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Penghu 1 is a fossil jaw (
mandible In jawed vertebrates, the mandible (from the Latin ''mandibula'', 'for chewing'), lower jaw, or jawbone is a bone that makes up the lowerand typically more mobilecomponent of the mouth (the upper jaw being known as the maxilla). The jawbone i ...
) belonging to an extinct
hominin The Hominini (hominins) form a taxonomic tribe of the subfamily Homininae (hominines). They comprise two extant genera: ''Homo'' (humans) and '' Pan'' (chimpanzees and bonobos), and in standard usage exclude the genus '' Gorilla'' ( gorillas) ...
species of the genus ''
Homo ''Homo'' () is a genus of great ape (family Hominidae) that emerged from the genus ''Australopithecus'' and encompasses only a single extant species, ''Homo sapiens'' (modern humans), along with a number of extinct species (collectively called ...
.'' It was collected from seafloor sediments of the Penghu Channel off the coast of
Taiwan Taiwan, officially the Republic of China (ROC), is a country in East Asia. The main geography of Taiwan, island of Taiwan, also known as ''Formosa'', lies between the East China Sea, East and South China Seas in the northwestern Pacific Ocea ...
, dating to sometime in the
Middle Pleistocene The Chibanian, more widely known as the Middle Pleistocene (its previous informal name), is an Age (geology), age in the international geologic timescale or a Stage (stratigraphy), stage in chronostratigraphy, being a division of the Pleistocen ...
or
Late Pleistocene The Late Pleistocene is an unofficial Age (geology), age in the international geologic timescale in chronostratigraphy, also known as the Upper Pleistocene from a Stratigraphy, stratigraphic perspective. It is intended to be the fourth division ...
. The classification of the mandible was historically disputed. Some believed it to be the fossil of a '' H. erectus'', an archaic
Human Humans (''Homo sapiens'') or modern humans are the most common and widespread species of primate, and the last surviving species of the genus ''Homo''. They are Hominidae, great apes characterized by their Prehistory of nakedness and clothing ...
or possibly a
Denisovan The Denisovans or Denisova hominins ( ) are an extinct species or subspecies of archaic human that ranged across Asia during the Lower and Middle Paleolithic, and lived, based on current evidence, from 285 thousand to 25 thousand years ago. D ...
. Analysis of protein sequences in 2025 confirmed that the mandible represented that of a male Denisovan.


History and discovery

The fossil was recovered sometime before 2008 by fishermen working in the Penghu Channel between the
Penghu Islands The Penghu ( , Hokkien POJ: ''Phîⁿ-ô͘''  or ''Phêⁿ-ô͘'' ) or Pescadores Islands are an archipelago of 90 islands and islets in the Taiwan Strait, about west of the main island of Taiwan across the Penghu Channel, cove ...
and mainland
Taiwan Taiwan, officially the Republic of China (ROC), is a country in East Asia. The main geography of Taiwan, island of Taiwan, also known as ''Formosa'', lies between the East China Sea, East and South China Seas in the northwestern Pacific Ocea ...
and acquired by
Tainan Tainan (), officially Tainan City, is a Special municipality (Taiwan), special municipality in southern Taiwan, facing the Taiwan Strait on its western coast. Tainan is the oldest city on the island and commonly called the "Taiwan Prefecture, ...
citizen Kun-Yu Tsai. The fossil was found 60–120 meters below the water's surface and about 25 kilometers off the western coast of Taiwan in an area which was once part of the mainland. Sea levels have risen since the last ice age and in consequence have submerged the area where the fossil was recovered. After Tsai donated the fossil to the National Museum of Natural Science, it was described in 2015 by an international team of Japanese, Taiwanese, and Australian scientists. Penghu 1 is currently housed at the National Museum of Natural Science in
Taichung Taichung (, Wade–Giles: '), officially Taichung City, is a special municipality (Taiwan), special municipality in central Taiwan. Taichung is Taiwan's second-largest city, with more than 2.85 million residents, making it the largest city in Ce ...
. The fossil is stratigraphically dated to younger than 450 kya, based on prehistoric sea-level lowering to either between 190 and 130 kya, or to between 70 and 10 kya.


Fossil morphology

The fossil consists of a nearly complete right lower jaw with four teeth, including worn molars and premolars. The
mandible In jawed vertebrates, the mandible (from the Latin ''mandibula'', 'for chewing'), lower jaw, or jawbone is a bone that makes up the lowerand typically more mobilecomponent of the mouth (the upper jaw being known as the maxilla). The jawbone i ...
has a high index of robustness, a robust lateral torus, large molars, and with the help of 3D reconstruction it was revealed to have a large bicondylar breadth. These features help confirm that the fossil was from the middle-late Pleistocene era. The alveoli of its four incisors and right canine have been preserved as well showing their great length. The specimen was assigned to the genus ''Homo'' based on its jaw and tooth morphology. The mandible shows a receding anterior surface and lacks a pronounced chin which has helped distinguish it from the species ''Homo sapiens''. However, the fossil exhibited derived traits similar to early ''Homo habilis'' including the shortness and width of its jaw. These and other characteristics such as the agenesis of the M3 molar have been sufficient evidence to classify the specimen of the genus ''Homo''.


Classification

Although the genus of the Penghu 1 has been widely accepted, there is much discussion on the potential species of the specimen. The Penghu 1 mandible has been described as most similar to Hexian fossils of ''
Homo erectus ''Homo erectus'' ( ) is an extinction, extinct species of Homo, archaic human from the Pleistocene, spanning nearly 2 million years. It is the first human species to evolve a humanlike body plan and human gait, gait, to early expansions of h ...
.'' Both Penghu 1 and the Hexian mandible share a similar crown size, mandibular prominence, and general robustness. As a result of these similarities and their late presence in Eastern Asia, the authors of "The first Archaic Homo of Taiwan" proposed several models for their existence. The features the mandibles' shared could be explained by either the retention of primitive characteristics of early Asian ''Homo erectus,'' a migration of ''Homo'' with robust jaws from Africa, inclusion in the species ''Homo heidelbergensis,'' or they could have been an adapted form of ''Homo erectus.'' However, the species identity or taxonomic relationships lack consensus due to limited material. Co-author Yousuke Kaifu cautioned that additional skeletal parts are needed before species evaluation. Chinese anthropologists Xinzhi Wu and Haowen Tong did not agree with the naming of a new species, tentatively assigning the mandible to archaic ''Homo sapiens'', leaving open the possibility of elevating it to a distinct species should more fossils be discovered. In 2019, Chen Fahu along with a group of co-authors presented a piece suspecting the Penghu 1 mandible to belong to a
Denisovan The Denisovans or Denisova hominins ( ) are an extinct species or subspecies of archaic human that ranged across Asia during the Lower and Middle Paleolithic, and lived, based on current evidence, from 285 thousand to 25 thousand years ago. D ...
. This conclusion has been supported through its comparison with the Denisovan Xiahe mandible. The Xiahe mandible was discovered on the
Tibetan Plateau The Tibetan Plateau, also known as the Qinghai–Tibet Plateau or Qingzang Plateau, is a vast elevated plateau located at the intersection of Central Asia, Central, South Asia, South, and East Asia. Geographically, it is located to the north of H ...
and is dated to be around 160,000 years old. The Xiahe specimen has similar dental morphology compared to Penghu 1. They share 4 distinct characteristics: their M2's are close in mesiodistal width, they both show the agenesis of the M3 molar, they have a similar unique M2 root structure which relates to modern Asian populations, and the P3 displays Tomes' root, which is rarely found in other fossil hominins. Wu & Bae (2024) assigned Penghu 1 to the new species '' Homo juluensis'', as Xujiayao hominin, Xiahe mandible and Denisovans. A 2025 proteomic analysis study confirmed that the remains represented those of a male Denisovan.


See also

* Dispersal of ''Homo erectus'' * Prehistory of Taiwan


References


External links


Human Timeline (Interactive)
Smithsonian,
National Museum of Natural History The National Museum of Natural History (NMNH) is a natural history museum administered by the Smithsonian Institution, located on the National Mall in Washington, D.C., United States. It has free admission and is open 364 days a year. With 4.4 ...
(August 2016). {{Authority control History of Penghu Early species of Homo Pleistocene primates Homo fossils Prehistoric Taiwan