Are Denisovans Able to Cross the Wallace’s Line?

Author : Wahid Ahmad

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Early modern humans departing Africa likely made their initial stop in India, where the first diversifications of mitochondrial and Y chromosome lineages outside Africa occurred. From India, one group moved eastward to Southeast Asia. This coastal migration facilitated the exploration of the lost continents of Sunda and Sahul.

 

During periods of low sea levels in the past, Southeast Asian regions, including islands like Sumatra, Java, Bali, and Borneo, formed an extended landmass known as Sundaland or Sunda. This landmass was linked by substantial land bridges during the last glacial maximum, around twenty thousand years ago.

Over time, Sundaland experienced sea-level rises resulting in a sea-level increase of one hundred and twenty meters that shaped the present-day geographical layout of Southeast Asia. Sahul comprised regions like Australia, New Guinea, and Tasmania, connected during the last glacial maximum due to lower sea levels.

 

Named after British naturalist Alfred Russel Wallace, Wallacean is a transitional biogeographic region located between Sunda and Sahul. It is bounded by two key biogeographic lines: Wallace’s Line to the west and Lydekker’s Line to the east.

Wallacean comprises a collection of islands, including Sulawesi, Lombok, Flores, Timor, and the Moluccas, and is characterized by its distinct flora, fauna, and geological history.

The Wallacean islands contain a mix of animals from both regions, including marsupials and placental mammals, each island having its own unique combination of species. To the west of Wallace’s Line, ecosystems are dominated by placental mammals, while to the east near Australasia, ecosystems are mainly marsupial-dominated.

 

The separation of the Sunda and Sahul landmasses by sixty miles of Wallace’s Line led to the independent evolution of Asian and Australian fauna. Asia developed placental mammals, while Sahul hosted marsupials.

 

The island of Flores revealed evidence of Stegodon elephants and Homo floresiensis. Besides Homo floresiensis, Southeast Asia’s human habitation history of nearly two million years includes Homo erectus and potentially Denisovans. Anatomically modern humans reached Australia through ancient seafaring, supported by archaeological finds.

The earliest human occupation on Sahul offers indirect evidence of advanced seafaring skills. Crossing the 60-mile-wide channel between Sunda and Sahul required intentional voyaging and likely guidance from indicators like forest fires or bird migrations. 

Around 50,000 to 65,000 years ago, modern humans spread rapidly through Southeast Asia due to lower sea levels. They crossed Wallacea's islands to reach Sahul.

 

The necessity of intentional voyaging is highlighted by the absence of archaic humans and large mammals on Sahul, suggesting directed efforts. This migration from Sunda to Sahul was a remarkable feat for several reasons. Firstly, it required advanced maritime skills to navigate the waters between islands.

Secondly, the migrants had to adapt to new environments, including unfamiliar plants and animals, as they settled on each new island or continent. Lastly, this migration occurred tens of thousands of years before the settlement of the Americas, making it an early and significant human movement. 

Considering factors like island distances, terrain, and sea crossings, two main migration routes have been suggested for the migration of early humans from the Sunda region to Sahul: the northern route and the southern route. The northern route followed the northern part of Wallacea. It was preferred due to shorter distances between islands and better visibility.

Around sixty-five thousand years ago, during lower sea levels, the Sunda coastline extended along Borneo's eastern side, reaching Java and Bali before curving west. The route likely began near the Balabalagan islands and extended to Misool Island. Potential paths included islands like Obi, Kofiau, Buru, Seram, Peleng, and Sula. This route involved three crossings after Sulawesi, each taking two to three days, with the destination island visible throughout the journey. The southern route involved longer, riskier voyages between Timor and Australia, requiring well-planned sea crossings. Islands on Sahul Banks were visible from high points on Timor and Roti.

Around seventy thousand years ago, due to higher sea levels, the Sunda coastline was different. Another route through the Nusa Tenggara archipelago was possible, starting from the southern tip of Sumatra, passing Timor, and reaching Sahul. The successful settlement of Sahul likely involved intentional, coordinated voyages by distinct populations over centuries.

 

They might have followed ocean currents and wind patterns. The southern route's past savanna corridor aided the migration of people adapted to such environments. After reaching Sahul, colonists occupied Australian deserts, and Indic sites dating back to fifty to forty-five thousand years ago. The northern route's environment resembled New Guinea, requiring coastal or forest travel.

Initial arrival into Sahul included two groups within fifty to sixty-five thousand years. One settled in Northern Sahul, which includes New Guinea and Near Oceania, and the other in Southern Sahul, which includes Australia and nearby islands. The two groups remained in relative isolation in the postglacial period after thirty thousand years ago, causing further diversification of human lineages in the region.

In Australia, evidence suggests that humans arrived around fifty to sixty thousand years ago. The Madjedbebe rock shelters provide some of the earliest proof of human presence, with artifacts dating back to that time. Skeletal remains from Lake Mungo display modern physical features, and ancient DNA from these remains reflects a unique Aboriginal lineage that still exists in today's population.

Around forty-five thousand years ago, people began colonizing Australia, adapting to diverse environments like deserts, grasslands, and coasts. They expanded gradually, leading to the unique biological and cultural diversity that characterizes Australia. In northeast Australia around forty-one thousand years ago, a decline in large animals and increased fires occurred, likely due to human hunting amid drying conditions. As people spread across the region, they relied on available animals.

They used various strategies to hunt small to medium-sized game and utilized permanent desert lakes for resources. Fishing tools like nets and spears, along with plant foods like yams, were used.

 

The Denisovans were an ancient human population that once roamed the Earth alongside the Neanderthals and modern Homo sapiens. However, nobody knew about them until a groundbreaking discovery in the Denisovan Cave in Siberia.

 

In two thousand and ten, an analysis of mitochondrial DNA extracted from a finger bone found in the Denisova Cave in the Russian Altai revealed the existence of a new hominin taxon. This newly described group was found to be genetically distinct from both Homo sapiens and Homo neanderthalensis. The mitochondrial DNA sequences showed haplotypes outside the range of variation of modern humans and Neanderthals.

 

Outside of Denisova Cave, a mandible found in Baishiya Cave in China was tentatively attributed to Denisovans based on proteomic evidence. Sediment DNA extracted from the site also confirmed the presence of Denisovans. These limited fossil findings suggest that Denisovans may have been widespread across continental Asia, island Southeast Asia, and Near Oceania.

During the Late Middle to Early Upper Pleistocene, three distinct groups emerged: early modern humans in Africa, Neanderthals in Europe, and Denisovans in Asia. Molecular data suggests that the split between Neanderthals and Denisovans occurred between three hundred and eighty to four hundred and seventy thousand years ago, while the branch leading to Denisovans and modern humans diverged around eight hundred thousand years ago. Homo heidelbergensis migrated to Eurasia around eight hundred thousand years ago, as indicated by the sites in Israel in the Levant region.

Around four hundred and fifty thousand to three hundred and fifty thousand years ago, Homo heidelbergensis from the Levant migrated to the Iranian Plateau and further to Central and North Asia, including Denisova Cave in southern Siberia, giving rise to Denisovans. This migration marked the splitting of late Homo heidelbergensis into Neanderthals and Denisovans.

Moreover, late Homo heidelbergensis from the Levant also reached to regions such as Turkmenia, Kazakhstan, and Mongolia, as indicated by the appearance of Acheulean industries in those areas. Their likely route was north of the Himalayas and Tibet. The mandible found in Baishiya Cave on the northeastern Tibetan Plateau dates back to at least one hundred and sixty thousand years ago.

The presence of Denisovans in the region is also supported by the adaptation of Tibetans to high altitudes possibly through introgression from Denisovans.

Hybridization events occurred between these groups, suggesting a meta population that includes all three taxa. The Altai region, where Denisova Cave is located, was occupied by Neanderthals, Denisovans, and modern humans, although the precise timing and coexistence of these groups remain uncertain. The Denisova 11 girl represents an example of hybridization, with a Neanderthal mother and a Denisovan father.

The study of ancient DNA has revealed significant genetic differences between Denisovans and modern humans. Denisovan genomes differ from the standard human genome by 11.7 per cent, while the difference between Neanderthals and modern humans is 12.2 per cent. Denisovans and Neanderthals are closely related but distinct from Homo sapiens.

Denisovans show higher genetic diversity than Neanderthals but lower diversity than modern humans, indicating a larger and more diverse population. Their geographic distribution ranged from North Asia to Southeast Asia.

Denisovans carried genetic traits associated with dark skin, brown hair, and brown eyes, which are also present in modern humans.

Denisovans likely played a role in adaptive introgression, where early Homo sapiens acquired genes from Denisovans that enhanced adaptation, disease resistance, and immune systems. The presence of human leukocyte antigens in modern humans, inherited from Denisovans, supports this hypothesis. For example, there are evidences that adaptation to high altitude hypoxia among modern day Tibetans is a result of introgression from Denisovans.

The populations of eastern Indonesians, Papuans, Philippine "negritos," Siberians, South Asians, and East Asians, are among the few existing groups that exhibit substantial traces of genetic material from Denisovans. It is worth noting that the region of Island Southeast Asia and Papua which consists of numerous densely inhabited archipelagos, holds some of the earliest evidence of early archaic humans outside Africa. Additionally, it is believed that archaic hominins like Homo floresiensis coexisted with modern humans in this region.

Modern Australian aboriginials and Papuans have the highest proportion of arround 5 to 6% of Denisovan ancestry, while American and mainland Asian populations have a smaller proportion about 0.2%.

The genetic diversity observed within the Denisovan lineage suggests their deep divergence and separation into three distinct branches across different regions. Three branches, D-one, D-two, and D-zero, have been identified. Branch D-two contributed to the introgression signal in Oceania and, to a lesser extent, in Asia. Branch D-one appears to be primarily confined to New Guinea and nearby islands, while branch D-zero is found in East Asia and Siberia. This indicates that Denisovans were capable of crossing significant geographical barriers, inhabiting a wide range of environments. The subgroups D-one and D-two diverged from the Altai Denisovan lineage approximately two hundred and eighty thousand and three hundred and sixty thousand years ago, respectively.

 

The genetic mixing with D-one Denisovans, restricted to New Guinea and nearby islands, may have occurred as recently as the end of the Pleistocene epoch, making them among the last surviving archaic hominins in the world.

There is an argument that interbreeding between humans and Neanderthals occurred east of the Wallace’s Line and another group argues that interbreeding occurred before humans crossed the Wallace’s Line.    

The distribution of Denisovan DNA in modern populations presents suggests significant interbreeding may have occurred after early humans crossed Wallace’s Line. Indigenous groups in New Guinea, Australia, and nearby islands exhibit the highest levels of Denisovan ancestry, reaching 3 to 4 percent. Interestingly, populations in mainland Asia, despite Denisovans’ historical range, show lower levels of Denisovan DNA.

 This disparity indicates that Denisovan-modern human interactions were concentrated east of Wallace’s Line, likely shaped by migration and ecological boundaries​ Mainland populations may have experienced a "replacement" effect, where later waves of East Asian populations diluted or replaced Denisovan DNA. Even in isolated groups like the Andaman Islanders, who show no admixture with other populations, no Denisovan DNA has been found.

 

Denisovans are thought to have lived across both mainland and Island Southeast Asia, thriving in diverse environments. During the Pleistocene, when sea levels dropped, vast savannah regions on the Sunda shelf (now submerged) likely served as migration routes and refuges for Denisovans during climate changes. This mobility may have allowed them to interact with modern humans in Island Southeast Asia, resulting in genetic mixing.

The only ancient human species clearly known to have crossed Wallace’s Line before modern humans is Homo floresiensis. This species remains mysterious, as scientists are unsure of its exact evolutionary origins. Some studies suggest it evolved from an early ancestor of Homo erectus, or perhaps from an even older and more primitive human species. Evidence of stone tools on Flores, dated to over 1 million years ago, indicates that these hominins arrived on the island very early.

However, it's difficult to link Homo floresiensis to the Denisovans. The Denisovans had larger molars (teeth), which don’t match the anatomy of Homo floresiensis. Moreover, genetic studies suggest that Denisovans and modern humans shared a common ancestor between one hundred seventy thousand  and 1 million years ago. This timeline makes it unlikely that Homo floresiensis, which had been isolated in the Wallacea region for so long, was directly related to the Denisovans.

Given the evidence, it’s likely that Homo floresiensis was a species uniquely adapted to its small island habitat, evolving separately for over a million years. The Denisovans, on the other hand, likely arrived later, around six hundred thousand years ago, and spread across a wider area of Asia and Southeast Asia. Some Denisovan fossils may yet be undiscovered, but possible candidates could include the Homo luzonensis from Philippines or other mysterious human fossils found in places like Narmada and Dali, Jinniushan, Maba, and Xujiayao (China). These sites suggest that Denisovans, or other ancient human species, once had a much broader range than previously thought. Recently evidence of human activity has in Kashmir valley, where around 300 to 400 thousand years ago humans relied on mammoth carcasses further supports this claim.

The Denisovan genome reveals evidence of interbreeding with another ancient population, but the identity of this group remains a mystery. Scientists speculate that Denisovans might have mixed with older human species like Homo erectus, Homo antecessor, or possibly a late-surviving Homo heidelbergensis in Asia. This raises intriguing questions about how many ancient human groups coexisted and interacted.

The fossil evidence from East Asia is incomplete and scattered, but it hints at a rich diversity of hominin species. Discoveries like Homo floresiensis on Flores and the Homo luzonensis remains from Callao Cave in the Luzon island of Philippines suggest that multiple human groups once inhabited Wallacea. This raises the possibility that some early hominins might have even reached New Guinea and Australia. If true, it would mean ancient humans were far more adaptable and capable of long-distance migration than previously thought.

Gene flow between Denisovans and modern humans occurred primarily east of Wallace’s Line, likely because the circumstances of these encounters were unique. Crossing Wallace’s Line required watercraft, which suggests that the first modern human groups reaching Wallacea were small and isolated. In such scenarios, interbreeding with the established Denisovan populations may have been more likely, as smaller populations often rely on intergroup mating to maintain genetic diversity. Additionally, genetic signals from these early interactions would be preserved more easily in the descendants of such small founding populations.

Interestingly, evidence suggests that this gene flow was predominantly male-mediated, meaning Denisovan males may have interbred with modern human females. This provides clues about the nature of these interactions, though much remains speculative.

Rapid dispersal into the tropical regions of Wallacea likely exposed early modern humans to unfamiliar pathogens. Hybridization with Denisovans, who were already adapted to the local environment, may have provided disease resistance genes that offered a survival advantage. This could have made interbreeding beneficial and increased the likelihood of these genetic traits being passed on and preserved.

As genomic research progresses, it is expected to shed more light on these interactions, revealing how Neanderthal and Denisovan DNA contributed to the genetic makeup of modern humans and whether some of the exchanged DNA had functional significance, such as aiding in disease resistance or environmental adaptation.

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