homo , H. erectus / ergaster and H. naledi ), as well as H. floresiensis from Liang Bua (LB1) and a series of modern human samples including some short-statured populations. The individual specimens included for linear metric comparison and geometric morphometric analysis are shown in Supplementary Data  2 and 3 , respectively. The modern human samples used for the linear metric analysis are in Supplementary Data  2 , while the modern human sample for the geometric morphometric analysis (Supplementary Data  3 ) is a mixed-sex sample of adults collected (by J.M. Plavcan) at the Smithsonian National Museum of Natural History (Washington, DC) 44 . As for the geometric morphometric analysis, the fossil sample was divided into six morphological groups, five of which have been established by previous studies 29 , 44 , 45 , 46 , while a sixth group consists of five specimens attributed to Homo naledi . We also collected outline data from a scan of the humerus of the LB1 skeleton (i.e., LB1/50) attributed to H. floresiensis 1 . The adult/subadult status for each specimen was determined by the epiphysial fusion of the proximal and/or distal ends, or other information if available (e.g., dental development). The linear metric comparisons focus on the adult samples, while the geometric morphometric analysis contains some subadult specimens, as noted in Fig.  4 and Supplementary Data  3 . See Supplementary Note  7 for additional information about the H. naledi sample.

Key landmarks

SOA-MM9 lacks most of the widely used osteometric landmarks, but the following points are usable:

Nutrient foramen (NF): the distal margin of the nutrient foramen on the midshaft.

hOF point: the proximal margin of the hollow leading to the olecranon fossa.

Developmental age

Human bones undergo substantial histomorphic changes during development and much of adulthood. In a limb bone shaft, periosteal cortical bone growth occurs as deposition of circumferential lamellar bone and primary osteons with non-Haversian canals. The proportion of these primary structures decreases as secondary osteons (Haversian canal surrounded by concentric rings of lamellar bone) appear and increase through bone remodeling. In late adulthood, the cortical bone is dominated by secondary osteons 17 , 47 . This process is numerically demonstrated by the count or density of the elements of secondary osteon (intact osteons, fragmentary osteons, Haversian canals, resorption bay, etc.) 20 , 48 , 49 , 50 . However, because the rate of such histomorphic change varies considerably depending on the locus in a bone, regional differences must be considered in histomorphological age estimation 21 , 51 . Because no histomorphometric data was available in the literature for human humeral midshaft, we collected our referential data using modern Japanese humeri. Our sample, which consists of 10 adults, 6 adolescents, and 4 child individuals, was unearthed from cemeteries of the Edo period (17th-19th centuries A.D.) in Tokyo City and is stored at the National Museum of Nature and Science, Japan. Based on the standard ossification procedure 52 , we categorized those humeri with completely fused epiphyses to ‘adult,’ those with unfused proximal epiphysis and completely fused distal epiphysis to ‘adolescent,’ and those with separate epiphyses as ‘child.’ The adolescent category was further subdivided into ‘early adolescent’ (fusion at distal epiphysis only, N = 2), ‘mid adolescent’ (fusion at distal epiphysis and medial epicondyle, N = 2), and ‘late adolescent’ (proximal epiphysis partially fused, N = 2). We (J.S.) cut out a small piece of the bone from the mid-posterior shaft of the SOA-MM9 humerus, to prepare a sectional sample (‘HS’ in Fig.  1c ). The location of this section is 6.5 mm distal to the preserved proximal edge, and 14.5 mm proximal to the distal margin of the nutrient foramen, and is assumed to be slightly distal to the missing deltoid tuberosity. The obtained section covers a full thickness from its outer (periosteal) to inner (intrathecal) surfaces. After embedding in resin, we prepared a polished surface to observe with an ultra-high-definition microscope (VHX-7000, Keyence). From each of the modern human humeri, we prepared a cortical section at the posterior surface 10 mm distal to the lower end of the deltoid tuberosity. First, we cut out a small piece of the bone using a diamond cutter. After embedding in resin, a transverse section of 70 μm-thickness was made by a microtome (SP-1600, Leica) to observe under an ordinary light microscope (ImagerA1, Leica). We focused on the outer one-third of the cortical bone, because the periosteal region is an active bone growth field and is useful for histomorphometric growth studies 20 . To allow for regional variation mentioned above, we examined three adjacent loci: one on the midsagittal line (middle field), and the others on either side of it (lateral and medial fields) as illustrated in Fig.  3 . The section prepared for SOA-MM9 corresponds to the lateral field. In each observation field, we counted the numbers of intact secondary osteons (N.On), osteon fragments (N.Fr), resorption bays (N.Re), Haversian canals of the secondary osteons/osteon fragments (N.Ca), and non-Haversian canals (N.nCa), as defined elsewhere 20 , 53 . An ‘osteon fragment’ is a secondary osteon eroded by later-formed osteons. A structure straddling the border of the observation field was counted only if more than half of it was inside. We use the following three size-free parameters as measures of cortical bone growth and pathology.

1)Osteon Population Density (OPD): This widely used index, calculated here as (N.On+N.Fr+N.Re) per area (mm2) 21 , monitors the increase of secondary made structures. A greater OPD value reflects advanced growth stage.

2)Haversian Canal Index (HCI): This is a ratio of the secondary made canals. It is calculated as (N.Ca/(N.Ca+N.nCa)), and increases from 0 to 1 with bone growth.

3)Relative Cortical Bone thickness (rCBt): We define this index as the mean cortical bone thickness divided by the minimum circumference of each humeral shaft. The former is the average of the three cortical bone thicknesses at the medial, middle and lateral fields in Fig.  3 , which we measured using a public-domain ImageJ (U.S. National Institutes of Health, available at https://imagej.nih.gov/ij/ ).

Length estimation

The extant and fossil hominin humeri exhibit a uniform pattern of transition in cross-sectional shape (i.e., flatness, angle of the long axis, ratio between the cortical and total areas, and other features) from the mid- to the distal shaft levels (Supplementary Figs.  1 and 2 ). We (S.M. and Y.K.) refer to this information to reconstruct the original maximum humeral length (Martin no. 1a) of SOA-MM9.

Cross-sectional properties of the shaft

We (S.M.) used CT-Rugle 1.2 (Medic Engineering Co.) to calculate the cross-sectional properties of the humeral shaft.

Cross-sectional geometry of the shaft

Previous studies of fossil hominin humeri have demonstrated the taxonomic utility of the cross-sectional shape of the distal diaphysis sampled at ~19% of total (biomechanical) humerus length from the distal end 29 , 44 , 46 , 54 , 55 , 56 . The ~19% level of SOA-MM9 was located by Y.K. based on our estimate of its maximum length (211–220 mm), which was converted to the biomechanical humeral length using the ratio between the two (the former is 1.08% longer on average in our mixed-sex, prehistoric modern human (Jomon) sample: N = 88). The 19% level thus located is within the CT slice nos. 573–639. Therefore, we chose three slices, nos. 573, 607 (best estimate), and 639 for the present analysis. Two-dimensional coordinates were collected by M.L. from all three sections of SOA-M9 following the procedure described previously 44 (i.e., two Type 2 landmarks on the medial and lateral extremes of the specimen and 58 sliding semilandmarks on the anterior and posterior surfaces). Raw landmark configurations were superimposed into the same shape space using orthogonal least-squares generalized Procrustes (GPA) superimposition 57 , GPA was performed using tpsRelw software 58 and semilandmarks were allowed to slide along the diaphyseal outline using the criterion of minimized bending energy 59 . Subsequent to GPA, morphometric relationships were assessed with the use of Procrustes distances (Dp) as a measure of shape dissimilarity 60 and principal component analysis (PCA) as a means of visual summary (via ordination) of shape variation among the individual specimens.

Dental analysis

SOA-MM10 (maxillary deciduous canine: d c ) was compared with the available sample of fossil hominins ( Australopithecus and Early Pleistocene Homo ), as well as a sample of H. sapiens (Supplementary Table  2 ) by Y.K. Unfortunately, there is no deciduous teeth in the existing H. floresiensis assemblage from Liang Bua. SOA-MM11 (mandibular third molar: M 3 ) was compared with Liang Bua H. floresiensis and its claimed two major ancestral candidates, H. habilis and early Javanese H. erectus (Supplementary Table  2 ). The early Javanese H. erectus dental sample examined in this report is from Sangiran, Central Java. We divided this sample into two chronological subsamples, Sangiran Lower and Sangiran Upper, following the previous report that demonstrated significant morphological differences in tooth size, mandible features, cranial capacity, etc. 61 , 62 . Linear measurements were taken based on the original specimens or high-quality casts by Y.K. using a digital caliper (Mitsutoyo Inc.) or otherwise collected from the literature (Supplementary Table  2 ). Occlusal crown contours of SOA-MM11 was further analyzed by normalized elliptic Fourier analysis (size-standardized EFA), using the comparative samples shown in Supplementary Table  2 and based on the methods detailed elsewhere 37 . In brief, the occlusal contour of each specimen was obtained from a photograph of the original specimen or high-quality cast, in a way which minimizes the error derived from parallax effect and orientation of the tooth or scale. Images were uploaded into Canvas X software (ACD Systems) to extract the occlusal contour and, for a worn tooth, to reconstruct small parts of the crown lost by interproximal wear. Then, normalized elliptic Fourier analysis was conducted using the software SHAPE 1.3 63 , after each crown contour was aligned along its mesiodistal axis. We did not assess sex for these materials because of the small sample and the reported low sexual dimorphism in modern human deciduous teeth 64 .

Reporting summary

Further information on research design is available in the  Nature Portfolio Reporting Summary linked to this article.

Data availability

All data generated or analyzed during this study are included in this published article (and its supplementary information files) or as a Source Data file. The Source Data file includes raw data used for Figs.  6 and 7 , and Supplementary Figs.  2 , 3 , 4 and 6 . The Mata Menge hominin fossils are housed at the Geological Museum, Bandung. The 3D data of SOA-MM9 humerus may be shared on request to Unggul P. Wibowo ([email protected]).  Source data are provided in this paper.

Brown, P. et al. A new small-bodied hominin from the late Pleistocene of Flores, Indonesia. Nature 431 , 1055–1061 (2004).

Article   ADS   CAS   PubMed   Google Scholar  

Morwood, M. J. et al. Further evidence for small-bodied hominins from the late Pleistocene of Flores, Indonesia. Nature 437 , 1012–1017 (2005).

Detroit, F. et al. A new species of Homo from the Late Pleistocene of the Philippines. Nature 568 , 181–186 (2019).

Aiello, L. C. Handbo ok of Pa leoan thropology (eds W. Henke & I. Tattersall) 2281–2297 (Springer, 2014).

Brumm, A. et al. Age and context of the oldest known hominin fossils from Flores. Nature 534 , 249–253 (2016).

van den Bergh, G. D. et al. An integrative geochronological framework for the Pleistocene So’a basin (Flores, Indonesia), and its implications for faunal turnover and hominin arrival. Quat. Sci. Rev. 294 , 107721 (2022).

Article   Google Scholar  

Brumm, A. et al. Hominins on Flores, Indonesia, by one million years ago. Nature 464 , 748–752 (2010).

Brumm, A. et al. Stone technology at the Middle Pleistocene site of Mata Menge, Flores, Indonesia. J. Archaeol. Sci. 37 , 451–473 (2010).

van den Bergh, G. D. et al. Homo floresiensis -like fossils from the early Middle Pleistocene of Flores. Nature 534 , 245–248 (2016).

Article   ADS   PubMed   Google Scholar  

Sutikna, T. et al. Revised stratigraphy and chronology for Homo floresiensis at Liang Bua in Indonesia. Nature 532 , 366–369 (2016).

Matsu’ura, S. et al. Age control of the first appearance datum for Javanese Homo erectus in the Sangiran area. Science 367 , 210–214 (2020).

Argue, D., Groves, C. P., Lee, M. S. Y. & Jungers, W. L. The affinities of Homo floresiensis based on phylogenetic analyses of cranial, dental, and postcranial characters. J. Hum. Evol. 107 , 107–133 (2017).

Article   PubMed   Google Scholar  

Dembo, M., Matzke, N. J. & Mooers, A. O. & Collard, M. Bayesian analysis of a morphological supermatrix sheds light on controversial fossil hominin relationships. Proc. Biol. Sci. 282 , 20150943 (2015).

PubMed   PubMed Central   Google Scholar  

Dembo, M. et al. The evolutionary relationships and age of Homo naledi : an assessment using dated Bayesian phylogenetic methods. J. Hum. Evol. 97 , 17–26 (2016).

Yurnaldi, D., Setiawan, R. & Patriani, E. Y. The magnetostratigraphy and the age of So’a basin fossil-bearing sequence, Flores, Indonesia. Indones. J. Geosci. 5 , 221–234 (2018).

Powley, M. J., Sutisna, I., Mikac, K. M., Wibowo, U. P. & van den Bergh, G. D. The stegodon bonebed of the Middle Pleistocene archaeological site Mata Menge (Flores, Indonesia): taphonomic agents in site formation. Quaternary 4 , 31 (2021).

Gocha, T. P., Robling, A. G. & Stout, S. D. Biological Anthropology of the Human Skeleton , 3rd Edition (eds M. A. Katenberg & A. L. Grauer) 145–187 (John Wiley & Sons, 2018 ) .

Martínez-Maza, C., Rosas, A. & García-Vargas, S. Bone paleohistology and human evolution. J. Anthropol. Sci. 84 , 33–52 (2006).

Google Scholar  

Wu, K., Schubeck, K. E., Frost, H. M. & Villanueva, A. Haversian bone formation rates determined by a new method in a mastodon, and in human diabetes mellitus and osteoporosis. Calcif. Tissue Res. 6 , 204–219 (1970).

Article   CAS   PubMed   Google Scholar  

Kerley, E. R. The microscopic determination of age in human bone. Am. J. Phys. Anthropol. 23 , 149–163 (1965).

Gocha, T. P. & Agnew, A. M. Spatial variation in osteon population density at the human femoral midshaft: histomorphometric adaptations to habitual load environment. J. Anat. 228 , 733–745 (2016).

Schultz, M. Forensic Taphonomy: The Postmortem Fate of Human Remains (eds W. D. Haglund & M. H. Sorg) 201–222 (CRC Press, 1997).

Carroll, S. E. A study on the nutrient foramina on the humeral diaphysis. J. Bone Jt. Surg. 45B , 176–181 (1963).

Larson, S. G. et al. Descriptions of the upper limb skeleton of Homo. floresiensis. J. Hum. Evol. 57 , 555–570 (2009).

Walker, A. & Leakey, R. The Nariokotome Homo erectus Skeleton (eds A. Walker & R. Leakey) 7 , 95–160 (Harvard University Press, 1993).

Churchill, S. E. et al. The shoulder, arm, and forearm of Australopithecus sediba. PaleoAnthropology 2018 , 234-281 (2018).

Ward, C. V., Kimbel, W. H., Harmon, E. H. & Johanson, D. C. New postcranial fossils of Australopithecus afarensis from Hadar, Ethiopia (1990-2007). J. Hum. Evol. 63 , 1–51 (2012).

White, T. D. et al. New discoveries of Australopithecus at Maka in Ethiopia. Nature 366 , 261–265 (1993).

Lague, M. R. et al. Cross-sectional properties of the humeral diaphysis of Paranthropus boisei: Implications for upper limb function. J. Hum. Evol. 126 , 51–70 (2019).

Grine, F. E. Comparison of the deciduous dentition of African and Asian hominids. Cour. Forsch. Senckenberg 69 , 69–82 (1984).

Jacob, T. et al. Pygmoid Australomelanesian Homo sapiens skeletal remains from Liang Bua, Flores: population affinities and pathological abnormalities. Proc. Natl Acad. Sci. Usa. 103 , 13421–13426 (2006).

Article   ADS   CAS   PubMed   PubMed Central   Google Scholar  

Kaifu, Y. et al. Descriptions of the dental remains of Homo floresiensis . Anthropol. Sci. 123 , 129–145 (2015).

Tobias, P.V. Olduvai Gorge, 4: the skulls, endocasts and teeth of Homo habilis . 4 (Cambridge University Press, 1991).

Wood, B. Koobi Fora Research Project 4: Hominid Cranial Remains . 4 (Clarendon Press, 1991).

Kaifu, Y., Aziz, F. & Baba, H. Hominid mandibular remains from Sangiran: 1952–1986 collection. Am. J. Phys. Anthropol. 128 , 497–519 (2005).

Hens, S. M., Konigsberg, L. W. & Jungers, W. L. Estimating stature in fossil hominids: which regression model and reference sample to use? J. Hum. Evol. 38 , 767–784 (2000).

Kaifu, Y. et al. Unique dental morphology of Homo floresiensis and its evolutionary implications. PLoS ONE 10 , e0141614 (2015).

Article   PubMed   PubMed Central   Google Scholar  

Lyras, G. A., Dermitzakis, M. D., Van Der Geer, A. A. E., Van Der Geer, S. B. & De Vos, J. The origin of Homo floresiensis and its relation to evolutionary processes under isolation. Anthropol. Sci. 117 , 33–43 (2009).

Gordon, A. D., Nevell, L. & Wood, B. The Homo floresiensis cranium (LB1): size, scaling, and early Homo affinities. Proc. Natl Acad. Sci. Usa. 105 , 4650–4655 (2008).

Kaifu, Y. et al. Craniofacial morphology of Homo floresiensis : description, taxonomic affinities, and evolutionary implication. J. Hum. Evol. 61 , 644–682 (2011).

Baab, K. L., McNulty, K. P. & Harvati, K. Homo floresiensis contextualized: a geometric morphometric comparative analysis of fossil and pathological human samples. PLoS One 8 , e69119 (2013).

Schroeder, L. et al. Skull diversity in the Homo lineage and the relative position of Homo . naledi. J. Hum. Evol. 104 , 124–135 (2017).

Kubo, D., Kono, R. T. & Kaifu, Y. Brain size of Homo floresiensis and its evolutionary implications. Proc. R. Soc. B: Biol. Sci. 280 , 20130338 (2013).

Lague, M. R. Taxonomic identification of Lower Pleistocene fossil hominins based on distal humeral diaphyseal cross-sectional shape. PeerJ 3 , e1084 (2015).

Lague, M. R. et al. Humeral anatomy of the KNM-ER 47000 upper limb skeleton from Ileret, Kenya: Implications for taxonomic identification. J. Hum. Evol. 126 , 24–38 (2019).

Lague, M. R. & Menter, C. G. Hominin Postcranial Remains from Sterkfontein, South Africa (eds C. V. Ward & B. Zipfel) (Oxford University Press, 2020).

Jee, W. S. S. Cell a nd Ti ssue Bio logy , 4th Edition (ed. L. Weiss) 212–254 (Urban and Schwarzenberg, 1983).

Thompson, D. D. The core technique in the determination of age at death in skeletons. J. Forensic Sci. 24 , 902–915 (1979).

Ericksen, M. F. Histologic estimation of age at death using the anterior cortex of the femur. Am. J. Phys. Anthropol. 84 , 171–179 (1991).

Stout, S. D. & Paine, R. R. Brief communication: histological age estimation using rib and clavicle. Am. J. Phys. Anthropol. 87 , 111–115 (1992).

Pfeiffer, S. Variability in osteon size in recent human populations. Am. J. Phys. Anthropol. 106 , 219–227 (1998).

Cunningham, C., Scheuer, L. & Black, S. Developmental Juvenile Osteology , 2nd Edition (Academic Press, 2016).

Dempster, D. W. et al. Standardized nomenclature, symbols, and units for bone histomorphometry: a 2012 update of the report of the ASBMR histomorphometry. Nomenclature Comm. J. Bone Miner. Res. 28 , 2–17 (2013).

Senut, B. Humeral outlines in some hominoid primates and in polio-Pleistocene hominids. Am. J. Phys. Anthropol. 56 , 275–283 (1981).

Susman, R. L., de Ruiter, D. & Brain, C. K. Recently identified postcranial remains of Paranthropus and early Homo from Swartkrans Cave, South Africa. J. Hum. Evol. 41 , 607–629 (2001).

De Lumley, M. A. & Marchal, F. Les sites préhistoriques de la région de Fejej, Sud-Omo, Ethiopie, dans leur contexte stratigraphique et paléontologique (eds H. de Lumley & Y. Beyene) 340–383 (Editions Recherche sur les Civilisations, 2004).

Rohlf, F. J. & Slice, D. E. Extensions of the Procrustes method for the optimal superimposition of landmarks. Syst. Zool. 39 , 40–59 (1990).

Rohlf, F. J. tpsRelw, relative warps analysis. version 1.49 ed. Department of Ecology and Evolution ( State University of New York at Stony Brook, 2010).

Bookstein, F. L. Morphometric Tools for Landmark Data: Geometry and Biology . (Cambridge University Press, 1997).

Rohlf, F. J. Shape statistics: procrustes superimpositions and tangent spaces. J. Classif. 16 , 197–223 (1999).

Kaifu, Y. et al. Taxonomic affinities and evolutionary history of the early Pleistocene hominids of Java: dentognathic evidence. Am. J. Phys. Anthropol. 128 , 709–726 (2005).

Kaifu, Y., Indriati, E., Aziz, F., Kurniawan, I. & Baba, H. Asian Paleoanthropology: From Africa to China and Beyond (eds C. J. Norton & D. R. Braun) 143–157 (Springer, 2010).

Iwata, H. & Ukai, Y. SHAPE: a computer program package for quantitative evaluation of biological shapes based on elliptic Fourier descriptors. J. Hered. 93 , 384–385 (2002).

Gil-Donoso, E. et al. Sexual dimorphism of deciduous canine dental tissues dimensions of modern human populations. Anthropol. Sci. 131 , 107–115 (2023).

Ubelaker, D. H. Human Skeletal Remains: Excavation, Analysis, Interpretation . 2nd edn (Taraxacum, 1989).

P, S., Bhavyaa, R., M, S. M., Nirmal, L. & Patil, S. S. Crown dimensions of primary teeth-A systematic review and meta-analysis. J. Forensic Sci. 67 , 1348–1356 (2022).

Ryan, W. B. F. et al. Global Multi-Resolution Topography (GMRT) synthesis data set. Geochem. Geophys. Geosyst. 10 , Q03014 (2009).

Article   ADS   Google Scholar  

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Acknowledgements

The former head of the Geological Agency, Banding, Dr. Surono, is acknowledged for his support. Geodetic recordings of excavated finds were conducted by Y. Sopyan (2013), and E.E. Laksmana and A. Rahmadi (2014). We thank the people from Mengeruda and Piga villages for their participation in the excavations and their ongoing support. We also thank the following persons for their contributions in the field during the 2013–2016 Mata Menge excavations: T. Suryana, E. Sukandar, A. Gunawan, Widji, A.T. Hascaryo, E. Setiyabudi, A.M. Saiful, B. Burhan, P.D. Moi, B. Alloway, B. Pillans, M. Storey, D. Yurnaldi, M. Moore, T. Sutikna, H. Insani, M.R.Puspaningrum, I. Yoga, H.J.M. Meijer, S. Hayes, and F. Aziz. We thank Takao Sato and Takashi Sano for their assistance in conducting morphological analyses, and J.M. Plavcan, C. Ward, M. Domínguez-Rodrigo, F. Di Vincenzo, and W. Kimbel for 3D scans and/or casts of fossil humeri. We are indebted to the following institutions: Pusat Penelitian Arkeologi Nasional, Senckenberg Research Institute and Natural History Museum Frankfurt, American Museum of Natural History, Sapporo Medical University, Niigata University, National Museum of Nature and Science, Tokyo, The University of Tokyo, St. Marianna University School of Medicine, Tahara Municipal Museum, Kyoto University, Kyushu University, Sasebo City Museum Shimanose Art Center, Okinawa Prefectural Museum and Art Museum, Okinawa Prefectural Archeology Center. Aspects of this research were financially supported by Australian Research Council Discovery grant (DP1093342 to the late M.J.M. Morwood and A.B.), Australian Research Council Future Fellowship (FT100100384 to G.D.vd.B.), Center for Geological Survey Bandung, Indonesia, Geology Museum Bandung, Indonesia (to I.K., R.S., I.S. and U.P.W.), JSPS KAKENHI grant (22H00421 to Y.K. and 23K17521 to J.S.) and National Science Foundation (BCS-0647557 to M.L.).

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Y.K., G.D.v.d.B., and I.K. conceived and led the study. Excavations were led by I.K. and R.S., with assistance from I.S. U.P.W. A.B. and G.D.v.d.B. Morphological analyses were conducted by Y.K., S.M., J.S., and M.L. with assistance from G.S., R.T.K. and T.S. Y.K., S.M., J.S., and M.L. wrote the manuscript with editorial inputs from all co-authors.

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Kaifu, Y., Kurniawan, I., Mizushima, S. et al. Early evolution of small body size in Homo floresiensis . Nat Commun 15 , 6381 (2024). https://doi.org/10.1038/s41467-024-50649-7

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