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    15 October 2025, Volume 44 Issue 05
    Human Fossils and Dating
    The hominin fossils from Hualongdong and their significance on human evolution
    LIU Wu, WU Xiujie
    2025, 44(05):  727-741.  doi:10.16359/j.1000-3193/AAS.2025.0065
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    Starting in 2014, ten years field work of excavations and deposit cleaning at the Hualongdong site, Dongzhi County, Anhui Province have resulted in more than fifty pieces of hominin fossils (including one well preserved skull), and stone tools and other evidence of hominin activities. Through fauna composition analysis and isotope dating, the hominin fossils were dated to about 300 ka. With these findings, the Hualongdong has been ranked as the most abundant hominin fossils and stone tool sites of Middle Pleistocene with Zhoukoudian. For the past ten years, a series of research with hominin fossils of cranium, facial bones, mandible, limb bones and teeth from Hualongdong have been conducted. Some new discoveries and understandings have been achieved from these studies. Our studies indicate the morphological traits of cranium, facial bones, mandible, limb bones and teeth from Hualongdong exhibit mosaics with combinations of East Asian Middle Pleistocene Homo erectus, Late Pleistocene hominin and modern human patterns. The cranium of Hualongdong has a suit of archaic features resembling those of East Asian Homo erectus, including low cranial vault, frontal keel, pronounced supraorbital torus, low position of maximum cranial breadth, slopping nasal floor and relatively small cranial capacity. Moreover, the cranium, facial bones, mandible and teeth show some derived features linking with Later Pleistocene hominins and modern humans. The modern human features in the Hualongdong hominin fossils include thin cranial wall, flat and gracile face, nearly vertical mandible symphysis, occurrence of initial mandibular trigon, and small and simple morphology teeth.

    The hominin fossils discovered from the Hualongdong site were regarded as the one population living in the same time from 270~330 ka. During the studies of the hominin fossils, we noticed some variabilities in the expressions of some morphological features. The lower second molar and the frontal bone discovered in 2006 exhibit a suit of feature patterns that are usually found in East Asian Homo erectus. The frontal bone shows a robust temporal line and thick cranial wall. Besides, the frontal bone has a metopic suture and enlarged frontal sinus. The occlusal surface of the molar has cusp 5, cusp 6 and cusp 7 plus complicated occlusal morphological pattern. Crown dimensions are within variations of Homo erectus and much larger than those of early modern human, contemporary human and European Middle Pleistocene humans. With these feature patterns, we proposed they may represent Homo erectus living during the Middle Pleistocene. In addition, the three pieces of Hualongdong femurs exhibit mosaic patterns with resemblance to Middle Pleistocene archaic hominins and Late Pleistocene early modern humans respectively, including lacking a pilaster, subtrochanteric pieces similar to Middle Pleistocene, and a prominent gluteal buttress and a small third trochanter resembling to many Upper Paleolithic hominins.

    These morphological patterns suggest that the 300 ka Hualongdong hominin fossils already exhibit the evolutionary trend towards the modern humans, and the Hualongdong hominins are the earliest Middle Pleistocene hominins in East Asia with most modern human features. The discoveries and related studies of Hualongdong hominin fossils provide new evidence for the hominin evolutionary diversities of East Asian late Middle Pleistocene.

    Sedimentary feature and chronology of Hualongdong site in Dongzhi, Anhui Province
    CAI Yanjun, PEI Shuwen, JIN Zetian
    2025, 44(05):  742-753.  doi:10.16359/j.1000-3193/AAS.2025.0075
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    Resolving sedimentary processes and constraining the ages of the Hualongdong site in Dongzhi County, Anhui Province, are crucial for researching ancient human evolution, as well as its living environment and adaptive survival behaviors. This paper comprehensively analyzes the geological, geomorphological, and sedimentary characteristics of the deposits at the Hualongdong site and its surroundings. In addition, we also examine the 230Th-234U-238U dating results of the speleothem debris and animal fossils collected during the excavation. It is confirmed that the deposits at the Hualongdong site formed from a mixture of cave deposits and surrounding rock fragments during the collapse of an ancient karst cave. The transition from the ancient karst cave to the accumulation at the Hualongdong site occurred in three main stages: the cave formation and development period, a relatively stable deposition period, and a rapid collapse and accumulation period. The collapse of the ancient karst cave and the formation of mixed deposits at the site occurred later than approximately 216 thousand years before present (ka BP), and ancient cave deposits containing fossils are the main components of the mixed accumulation at the site. Along with the dating results attained previously, the 230Th-234U-238U dating results of the recently discovered flowstone within the deposits indicate that the cave underwent at least two accumulation periods during its stable sedimentation phase, with early deposition occurring prior to 600 ka BP and later deposition spanning from around 331 to 275 ka BP. The mixing of deposition across at least two depositing episodes adds complexity to the chronological study of the site. By integrating the 230Th-234U-238U dating results of all available speleothem, fossil teeth and bones, this study speculates that the burial age of the most complete human skull fossil discovered at the Hualongdong site can be bracketed between 275 and 331 ka BP, placing it in the late Middle Pleistocene. We suggested that Uranium-series dating of representative fossil specimens can offer vital insights for the chronological study of the site and is strongly recommended for understanding the evolution of the fauna and ancient humans at this site.

    Craniofacial reconstruction and morphological characteristics of Hualongdong No.6 hominid skull
    JIN Zetian, WU Xiujie, DENG Guodong, LIU Wu
    2025, 44(05):  754-764.  doi:10.16359/j.1000-3193/AAS.2025.0063
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    The Hualongdong (HLD) site is situated in Wangcun Village, Yaodu Town, Dongzhi County, Anhui Province, China, on the southern slope of Meiyuan Hill. The site is characterized by a collapsed cave comprising carbonate-cemented cavern breccia, unconsolidated clay and gravel. It was discovered in late 1988. Following several excavation campaigns between 2006 and 2024, approximately 20 human individuals, more than 400 artificial stone tools, numerous bone fragments with signs of manual cutting and chopping, and over 80 species of animal fossils have been uncovered. These findings date back to between 331,000 and 275,000 years ago.

    In this paper, we carried out facial restoration on the HLD No.6 skull fossil unearthed at this site. The HLD 6 hominid skull was found in 2015. When unearthed, the skull was broken into 20 fragments of different sizes, among which 7 were wrapped in a large cementing material, and the other 13 dispersed around the large cementing material. According to the bone thickness, bone sutures, cross-sections, the middle cerebral artery and the occlusal state of the teeth, a three-dimensional model of the HLD skull was 3D virtual restored. After reconstruction, it preserves almost entire face, most of the mandibular and a largely cranium.

    To show the pre-life appearance of HLD hominid, according to the relationship between facial bones and soft tissues, a manual method was adopted to reconstruct the outer surface of the skull with muscle, skin, and facial features using plasticized materials (silly putty and plaster) on the 3D virtual reconstruction of the 3D printed skull model. After craniofacial reconstruction, the HLD hominid showed more modern human features with a flat glabella, a backward forehead, a flat face, and the appearance of a chin, however, it still retained some archaic features such as a developed brow ridge, a low cranial height. The HLD human fossils exhibit a mix of physical characteristics transitional between Homo erectus and modern humans. The cranium displays more primitive traits, the facial structure resembles that of modern humans, and the mandible shows early signs of modern human chin. Compared with the "pre-Homo sapiens" Jebel Irhoud found in Morocco, Africa, dated to the same age, the two show similar transitional physical features from archaic to Homo sapiens. The HLD hominin fossils provide crucial evidence of the diversity of late Middle Pleistocene human evolution in East Asia, indicating that modern forms related to Homo sapiens emerged in this region around 300,000 years ago.

    Stone Artifacts, Animal Fossils
    Lithic assemblage and adaptive behaviors of hominins at Hualongdong site
    PEI Shuwen, DONG Zhe, GENG Shuaijie, YE Zhi, MA Dongdong, ZHANG Yameng, JIN Zetian
    2025, 44(05):  765-778.  doi:10.16359/j.1000-3193/AAS.2025.0070
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    The Hualongdong (HLD) site (latitude 30°06′34.1′′N, longitude 116°56′54.2′′E, 40 m above sea level) is located in Dongzhi County, Anhui Province, China. It was initially discovered in 1988, the excavations at the site in 2006, and from 2014 to 2024 resulted in the discovery of more than 20 individual human fossils, an abundance of lithic artifacts and mammalian fauna. The original cave deposits was about 20 m higher above its current location. Synsedimentary dismantlement, downward slippage, and bedrock weathering are indicated by the brecciated arrangement of the limestone rock blocks, cemented angular, subangular clasts and archaeological remains within the excavation area. Uranium-Thorium dating of speleothems and animal teeth from the brecciated deposits indicated that early human occupied the site most probably took place in the late Middle Pleistocene, ca. 300 ka (270~330 ka). This paper presents the lithic technology and the adapted behaviors adopted by humans at the site.

    The Hualongdong lithic assemblage consists of 322 artifacts which can be divided into Detatched pieces [flakes (N=79; 24.6%), debitage shatters (N=96; 29.8%), bipolar elements (N=45; 14.0%)], Flaked pieces [cores (N=65, 20.2%), core fragments (N=4; 0.3%), bipolar cores (N=2; 0.6%), retouched pieces (N=22; 6.8%)] and Pounded pieces (8 hammerstones and 1 anvil). Lithic raw material includes quartz/quartzite and lava show fluvial cortex and suggests sourcing from streams, while chert nodules and some quartz rock block derive from the outcrop of chert and quartz -bearing bed in bands within siliceous carbonate-clastic strata formation in the surrounding landscape. Quartz is the predominant (87.9%) rock type. Technologically, cores, debitage shatter, and bipolar elements from HLD indicate that freehand percussion was the major knapping technique, while the use of hammer-and-anvil technique is the complementary technique to knap cobble and low-quality quartz. HLD reduction sequences are relatively short, non-organized and without core preparation, resulting in a low degree of standardization in flake morphologies and dimensions. Albeit scarce, retouched pieces are present in HLD lithic assemblage, with an average proportion of 6.8%. Retouch is normally on flakes or flake fragments even cobbles. Scraper dominants the tool type, followed by chopper, denticulate, notch and point. Retouch is casual in HLD assemblages, with no imposition of standardized shapes on blanks. The HLD lithic assemblage can be assigned to Oldowan-like or Mode 1 technology in east Asia as characterized by a simple technological design, low degree of standardization, expedient flaking, and a few non-standardized retouched flakes.

    All in all, the HLD core and flake technology includes bipolar flaking of low-quality quartz rock block and cobbles, and freehand flaking of chert, lava and quartz block fragments, which demonstrate knappers intentionally selected good-quality raw materials to manufacture small flakes and use them for some function. Most chert and relatively good quality of quartz flakes show use wear, probably associated with carcass processing given the presence of cutmarks on some bones. Furthermore, it can be inferred that the general feature of HLD toolkits were expediently made for the function of getting food from animal resources and the knapping strategy was adapted to the local resources in the HLD archaeological landscape.

    New mammalian fossils from the Hualongdong site in Dongzhi County, Anhui
    TONG Haowen, JIANGZUO Qigao, LI Qiang, CHANG Meijing, LIU Boxuan, JIN Zetian
    2025, 44(05):  779-798.  doi:10.16359/j.1000-3193/AAS.2025.0066
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    The Hualongdong site (Hualongdong Loc.1) in Dongzhi County, Anhui Province is the most important fossil site of mid-Pleistocene Homo found in China in recent years. The Hualongdong site was discovered in 1988 and has undergone 9 formal excavations in 2006, 2014~2016, 2017~2019 and 2023 respectively. The mammalian fossils unearthed during the periods 2014~2016 have been preliminarily reported; Since 2017 onward, a large number of mammal fossils have been discovered, and more species have been added to the fauna list: Rhinopithecus sp., Canis sp., Ailuropoda wulingshanensis, Panthera pardus, Neofelis nebulosa, Prionailurus kurteni, Prionodon hualongensis, Paguma larvata, Urva urva, Crocuta ultima, Equus sp., Stephanorhinus sp., Rusa unicorlor, Hydropotes sp. and Capricornis sp. At present, there are as many as 93 species, including undetermined species of mammals have been recognized from the Hualongdong site. More and more complete fossil specimens have been added to the previously reported genera and species, such as the Ursus thibetanus, Ailuropoda baconi, Panthera tigris, Sus lydekkeri, Cervus grayi, and Bos (Bibos) sp., among which Sus lydekkeri and Bos (Bibos) sp. have the largest body sizes for their kinds ever known. The fossils of brown bear (Ursus arctos), horse (Equus sp.), Lydekker's pig (Sus lydekkeri) and giant deer (Sinomegaceros sp.) from Hualongdong represent the southernmost occurrence of their kinds in eastern China, and they are more frequently recovered in the northern faunas. The age of the fauna of the Hualongdong site is complicated by the discovery of the fossil of Ailuropoda wulingshanensis, because this species was only found in the late Early Pleistocene sites, which is obviously inconsistent with the formation time (ca. 300 ka) of the Hualongdong site as currently dated. The explanation for this phenomenon is most likely due to the collapse and re-accumulation of the original cave deposits, resulting in a mixture of old and new strata. The dominant species of the Hualongdong fauna are gaurs and diverse cervids, which are very similar to those of the Bailongdong fauna in Hubei Province. The Hualongdong fauna is different from that of the nearby Hexian Man Site in the lackness of gaurs in the latter. In general, the dominant species of the Hualongdong fauna is comparable to the Ailuropoda-Stegodon fauna of Middle Pleistocene in southern China, but with a few invaders from the North, which suggests a relatively cold substage at which the Hualongdong Man once experienced. The Hualongdong fauna is different from the Late Pleistocene fauna of southern China, the latter is usually dominated by the Sus scrofa-cervids assemblage.

    Survival paleoenvironment of the Hualongdong Hominid reflected by non-volant micromammalian fossils
    CHANG Meijing, LI Qiang, NI Xijun, ZHANG Xuan, TONG Haowen
    2025, 44(05):  799-815.  doi:10.16359/j.1000-3193/AAS.2025.0067
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    Hualongdong (HLD) site in Dongzhi County, Anhui Province is another important locality for early hominin skull fossils following notable sites such as Zhoukoudian in Beijing, Lantian in Shaanxi, Hexian in Anhui and Tangshan in Nanjing. The geological age of the site is approximately 300 ka. Since its discovery in 2004, multiple excavations have been conducted at HLD site, yielding abundant hominin and associated vertebrate fossils. This study focuses on the taxonomic identification, faunal analysis of non-volant small mammal fossils associated with hominin fossils from the 2015-2017 excavation, and paleoenvironmental reconstruction. A total of 5,645 non-volant micromammal fossil specimens have been discovered, primarily consisting of isolated teeth. These fossils represent 3 orders, 11 families, 38 genera, and 49 species (including two species indeterminata). In terms of both the number of species and the number of identified specimens, Rodentia is dominant at the ordinal level; Muridae is the most diverse at the familial level; and Apodemus is the most abundant at the generic level. The extinction proportion of HLD site is low (4.26%), which is obviously different from the Early Pleistocene and the early Middle Pleistocene non-volant small mammal assemblages with abundant Tertiary relict species, and the Late Pleistocene animal assemblages with almost complete disappearance of extinct species, but more similar to the late Middle Pleistocene small non-volant mammal assemblage in southern China. In terms of faunal composition, the non-volant micromammalian assemblage of the HLD site shows the higher similarity to that of Yumidong of Chongqing, Xinglogndong of Chongqing, Yanhuidong of Guizhou, and Baotansi of Chongqing. The comparison with the present small mammal assemblages shows that remarkable species alternations have taken place in this region, with the decline in species quantity of Soricinae, flying sciurids and Apodemus, the emergence of temperate species Myospalax and the disappearance of subtropical and tropical species since the late Middle Pleistocene to the present.

    The 45 extant species at the HLD site collectively reflect a forest-dominated habitat; however, the top eight dominant species, as determined by minimum number of individuals (MNI), indicate a mixed forest-grassland environment. Based on the MNI, we analyzed the habitat preferences of these dominant species. Among them, Pteromys volans and Lasiopodomys brandtii are primarily distributed in temperate regions; Eothenomys melanogaster is usually inhabited subtropical and tropical shrublands; species of Apodemus and Niviventer andersoni are mainly found in temperate and subtropical areas; while Mus musculus is widely distributed from tropical to temperate zones. These species typically inhabit various forest types and meadow environments at forest edges. In light of the small mammal assemblage, we infer that the HLD site in Dongzhi, Anhui, during the late Middle Pleistocene, featured a mixed environment of forests and meadows. The mean annual temperature (MAT) and mean annual precipitation (MAP) reconstructed using the Random Forest algorithm for the late Middle Pleistocene HLD site were 12.57°C and 909.81 mm (based on the eight MNI dominant species) and 13.38°C and 1014.70 mm (based on all 45 extant species), respectively. These values are lower than the meteorological data from present-day Dongzhi (16.1°C and 1628.3 mm), indicating that the climate at that time was somewhat cooler and drier.

    Mammalian astragalus fossils excavated at Hualongdong site
    LIU Sitong, LIU Boxuan, JIN Zetian, DENG Guodong, TONG Haowen, WU Xiujie
    2025, 44(05):  816-835.  doi:10.16359/j.1000-3193/AAS.2025.0072
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    The astragalus is one of the tarsal bones and anatomically subdivided into three distinct regions: the head, neck, and body. This critical bone serves as the primary osseous interface between the distal tibia and proximal tarsal bones, forming the talocrural articulation that facilitates the characteristic hinge-like motion of the tetrapod ankle joint. Furthermore, comparative osteological analyses reveal that the astragalus typically exhibits greater structural density compared to other skeletal elements, a morphological adaptation likely associated with its weight-bearing function within the ankle complex. This increased bone density, combined with its strategic anatomical position protected by adjacent articulating surfaces, significantly enhances its preservation potential in both modern and paleontological contexts. The exceptional preservation frequency of astragali in vertebrate fossil records has consequently established this element as a crucial anatomical marker for phylogenetic studies and functional morphology analyses. In addition, since the structure of the astragalus is directly related to the locomotion patterns of animals and reflects the range of motion of the hind limbs, it is often used to analyze the characteristics of species and their ecological and functional adaptability. The Hualongdong Site is an important comprehensive ancient human site from the late Middle Pleistocene in China, where a large number of mammalian fossils have been unearthed. This paper takes 316 specimens of mammalian astragalus fossils excavated at the site from 2014 to 2024 as research materials. The investigation methodologically integrates the measurement, the identification of morphological characteristics and the analysis of surface traces, This systematic approach facilitates a comprehensive exploration of the animal group, the ancient human behaviors, and the background of burial. The mammalian astragalus bones unearthed at the Hualongdong reflect a fauna composition dominated by the Cervidae and the Bovidae, accounting for 84.5% of the identifiable specimens and 80.5% of the minimum number of individuals respectively. Sus lydekkeri and carnivores, predominantly from the Ursidae also account for a certain number. There are also a few specimens from the Rhinocerotidae, Equidae, Tapiridae and Stegodontidae. The presence of the austral Ailuropoda-Stegodon faunal elements is represented by Ailuropda, Stegodontidae, and Tapiridae, while the presence of boreal taxa is represented by Cervus (Sika) grayi, Equidae, Stephanorhinus sp. In fact, the NISP and MNI of the boreal taxa are higher than the austral taxa. The astragalus measurement data show that Bos (Bibos) sp., Equidae, Rhiocerotidae, and Sus lydekkeri at the Hualongdong site represent the larger body sizes comparative to their relatives during the same period. Overall, the composition of the animal group reflected by the astragalus indicates a relatively open forest-grassland landform, with the presence of mountain animals. Burial analysis indicates that the astragalus is less affected by natural forces such as weathering and erosion by water flow. A thorough examination of the surface marks reveals that the astragali of the Cervidae and Bovidae families exhibited artificial cutting marks, chopping marks, carnivorous biting marks, and rodent gnawing marks. This indicates the involvement of humans, carnivores, and rodents in the accumulation of astragali in the cave. This study is the first to systematically reveals the composition of the animal group and the evidence of human behavior reflected by the astragalus fossils at the Hualongdong site. It provides reference materials for the research on the survival strategies and animal resource utilization of ancient humans in East Asia during the late Middle Pleistocene.

    Fossil carnivora from the Hualongdong Site Loc.1 and their chronological and paleoenvironmental implications
    JIANGZUO Qigao, LIU Boxuan, LIU Sitong, HE Letian, TONG Haowen
    2025, 44(05):  836-849.  doi:10.16359/j.1000-3193/AAS.2025.0064
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    The Hualongdong Hominid Site is an important locality for ancient human fossils in the middle and lower reaches of the Yangtze River in China, where several well-preserved ancient human fossils and many mammal fossils have been discovered. This paper systematically studies the Carnivora fossils excavated from Hualong Cave from 2014 to 2024, including 8 families, 14 genera, and 17 species, including Cuon alpinus, Ursus thibetanus, Ursus arctos, Ailuropoda melanoleuca baconi, Arctonyx cf. collaris, Catopuma teilhardi, Prionailurus kurteni, Prionailurus sp., Panthera tigris, Panthera pardus, Neofelis nebulosa, Prionodon hualongensis sp. nov., Pachycrocuta brevirostris sinensis, Crocuta ultima, Viverra zibetha, Paguma larvata, Urva urva. Felidae are the most diverse family, with six species coexisted, ranging from about 1kg P. kurteni to around 200 kg P. tigris. Ursidae are also diversified, with three species, and two of them, U. thibetanus and A. m. baconi are the two abundant carnivore species in the fauna. Among these species, the newly identified species of linsang (Prionodon hualongensis) is the first fossil species discovered from the genus Prionodon, characteristic by having large body size, robust dentition and mandible, wide m1 protocone and m1 talonid, indicating a diet comprising more hard tissue. The presence of the abundant spotted hyena, indicating that the site is older than Zhoukoudian Loc.1 (very rare and only present in the upmost layer). On the other hand, the morphology of the brown bear and the golden cat is similar to those from the Zhoukoudian Loc.1, and the morphology of the dhole (relatively distinct m1 entoconid) and the hog-nosed badger (relatively large m1 paraconid) appears slightly more primitive than that of modern population. The archaic element giant hyena Pachycrocuta is still present in the locality, and there are also several extinct species, including the Hualong linsang and the Kurten's leopard cat. In summary, the composition and evolutionary level of carnivore guild from the Hualongdong site supports a late Middle Pleistocene age, which aligns with the results of previous dating. The Hualongdong carnivore guild mainly composes of species typical of woodland areas (no specific temperature preference), such as the dhole and the Asian black bear, followed by typical southern China components like the giant panda and the clouded leopard. However, there are also typical North China components, notably the brown bear, indicating a mosaic habitat primarily dominated by forests.

    Middle Pleistocene snakes from Hualongdong Site Loc.1, in Dongzhi County, Anhui
    SHI Jingsong, HU Haiqian, DENG Guodong, JIN Zetian
    2025, 44(05):  850-861.  doi:10.16359/j.1000-3193/AAS.2025.0073
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    The Hualongdong Site in Dongzhi County, Anhui Province, is a well-known Middle Pleistocene archaeological site in East Asia associated with late archaic hominin activities. Here we initially report several postcranial snake fossils that discovered from Loc.1. In this study, we conducted a preliminary identification of these specimens based on integrative osteological comparison between fossils and extant snakes. Three families, five genera, and six extant species were identified, which can be attributed to extant taxa. Diagnosis for each identified genus was summarized. The zoogeographic composition of the snake assemblage is primarily characterized by South China, Oriental, and Monsoonal elements. Among them, the Oriental tropical type Ptyas cf. carinata is most frequently recorded. The snake fauna might reveal a warmer and more humid climate in Hualongdong Site during the late Middle Pleistocene compared to the present.

    Hunting behavior of Hualongdong Hominid based on the teeth of gaur
    LIU Boxuan, LIU Sitong, JIN Zetian, DENG Guodong, WU Xiujie
    2025, 44(05):  862-873.  doi:10.16359/j.1000-3193/AAS.2025.0076
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    The Hualongdong site in Dongzhi County, Anhui Province has yielded abundant hominin fossils along with numerous mammalian bones and teeth. Among the large mammal fossils unearthed at the site, cervid remains are the most common, followed by those of large bovids. To investigate the relationship between the Hualongdong hominins and these mammals, this study focuses on the gaur [Bos (Bibos) sp.] dental fossils excavated from the site between 2014 and 2024. The subgenus Bos (Bibos) represents an extinct group of the Bos (Bibos) sp., distinct from extant Bos frontalis. Fossils of these bovids are frequently found in Pleistocene cave sites in southern China, with a temporal range spanning from the Early to the Late Pleistocene. In this study, the dental sequence was first identified, and the tooth dimensions were measured and compared with those of bovids from other sites as well as extant gaur. Based on this analysis, the minimum number of individuals (MNI) was calculated. The age-at-death was estimated using crown height measurements, and age groups were classified to construct the mortality profile. The mortality pattern of the Hualongdong bovids was identified using a ternary diagram and its possible causes inferred through comparative analysis.

    The results indicate that: 1) A total of 1,205 bovid teeth were unearthed, including 139 incisors (Including incisiform canines), 202 fragmented cheek teeth, and 864 complete and measurable cheek teeth (69 deciduous teeth and 795 permanent teeth); 2) The MNI was estimated at 53, based on the combination of 39 left M3s and 14 right DP3s; 3) The tooth dimensions of the Hualongdong bovids most closely resemble those of Bos (Bibos) gaurus from the Bailong Cave site; 4) Using a quadratic crown height equation for permanent teeth and a linear equation for deciduous teeth, the age distribution of the Hualongdong bovids was determined to be 38% juveniles, 44% prime adults, and 18% old adults; 5) The age profile plotted on a ternary diagram closely aligns with that of extant wild gaur and European bison, indicating a catastrophic mortality pattern; 6) The relatively high proportion of juveniles suggests that the Hualongdong hominins may have employed multiple hunting strategies, such as a combination of non-selective ambush and endurance hunting, or that they may have targeted more juvenile bovids during the breeding season.

    Teeth are among the most well-preserved animal remains at archaeological sites and contain rich information about the animals themselves. By analyzing the dental fossils of large bovids from Hualongdong, this study reconstructs their mortality profile and provides new evidence for interpreting the hunting behavior of the Hualongdong hominins.

    Production and processing of large flakes by hominins during the Middle to Late Pleistocene in the Chaohu Lake Region, Anhui Province
    ZHAN Shijia, DONG Zhe, YI Shuangwen
    2025, 44(05):  874-883.  doi:10.16359/j.1000-3193/AAS.2025.0062
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    Although quantities of Acheulean large cutting tools discovered in different areas and sites in China have been insightfully analyzed, there have been few studies focusing on large flake production and large or giant core exploitation until now. In fact, producing large flakes with the expected morphology from giant or large cores through special percussion methods is a key criterion of Acheulean technology. During the 1980s-1990s, the Institute of Archaeology and Cultural Relics of Anhui Province conducted several Paleolithic investigations and excavations in the Chaohu Lake Region (CLR). A total of 319 stone artifacts embedded in red clay were discovered on the surface and in situ, among which 107 Large Flake Acheulean products could be identified. All of these artifacts were detached from quartzite river cobbles using free-hand stone hammer percussion technology. Traditionally, these artifacts were assigned to the Pebble Tool Industry. Based on lithic technological typological analysis, it can be deduced that hominins detached large flakes from large cores via the cobble-opening method and slab-slicing method, and further shaped them into large cutting tools (LCTs) such as bifaces, knives, cleavers, and picks—alongside Acheulean large tools, particularly massive scrapers and massive denticulates.

    Based on the OSL dating results from the Wangjia'ao Locality (near Wangchenggang, the 1980s' and 1990s' work area), early humans occupied this site around 70 kaBP. Recently, two additional OSL dating analyses were conducted at the Liujia 2 Locality and Zhangjiawan Locality, both approximately 4.5 kilometers from the Wangjia'ao Locality. At the Liujia 2 Locality, hominins exploited large cores, detached large flakes, and further used these flakes as blanks to manufacture LCTs during two distinct periods: 225~70 kaBP and 70~30 kaBP. In contrast, the OSL dating result from the Zhangjiawan Locality indicated that early humans detached large flakes no later than 125 kaBP.

    All the aforementioned OSL dating results from the CLR demonstrate that ancient humans in this region exhibited diverse adaptive behaviors, as they were capable of producing large flakes and manufacturing LCTs no later than Marine Isotope Stage 4 (MIS 4). A similar pattern has also been observed at the Wenquan Site in Henan Province, and an even earlier occurrence has been documented in the Luonan Basin and Bose Basin. This research holds great significance for studying the transition of lithic technology and the production of large flakes in Acheulean technology in South China from the late Middle Pleistocene to the Late Pleistocene.

    Isotope, Palynology, Paleoprotein
    Isotopic paleoecology of Ursidae from Hualongdong, Anhui
    MA Jiao, JIANGZUO Qigao, JIN Zetian, DENG Guodong, CHEN Yiying, YAN Yi
    2025, 44(05):  884-894.  doi:10.16359/j.1000-3193/AAS.2025.0068
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    This study investigates the dietary ecology of three sympatric Ursidae species— Ailuropoda melanoleuca baconi, Ursus thibetanus, and Ursus arctos — from two sites at Hualonglong (locality 1: ~300 ka; locality 3: late Middle Pleistocene to Late Pleistocene) in Anhui Province, southern China. Through stable carbon and oxygen isotope (δ13C, δ18O) analysis of tooth enamel, this study reveals distinct ecological strategies: at locality 1, U. arctos (n=3) occupied intermediate woodlands (δ13Cdiet= −23.1‰±1.9‰), while A. melanoleuca baconi (n=5) and U. thibetanus (n=4) inhibited close-canopy forests (δ13Cdiet values are −27.8‰±0.6‰ and −27.8‰±0.8‰, respectively). The latter exhibites more diverse dietary ecology, as reflected by their varied δ18O values (−6.9‰±0.3‰ and −8.3‰±1.1‰). By locality 3, U. arctos were absent, but A. melanoleuca baconi (n=5) and U. thibetanus (n=5) continue to inhabit close-canopy forests. The δ18O convergence between A. melanoleuca baconi (−7.0‰±0.4‰) and U. thibetanus (−7.2‰±0.8‰) suggests possible ecological shifts in U. thibetanus. Regional comparisons (Guangxi: Baxian Cave, Quzai Cave, and Yugong Cave from previous studies; Anhui: two localities at Hualongdong in this study) indicate that during the middle-late Pleistocene, both A. melanoleuca baconi (δ13Cdiet values were −27.1‰±1.0‰ (n=20) in Guangxi and −27.8‰±0.6‰ (n=10) in Anhui) and U. thibetanus (δ13Cdiet values were −27.7‰±1.7‰ (n=10) in Guangxi and −27.8‰±0.6‰ (n=9) in Anhui) consistently exploited close-canopy forests in Guangxi and Anhui, yet maintained niche partitioning at each sites. By integrating isotopic data from Pleistocene Ailuropoda, we observed a gradual shift towards more positive δ13Cdiet values from the Early Pleistocene Ailuropoda microta (data from Gigantopithecus Cave and Yanliang Cave in Guangxi, δ13Cdiet= −28.2‰±1.0‰, n=8) and A. melanoleuca wulingshanensis (data from Longgu Cave in Hubei, δ13Cdiet= −27.7‰±0.7‰, n=4) to the middle-late Pleistocene A. melanoleuca baconi (all aforementioned specimens from Guangxi and Anhui, δ13Cdiet= −27.3‰ ±0.9‰, n=30). Their δ18O values also varied by regions and periods. The spatiotemporal isotopic differences in Pleistocene Ailuropoda, linked to their specialized bamboo-diet, underscore the differences in regional paleoenvironments in southern China. In Guangxi, A. melanoleuca baconi exhibited slightly higher δ13Cdiet values and a broader range of δ18O values compared to A. microta. This may suggest that, with the increase in body mass and food intake, A. melanoleuca baconi occupied more open habitats and utilized bamboo more diversely, as potentially evidenced by a wider variety of bamboo species, parts, and distribution ranges. This study provides the first isotopic evidence for sympatric Ursidae niche partitioning in southern China, offering critical paleoenvironmental context for understanding the coevolution between human and non-human animals in this region.

    Dietary ecology of gaur from the Hualongdong site in Anhui Province
    ZHENG Mingcong, WANG Jingyi, YAN Yi, CHEN Yiying, WU Yan
    2025, 44(05):  895-905.  doi:10.16359/j.1000-3193/AAS.2025.0071
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    The Hualongdong site in Dongzhi County, Anhui Province, represents the most comprehensive Paleolithic site in China since Zhoukoudian, yielding abundant hominin fossils alongside stone tools and other evidence of human activities. Notably, it has also produced a substantial assemblage of mammalian fossils. As a key species in China's Quaternary mammalian fauna, the Pleistocene gaur [Bos (Bibos) sp.] shares close phylogenetic relationships with its extant counterparts. However, systematic understanding of this fossil Bos (Bibos) sp. species remains limited due to fragmentary fossil materials and methodological constraints. The dental fossils of Bos (Bibos) sp. from Hualongdong site provide crucial materials for investigating its dietary habits, ecological niche, paleoenvironment and ecosystem. This study employs multidisciplinary approaches including CT three-dimensional modeling, phytolith analysis, and stable isotope analysis to systematically investigate Bos (Bibos) sp. dental fossils. CT modeling precisely identified dental calculus locations, enabling subsequent phytolith extraction and morphological characterization from calculus deposits. Concurrently, carbon stable isotope analysis was conducted on tooth enamel. Results reveal that the late Middle Pleistocene Hualongdong site Bos (Bibos) sp. feed on herbaceous plants of varying heights. Carbon isotope data indicate a primary reliance on C3 plants from the site and surrounding areas, supplemented by limited C4 plant consumption (δ13C values: -10.3‰ to -4.6‰). Phytolith analysis specifically indicates frequent ingestion of Poaceae and Zingiberales species. These findings collectively indicate that Bos (Bibos) sp. primarily foraged in warm environments dominated by C3 vegetation during the late Middle Pleistocene. The diverse phytolith assemblages and isotopic signatures reflect rich regional vegetation heterogeneity, providing ample food resources. Furthermore, the common occurrence of Pleistocene Bos (Bibos) sp. fossils in alluvial plains, mountain valleys, and karst caves suggests their preference for open forest areas with grassy patches. This ecological pattern aligns with their mixed dietary strategy of feeding on both C3 and C4 plants. This research establishes novel phytolith and isotopic records for understanding Bos (Bibos) sp. paleodietary ecology while contributing to Middle-Late Pleistocene paleoclimatic and paleoecological reconstructions in Anhui Province. The integrated methodology demonstrates the efficacy of combining Plant microfossils and geochemical proxies in paleodietary studies, offering insights into herbivore-environment interactions during the Pleistocene.

    Vegetation and environmental context of archaic human occupation at the Hualongdong site, Anhui
    ZHAO Keliang, BAI Guangyi, HUI Jiaming, ZHANG Xuan, LI Xiaoqiang
    2025, 44(05):  906-916.  doi:10.16359/j.1000-3193/AAS.2025.0074
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    As a crucial Middle Pleistocene hominin activity site, Hualongdong (HLD) provides pivotal evidence for understanding ancient human evolution and environmental adaptation in East Asia. The site's hominin fossils and stone tools were unearthed from breccia and brownish-red clay deposits. To reconstruct the paleovegetation and better understand the paleoenvironmental context of the Hualongdong hominins, this study conducted pollen analysis on 20 sediment samples collected from these strata in the 2016 excavation area. A total of 41 pollen taxa were identified across the samples. From the breccia deposits (7 samples), 94 pollen grains were recovered, whereas 1,148 grains were counted in the 13 samples from the HLD2016 section. Uranium-series dating of stalagmites and animal teeth fossils indicates that the breccia deposits formed between 331 and 275 ka BP. Although the HLD2016 section lacks direct dating, published chronostudies suggest that the brownish-red clay was deposited during the late Middle Pleistocene, approximately 0.4~0.1 Ma. Notably, the pollen assemblages from the lower portion of the HLD2016 section (below 150 cm) closely resemble those of the breccia deposits. This similarity suggests that the sedimentary layer between 240 and 150 cm may correspond temporally to the breccia deposits.

    In the breccia samples and the HLD2016 sediments below 150 cm, arboreal pollen dominates (67.7%), followed by herbaceous (31.9%) and shrub pollen (0.4%). The assemblage reflects a deciduous broad-leaved forest dominated by Ulmus and Quercus, with an understory and surrounding grasslands composed of Poaceae, Chenopodiaceae, and Artemisia. This vegetation structure is distinct from the region's current subtropical environment but similar to the warm-temperate deciduous broad-leaved forests of southern North China suggests that the climate during the Hualongdong hominin occupation was cooler and drier than today. The landscape likely consisted of a forest and grassland mosaic, offering diverse ecological resources. Comparative analysis reveals that Hualongdong's paleoenvironment shares similarities with that of the Nanjing site but exhibits denser vegetation cover than Zhoukoudian and Dali, along with intermediate openness compared to Miaohoushan and Xujiayao. These environmental features may reflect the ecological adaptation strategies of Middle Pleistocene hominins in East Asia, particularly their utilization of mixed forest-grassland ecotones. The mosaic of deciduous broad-leaved forests and open grasslands would have provided both shelter and hunting opportunities, supporting a dynamic subsistence strategy.

    A preliminary study of protein preservation in animal fossils from the Hualongdong site in Dongzhi County, Anhui
    TANG Siyuan, LIU Sitong, ZHOU Kai, RAO Huiyun, WU Xiujie, FU Qiaomei
    2025, 44(05):  917-926.  doi:10.16359/j.1000-3193/AAS.2025.0069
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    The Hualongdong(HLD) site (30°06′34″N, 116°56′54″E, 40 m above sea level) is located in Dongzhi County, Anhui Province, China. Dated to 331~275 ka, it corresponds to the critical transition period from Homo erectus to early Homo sapiens in East Asia. Morphological studies of its hominin fossils suggest that these remains may represent the earliest pre-modern humans in the region.This study adopted a strategy of evaluating the protein preservation of animal fossils to preliminarily assess the overall protein preservation in the same site, aiming to investigate the feasibility of proteomic analysis for Hualongdong hominins without damaging hominin specimens. In this study, we analyzed 49 teeth and femoral bone specimens excavated from No.1 Site, which is the same site where the ancient human fossils were discovered.Mass spectrometry analysis detected no endogenous collagen markers in any of the 40 bone samples. Among nine dentine samples, two exhibited abundant collagen-specific peptide markers. Enamel protein analysis detected endogenous peptide signals in six out of seven enamel samples. These results indicate that part of dental specimens from Hualongdong site retain sufficient endogenous proteins to support large-scale screening of hominin fossils, thereby enabling further proteomic analysis of hominin fossils from this critical site to help track east Asian human evolution.