Course:ARCL140 Summer2020/TermProject Group26

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An Exploration on Pleistocene Non-Sapien Hominins

Contributors and Roles :

Zia Chapman : Rising Star Cave, South Africa

Caye Hing : Liang Bua Cave, Flores Island, Indonesia

Rory Mills : Neander Valley, Germany

Natalie Urquhart : Denisovan Cave, Altai Mountains, Siberia, Russia


A screenshot of our Google MyMaps with pinned locations: Germany, South Africa, Indonesia and Siberia

Google MyMaps link:


The evolutionary journey of the genus Homo, as we have examined, is not a straightforward nor linear process. As we continue to learn more about our genus, we are left puzzled by the sequence and timing of our origin. The earth from a millenia ago was walked by multiple species of humans, similar to how we see multiple species of animals today. Can we imagine a modern world if we live alongside Neanderthals and Denisovans? Is there a secret to our exclusivity in becoming the last remaining human species? In an effort to seek the answer to our particular exclusivity, we have designed this research project to shine the spotlight on our distant cousins, particularly the Denisovans, Homo floresiensis, Neanderthals and Homo naledi. The study of these numerous non-sapien hominins provides us insight into their unique lifeways. Further, we will be exploring themes such as morphological change due to selective pressures, biogeographical demands as well as behavioral and reproductive inclinations. The sites covered within this project range are respectively in Southeast Asia, Eurasia, Siberia and South Africa, with each individual site providing a unique perspective on human evolution.

To start off, the skeletal specimens of the Neanderthals within the small valley of Düssel in Germany was the first time that humans identified other non-sapien hominins. This discovery resulted in a widespread discussion on their role in human evolutionary history, which kickstarted the conversation about other non-sapien hominins (Schmitz et al., 13342).

In South Africa, Homo Naledi illustrates the phenomenon of mixed physical features, particularly shared features of Autralopithecus and modern features. This can be seen in their transitional form with humanlike body size, hands and wrists and an Australopith-sized brain and hip (Berger et al. 2015, 23).

In Flores, the evolution of Homo floresiensis serves as a primary example of how long-term isolation in an enclosed environment, like an island can cause nutritional deficiencies, which in the long run causes insular dwarfism. This exhibits how environmental pressures induces phylogenetic modification (Bergh et al. 2009, 535).

Finally, a Siberian cave provides the first instance of a human hybrid with a half Denisovan, half Neanderthal female child (Paskey& Cisneros 2019, 26). This case of interbreeding illustrates the process through which Denisovan adaptations-- such as hypoxia immunity--have passed down to modern humans in the area.

As we will see shortly, through careful examination of various sites, we will try to entangle this complex evolutionary web and iron out the famous fallacy of a linear model of evolution. Furthermore, we will delve into the vast diversity of our genus and see how geographical differences led to very different lifestyles: how some were relentless hunters while others were more docile and how some lived on a single island while others roamed continents.

SITE 1: Liang Bua Cave, Flores Island, Indonesia

Liang Bua Cave, Flores, Indonesia


Caye Hing


Coordinates: 8.5450° S, 120.4300° E

Liang Bua Cave is 7 km northwest of Ruteng, the regional capital of Manggarai Regency, west Flores. Flores sits east of Sumbawa and Komodo islands and west of Lembata island and the Alor Archipelago (Morwood et al. 2009, 441).


Based on stratified deposits of stone artifacts and faunal remains, Liang Bua Cave already existed approximately 900,000 years ago, around the Late Pleistocene or potentially even earlier. Homo skeletons, however, have been identified to be spanning the last 95,000 years.


Liang Bua is a limestone cave on the island of Flores, East Indonesia. Flores is part of Wallacea, the group of islands in between the Australian and Asian continental shelves and dates all the way back to the Late and Middle Pleistocene epoch. It has a (current period) land area of 5,230 square miles and is the last considerable island in the chain of the Lesser Sunda Islands in East Nusa Tenggara (Bergh et al. 2009, 527). During glacial periods, when sea levels dropped, the Sunda shelf to the west connected a land bridge to mainland Asia, however, due to present sea levels, anything east of the continental shelf, including Flores (which originally arose from the ocean floor) remains ceaselessly enclosed by water (Morwood et al. 2009, 438).

Renowned for its numerous inlets and bays, Flores’ ecology is largely mountainous, particularly in the west, where Mandasawu Peak reaches 7,900 feet and is considered the highest point of the island. The rivers in the island are for the most part unnavigable and several active volcanoes are present in the centre and east of Flores. Moreover, the island is filled with a lush ecoregion of flora and fauna which consists mostly of moist deciduous, dry thorn, and dry evergreen forests. The fauna includes a large diversity of species endemic to the island which include the Komodo Dragon, the world’s largest lizard and mammals like the Flores shrew, (Bergh et al. 2009, 533).


Between 1950 to 1960, a Dutch priest, Theodor Verhoeven carried out excavations and collections at many limestone rock reserves and caves in Flores. During the latter half of the 20th century, the next phase of archaeological and palaeontological research on Flores were spearheaded by both local and international teams in an effort to collect sediments from rivers and lakes, which they later established as deposits laid down during the Paleolithic Asian settlement (Westaway et al. 2007, 338).

Archaeological excavations in Flores have yielded a plethora of stone artifacts and faunal remains spanning the last 95,000 years, including the skeletal remains of two human species, Homo floresiensis (discovered in 2003) in the Pleistocene and Homo sapiens in the Holocene (Morwood and Jungers 2009, 640). In 2010 and 2011, archaeologists discovered two hominin teeth in the cave dating to around 46,000 BP, identified as belonging to Homo sapiens.


The Liang Bua Cave team discovered sub-fossil remains, coded LB1, LB2 which were later identified to be a new species of Homo, Homo floresiensis. They have so far only been found in Flores and are widely speculated to have become the Indonesian descendants of Homo erectus immigrants who crossed miles of ocean to colonize their new tropical home. In this new island habitat, H. floresiensis eventually underwent insular dwarfing and stood only at approximately 3 feet 6 inches tall. Despite their small body and brain size, H. floresiensis made and used stone tools, hunted, were speculated to have utilized fire and put up with predators like the giant Komodo dragons.

File:Homo floresiensis bust.jpg
A recreation of Homo floresiensis

Morwood and Jungers (2009) gathered that they show a mix of Australopiths-like ancient features including wide hips, a short collarbone and forward-positioned shoulder yet more contemporary features as well like brow ridges, skull thickness and brain shape resembling H. erectus and later species (643).

The excavated stone tools were manufactured predominantly from volcanic rocks, intentionally fractured to have sharp edges for slicing and dicing (Morwood et al. 2009, 440).These tools resemble the Oldowan technology which was predominant in Africa about 2.6 million years ago.

Many combined subfields of science have been able to identify a faunal assemblage through fossil remains, illustrating island gigantism in small mammals and the dwarfing of large taxa. According to Bergh et al. (2009), together with evidence from Middle-Late Pleistocene sites in Flores, the discovery of the “hobbit" has confirmed that long-term isolation, nutritional privation, and environmental pressures cause phylogenetic continuity, ultimately altering a single species and even an entire faunal community (535).

SITE 2: Denisova Cave, Altai Mountains, Siberia, Russia


Denisova Cave viewed from outside

Natalie Urquhart


Coordinates: 51.3975° N, 84.6761° E

The Denisova cave lies in Bashelaksky Range of the Altai mountains, Siberia, Russia. The Altai mountain range crosses from central to East Asia at the intersection of Russia, China, Mongolia, and Kazakhstan. The cave itself is located in Altai Krai, near the village of Chorny Anui. It sits 150 km South of the regional capital Barnaul and 28 m above the Anuy River on the right side.


The bone fragments, tools attributed to Denisovans, and various other decorative artifacts date to around 40,000 BP. Therefore researchers have situated the remnants in the cave from the Middle Palaeolithic period-- about 340,000 to 45,000 years ago-- to the Initial Upper Palaeolithic period, 30,000 to 48,000 years ago (Dennell 2019, 571).


The Denisova cave was originally named by the Russians after a hermit that inhabited it in the late 18th century, Saint Denis. The locals however, call it Aju-Tasch, meaning “Bear rock”. The hermit Denis was the last person known to inhabit the cave, but before him came the Denisovans of whom evidence dates back to at least 48 kya, though it is possible that they lived around there more than 200,000 years ago (Paskey&Cisneros 2019, 23).

This cave in the Altai mountains sits at more than a kilometre above sea level and has an annual temperature close to 0˚C. For most modern humans, hypothermia, altitude sickness, and hypoxia would become prevalent issues in long term inhabitation. Denisovans however, seem to have had multiple genetic adaptations which prevented such illnesses. Through interbreeding 30 kya to 40 kya, the Denisova have passed these adaptations down to modern Tibetans (Paskey&Cisneros 2019, 19).

This cave consists of three chambers, known as the Main, East, and South chambers (Douka 2019, 640).


Photograph of the 2 cm bone (Denisova 11) found in the Denisova caves, published in Scientific Reports in 2016

The first archeological studies of the cave began in the 1970’s when Russian scientists uncovered paleoarcheological remains. This discovery led to continued explorations, which have thus far found 22 strata layers containing artifacts ranging all the way back to approximately 180 kya. These strata have been dated using radiocarbon dating on charcoal, bone, and tooth as well as thermoluminescence dating on sediments (Oxford University, n.d).

The site garnered the most attention only recently in 2010 when the finger bone of a young girl and a large molar were found. Before this, the bones found had been from an undetermined archaic human group— the 2010 research team then naming them the Denisovans (Gibbons 2011, 1084). The remnants of 12 hominins in total have been found at the site so far, most of them fragmentary (Douka 2019, 640). However, even with the small and fragile state of these remains, many of them have well preserved DNA.


Denisova cave quickly became an important archaeological site in 2010 when the finger bone of a 13 year old female was found. Not only did this phalanx allow for the confirmation of a new ancient hominid group, but it was also the discovery of the first recorded human hybrid (Oxford University, n.d). This young girl was the product of a Neanderthal mother and a Denisovan father (Paskey&Cisneros 2019, 26).

The finger also provided a large sample of aDNA for study and, with its extremely distinct mtDNA sequence, researchers were able to determine that Denisovans diverged from modern humans and Neanderthals approximately one million years ago. However, soon after said research was published, contradicting evidence in the nuclear DNA of the Denisovans showed that they were actually a sister taxon of Neanderthals (Stringer& Barnes 2015). As such, Neanderthals and Denisovans may have only diverged between 381kya to 473kya (Paskey&Cisneros 2019, 23). Even after this divergence however, the existence of the hybrid girl confirms that the groups still intermingled.

We can be certain that Neanderthals only visited the area in the summer though (Oxford University, n.d) because they did not have the genetic adaptations required to survive in the area year round. To inhabit the Altai mountains year round, with the temperature so low and the elevation high, Denisovans developed adaptations which prevented hypoxia and altitude sickness. While there is no evidence of modern homo sapiens inhabiting the area at the time of the Denisovans, interbreeding amongst the Denisovans and other hominins 30,000 to 40,000 years ago has equipped modern Tibetans with similar modifications (Paskey&Cisneros 2019, 19).

SITE 3: Neander Valley, Germany

Fuhlrott's drawings of the Kleine Feldhofer Grotte

Rory Mills

Skullcap of Neanderthal 1 found in 1856


The Neander Valley is small valley of the Düssel River in the German state of North-Rhine-Westphalia, 12 km East of Düsseldorf. Originally a limestone canyon, extensive quarrying in the 19th and 20th centuries removed most of the limestone, including the Kleine Feldhofer Grotte, the original cave in which the Neanderthal bones initially resided (Trinkaus, 2008, 49).

Coordinates: 51.2206°N, 6.9756°SCoordinates: 51.2206° N, 6.9756° S


Radiocarbon dating of the original Neanderthal 1 remains indicates an age of ~40,000 years, identified through a mtDNA sequence in 1997 (Schmitz et al. 2002, 13342). This specimen was cross-referenced with a second individual yielded from further digs, shows a dating age of a similar range, along with sequences that cluster with other published Neanderthal sequences (Schmitz et al.,13342).


Originally known as “Hundsklipp,” or, “Gesteins,” the valley became known by about 1850 as the Neander Valley in honour of Joachim Neander, a local teacher and poet (Schmitz et al.,13342). The Kleine Feldhofer Grotte specifically was named because of its proximity to the nearby farm of Feldhof (Schmitz et al., 13342). Rich in limestone, the Neander valley hosted several quarries in the 1860s to supply local construction projects, the process by which the Kleine Feldhofer Grotte was unearthed (King, 1864, 88).


In August 1856, several bones were discovered during quarry work in the Feldhofer grotto. Workers dismissed the bones as belonging to a cave bear, but the co-owner of the quarry, Wilhelm Beckershoff noticed the bones and identified them as human (Trinkaus, 49).

The find was first reported scientifically by Fuhlrott and Schaaffhausen in 1857, was the subject of a detailed analysis by Schaaffhausen, becoming a focal point in debates of evolution and human anthropology in the late 19th and early 20th centuries (Schmitz et al., 13342). The initial discovery yielded a total of 16 bones, 15 postcranial, and one skull cap (King, 89).

Later digs in 1997 and 2000 digs yielded in total, 67 new fragments of human bones, and several fauna artifacts, with in addition to new pieces for Neanderthal 1, identifies at least one other adult, and one other subadult (Schmitz et al., 13343). However, more extensive possibilities of scientific discovery only become apparent after the 1997 and 2000 excavations. Providing evidence of Middle Palaeolithic tools and Late Pleistocene- age fauna as well as a direct AMS date of 40,000 years establishes, to our knowledge for the first time, an appropriate context for Neanderthal 1 (Ralf, 13347). With the recovery of the Neanderthal remains in 1856 being only a small piece of the possible artifacts not yet excavated at this site, key insights to the early palaeontologists of the 19th century were missed (Schmitz et al., 13342).

The skeletal specimens of Neanderthal 1 discovered in the contemporary digs exhibit anatomical features consistent with other Neanderthals, and significantly enhanced anatomical knowledge of the Neanderthal specimen. Other remains from this site also appear to derive morphology from Neanderthals. Access to more Neanderthal samples provides the opportunity to further investigate anatomical variation of the Neanderthal specimens (Schmitz et al., 13342).


The 1856 discovery of Neandertal 1 in western Germany marked the initiation of human palaeontology and ignited one of the fiercest debates in burgeoning field: the role of Neanderthals in human evolutionary history (Schmitz et al.,13347). The unfortunate circumstance of the bones discovery means that scientists and intellectuals of the time could only discuss implications through Neanderthal 1’s skeletal anatomy, as no archaeological or faunal materials were reported from the site in 1856. As no careful surveying was done, the exact location of the Kleine Feldhofer Grotte was no longer known by 1900. With no associated finds, and the cave being destroyed without any scientific geological analysis, the site had been considered undatable until the 1997 and 2000 excavations. (Schmitz et al., 13347).

Questions concerning the antiquity of the original Neanderthal specimen led many to reject any role for it in human evolution and generally impeded late 19th century progress in the study of human evolution (Schmitz et al., 13347).

The mtDNA sourced from one of second Neanderthal individuals also dates to 40,000 years ago, making this site the first Pleistocene dated location to yield sequences from more than one hominin. This allows for not only morphological, but also molecular genetics of the Neanderthal population before any credible contact with modern Sapiens in Europe (Schmitz et al.,13342).

SITE 4: Rising Star Cave, South Africa


Zia Chapman


Coordinates: 25.9254° S, 27.7674° E

The Rising Star Cave is located 1450-1480 m above sea-level in the Bloubank River valley, positioned within the Cradle of Humankind (CoH) which is a UNESCO World Heritage Site in South Africa and is approximately 2km west of the Sterkfontein Caves and 35km northwest of Johannesburg (Wiersma et al. 2019, 899).


Within the caves of the CoH, fossil assemblages from the late Pliocene to early Pleistocene age have been found (Dirks et al. 2017, 3). The fairly recent discovery of the Homo naledi fossils, which is characteristic to the Rising Star Cave, are said to be between 236,000 and 335,000 years old. These dates that were estimated by the researchers involved in the discovery of H. naledi are much more recent than was predicated which seems to indicate that the species was alive around the same time as the earliest members of our species which evolved approximately 300,000 and 200,000 years ago (Dirks et al. 2017, 2).


Within the Rising Star Cave lies the Dinaledi Chamber that is contextually important to the discovery of the new species Homo naledi. The Chamber is characteristically a sedimentary environment that is geochemically distinct from the remaining parts of the Rising Star Cave (Dirks et al. 2017, 2). The hominin deposits in Cave of H. naledi are mainly unconsolidated, mud-clast breccia in a muddy medium but there are no signs of clastic sediment taken by water flow even though the Cave is situated in the Bloubank River Valley (Dirks et al. 2017, 3). The unlitihified formation of the mud-clast breccia deposit is gradual and the first formation is approximately ca 8 x 10-4 mm year-1. Due to this gradual process, it seems to indicate that the Chamber has had dry and stable conditions for the last ca 300ka (Wiersma et al. 2019, 897).

Two team members out of the six cavers that collected the 1500 fossils from the Rising Star Cave, South Africa.


A reconstructed face of the Homo naledi

The Rising Star Cave was only seriously investigated in September 2013 by an excavation team under American paleo-anthropologist, Lee Rogers Berger (Kruger et al. 2016, 11). After determining the potential of the site and acknowledging the difficulties associated with accessing the Dinaledi Chamber due to its incredibly narrow width, Berger sent out a request for qualified archaeologists that were "skinny and preferably small...[and] must not be claustrophobic" (Leakey Foundation, episode 26). On receiving a much larger response than anticipated, he invited a team of international women joined the excavation process and in October 2013, the fossil hominins were first recognised (Berger et al. 2015, 2).


The Rising Star Cave was periodically visited by several amateur cavers for several years, but it was only in September 2013 that a caving team from the Evolutionary Studies Institute formally investigated the system for the first time to seek out fossil remains of early hominins. Between 2013-2014, the excavators yielded 1550 identifiable fossil elements that represented at least 15 individuals and several examples of a large amount of the bones in the skeleton. This collection is the largest of fossil hominin material found on the African continent to this date and the research team assigned them to a new species of the de novo hominin taxon - the Homo naledi (Kruger et al. 2016, 11).

The name came about as the word naledi means 'star' in the Sotho language (one of the 11 official languages of South Africa) which alludes to the Dinaledi Chamber's location in the Rising Star Cave system (Berger et al. 2015, 3). Anatomically, the H. naledi shows some shared features with Australopithecus, some shared features with Homo, and some that have not been seen in any hominin species prior to this (Berger et al. 2015, 17). Specifically, the species exhibits a combination of a human like body build and size with an australopith-sized brain as well as the shoulders and hands adapted to climbing, with humanlike hands and wrist adaptations. They also have australopith-like hip mechanics with humanlike terrestrial adaptations of the foot and lower limb (Berger et al. 2015, 23). The evidence of traits seem to indicate adaptations for creating material culture and the significance of the enormous amount of fossils within the Dinaledi Chamber have left paleo-anthropoligists with the question of whether it was a burial site for this species. Although this is still being debated, it is mostly agreed that such cultural behaviour was performed within the capabilities of the species (Berger et al. 2017, 12). The discovery of the H. naledi fossils within the Dinaledi chamber has been an incredible representation of skeletal elements across the lifespan in the hominin fossil record. While more analytical techniques and an understanding of aDNA is required to provide clarity on the several questions surrounding the H. naledi, the discovery of this large assemblage of fossils has strongly supported the development of an understanding that the genus Homo underwent a wide spectrum of evolutionary experiments (Berger et al. 2015, 24).


As we have outlined the sheer archaeological and paleo-anthropological potential of the four location sites above, the respective discoveries from each site indicate a closer relationship to the wider picture of human evolution. Thus, we conclude on a number of things: Firstly, that human evolution is not a linear process but instead happened mosaically or in a patchy, non-simultaneous manner. Secondly, the study of human evolution has been shaped by the philosophies of the times of their discoveries, and the subsequent scientific assumptions that were derived from incomplete archeological studies, or limited access to technologies. Thirdly, geographical variation and selective pressures play an unequivocal role in how we have evolved, influencing morphological traits and phylogenetic continuity. And lastly, through excavated stone tools and faunal assemblages, we are able to establish that a potential material culture existed amongst these hominin species, particularly through how they handcrafted non-biological instruments that aided in their overall survival.

Archaeological discoveries and faunal impacts led to the discoveries of Homo floresiensis in Southeast Asia, Neanderthals in Eurasia, Denisovans in Siberia and Homo Naledi in South Africa. All these pose significant implications that have contributed key factors about hominin evolution, reproduction, behaviour and ecology and how each respective species came to adapt, through time, to their environment. Denisovans, for example, provide the first known instance of a human hybrid-- the byproduct of interbreeding between various species. Additionally, as with the case of Homo naledi, this indicated the possible adaptations for creating material culture as seen through possible mortuary behaviour. Moreover, this not only elucidates upon archaic hominin evolution in the Pleistocene but also provides further evidence about the nature and timing of early hominin biogeography and dispersal, captivating global regard. As well, early to contemporary archeological techniques and methodologies help us continue to redefine our story of origin. The social impact of these burgeoning discoveries reflects upon the endless possibilities that Homo Sapiens pose in the modern world. Further, Pleistocene hominin skeletal and stone tool evidence from these respective sites infer paradigm-shifting implications for early hominin material culture, biogeography and occupation in Asia, Europe and Africa more generally. Conclusively, this makes us rethink and re-evaluate our evolution as a human species and consider how the story of evolution is an active effort by many subfields of science. Learning about our genus Homo's story is a comparative, interdisciplinary endeavour that continues to hold much more unanswered questions for the field of archaeology, paleoanthropology and overall, world prehistory.


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