A new Homo erectus (Zhoukoudian V) brain endocast from China

A new Homo erectus endocast, Zhoukoudian (ZKD) V, is assessed by comparing it with ZKD II, ZKD III, ZKD X, ZKD XI, ZKD XII, Hexian, Trinil II, Sambungmacan (Sm) 3, Sangiran 2, Sangiran 17, KNM-ER 3733, KNM-WT 15 000, Kabwe, Liujiang and 31 modern Chinese. The endocast of ZKD V has an estimated endocranial volume of 1140 ml. As the geological age of ZKD V is younger than the other ZKD H. erectus, evolutionary changes in brain morphology are evaluated. The brain size of the ZKD specimens increases slightly over time. Compared with the other ZKD endocasts, ZKD V shows important differences, including broader frontal and occipital lobes, some indication of fuller parietal lobes, and relatively large brain size that reflect significant trends documented in later hominin brain evolution. Bivariate and principal component analyses indicate that geographical variation does not characterize the ZKD, African and other Asian specimens. The ZKD endocasts share some common morphological and morphometric features with other H. erectus endocasts that distinguish them from Homo sapiens.     

High-precision U-series dating of Locality 1 at Zhoukoudian, China

Thermal ionization mass spectrometric(230)Th/(234)U dating has been carried out on intercalated speleothem samples from the limestone cave occupied by Homo erectus at Zhoukoudian, China. The samples were recently collected in proper stratigraphic context after detailed field examinations. The results show that the age of the No. 5 Skull from Layer 3 is >400 ka, possibly in the range of about 400-500 ka, and that the hominid fossils from the lower strata are at least 600 ka and possibly >800 ka, much older than previously thought. The near-equilibrium(230)Th/(234)U ratios and internal consistency of the dates and stratigraphy lend credence to the results and allow us to comment on their important implications for human evolution.     

Aeretesmelanopterus (Pteromyinae, Rodentia) from Tianyuan Cave near Zhoukoudian (Choukoutien) in China

The materials of Aeretesmelanopterus from the Tianyuan Cave, a Late Pleistocene site with human fossils recovered recently near Zhoukoudian, are the richest of this species ever reported in North China. Aeretesmelanopterus is an endemic species of China, its relationship with Petaurista and Trogopterus is still controversal. Studies based on the dental characters show that Aeretesmelanopterus resembles Petaurista more closely than Trogopterus. The earliest fossil record of Aeretesmelanopterus was found in South China in Middle Pleistocene deposits. The earliest records in the Beijing area are from Upper Cave and Tianyuan Cave at Zhoukoudian are of late-Late Pleistocene age, around 30 kyrs BP. The geographical distributions of this species are very limited. In evolution, Aeretesmelanopterus experienced both the increase and the decrease in tooth size; but its crown height of cheek teeth was continuously increasing.     

Palaeopathological and variant conditions of the Homo heidelbergensis type specimen (Mauer,Germany)

Although early Homo specimens are now known from a number of African, Asian and European Middle Pleistocene sites, the taxon Homo heidelbergensis was initially introduced for the Mauer jaw recovered in 1907. Fossil hominids from the earlier Middle Pleistocene of Europe are very rare and the Mauer mandible is generally accepted as one of the most ancient, with an age of approximately 700 kyr. A new preparation of the mandible was conducted in 1996 and gave rise to the detailed palaeopathological examination which is presented here. Based on comparative analyses, the extreme breadth of the mandibular ramus and its flat intercondylar incision, in conjunction with the flattening and broadening of the coronoid process tip, results either from an idiosyncratic pattern of the course and insertion of the temporalis muscle on the coronoid process or from the temporalis possessing an accessory head. The incidence of periodontal pocketing, together with a vertical reduction of the alveolar margin to approximately 3.00 mm, and a slight protuberance formed in vicinity of the right M(2)can safely be interpreted as pathognomonic indications of periodontal disease. The short distance between the enamel-dentine junction of the teeth and the horizontal alveolar margins could either be an inherited variant or may result from incipient osteoporosis. In addition, an arthrotic condition with slight osteophytic peripheral exostoses and an arthrolit (i.e. an articular calculus or "joint mouse") on the left condylus articularisand a depression in the medial part of the left mandibular condyle extending into the inferior part of the ramus are present. These features are indicative of a trauma-induced osteochondrosis dissecans. The diagnosis therefore suggests that the observed depression results from a well-healed fracture. This traumatic event illustrates the demanding living conditions endured by humans during the European Middle Pleistocene. The variations and pathological conditions observed in Mauer do not question the mandible's role as type specimen for the taxon Homo heidelbergensis.     

A re-evaluation of the metric diversity within Homo erectus

Previous work by several researchers has suggested that the cranial sample from Zhoukoudian possesses a unique metric pattern relative to the African and Asian specimens assigned to Homo erectus. The current study readdresses this issue with an expanded fossil sample and a larger and more comprehensive set of cranial measurements. To test the patterns present in the assemblage, canonical variates analysis was performed using a covariance matrix generated from the Howells data set. From this, interindividual Mahalanobis distances were computed for the fossils. Random expectation statistics were then used to measure statistical significance of the Mahalanobis distances. The results show that the Zhoukoudian hominids exhibit a unique metric pattern not shared by the African and Indonesian crania sampled. In these tests the Hexian calvaria resembled the African and Indonesian specimens and differed significantly from the craniometric pattern seen in the Zhoukoudian fossils. The Zhoukoudian specimens are characterized by a wide midvault and relatively narrow occipital and frontal bones, while the African and Indonesian crania (including Hexian) have relatively broad frontal and occipital dimensions compared to their midvaults. These results do not suggest that a multiple-species scenario is necessary to encompass the variation present in the sample. Based on the current evidence it is more probable that this variation reflects polytypism influenced by environmental adaptation and/or genetic drift.     

New discoveries of Gigantopithecus blacki teeth from Chuifeng Cave in the Bubing Basin, Guangxi, south China

Ninety two Gigantopithecus blacki teeth have recently been recovered in situ at Chuifeng Cave in the Bubing Basin, Guangxi, south China. The hominoid teeth are associated with a typical early Pleistocene fauna. In comparison with Gigantopithecus of known provenience, for which classification is problematic, the Chuifeng sample can be allocated definitively to G. blacki. The new collection represents the largest sample of this species known, with the exception of the material from Liucheng. Statistical analysis of the mammal fauna indicates that G. blacki is one of the dominant elements, comprising 9% of the fauna. Eleven teeth (12% of the sample) of G. blacki are diagnosed as having caries. In addition, wear on M₃ shows that G. blacki was adapted to consume tough or fibrous food and this wear may potentially imply relative longevity. Further study of this large sample of Gigantopithecus will provide additional insight into the paleobiology of this extinct hominoid.     

Energetics in Homo erectus and other early hominins: the consequences of increased lower-limb length

Previous studies of daily energy expenditure (DEE) in hominin fossils have estimated locomotor costs using a formula that was based on six species, all 18 kg or less in mass, including no primates, and that has a number of other problems when applied in an ecological context. It is well established that the energetic cost of human walking is lower than that of representative mammals, particularly for individuals with long lower limbs. The current study reevaluates the daily energy expenditures of a variety of hominin species using more appropriate approaches to estimating locomotor costs. To estimate DEE for primates, I relied on published data on body mass, day range, and the percentage of time spent in various activities. Based on those data, I calculated a value for nonlocomotor DEE. I then used a variant of a method that I have suggested elsewhere to calculate the daily cost due to locomotion (DEEL) and summed the two to calculate total DEE. The more up-to-date methods for calculating the cost of travel result in lower estimates of this aspect of the energy budget than seen in previous studies. Values obtained here for DEE in various representatives of Australopithecus are lower than reported previously by around 200 kcal/day. Taking into account the greater economy of human walking, particularly the effect of the longer lower limbs found in many later Homo species, also results in lowered estimates of DEE. Elongation of the lower limbs in H. erectus reduced relative travel costs nearly 50% in comparison to A.L. 288-1 (A. afarensis). The present method for calculating DEE indicates that female H. erectus DEE was 84% greater than that of female Australopithecus; this disparity is even larger than that suggested by previous workers.     

Early Pleistocene Dicerorhinus sumatrensis remains from the Liucheng Gigantopithecus Cave, Guangxi, China

The Liucheng Gigantopithecus Cave is very famous for its rich mammalian fauna and Gigantopithecus fossils. The Gigantopithecus Cave Fauna has been regarded as the typical Early Pleistocene fauna in South China. The majority of the fossils unearthed has been studied and published during the past decades. The only group remaining unpublished is the rhinoceros, which is studied in this paper. The rhino materials available today are only mandibles and lower cheek teeth. The dimensions of the tooth rows and the isolated teeth fall well within the range of the recent Dicerorhinus sumatrensis. Therefore, the rhino fossils from the Liucheng Gigantopithecus Cave can be referred to this living species. The reduced lower incisors and the U-shaped lingual contour of the mandible also support this taxonomic determination. The rhino materials from the Liucheng Gigantopithecus Cave represent the earliest known record of this species and the smallest Pleistocene rhinocerotid in China.     

Preface: research at Liang Bua, Flores, Indonesia

Excavations at Liang Bua, Flores, Indonesia, have yielded evidence for an endemic human species, Homo floresiensis, a population that occupied the cave between ∼95-17 ka. This discovery has major implications for early hominin evolution and dispersal in Africa and Asia, attracting worldwide interest. This preface describes the rationale for the excavations in historical, geographical, and wider research contexts, as well as the methods used. It also introduces the other papers on aspects of Liang Bua research that feature in this edition of the Journal of Human Evolution.     

广西崇左木榄山智人洞的鼠科化石

最近在广西崇左木榄山智人洞遗址采集到1件具有现代智人解剖特征初始状态的下颌骨和丰富的哺乳动物化石。啮齿类中的鼠科化石有锡金小鼠(Mus pahari)、中华姬鼠(Apodemus draco)、黑线姬鼠(Apodemus agrarius)、似德氏狨鼠(Hapalomys cf.H.delacouri)、笔尾树鼠(Chiropodomys gliroides)、社鼠(Niviventer confucianus)、针毛鼠(Niviventer fulvescens)、爱氏巨鼠(Leopoldamys edwardsi)、印度板齿鼠(Bandicota indica)、褐家鼠(Rattus norvegicus)和黄毛鼠(Rattus losea)共8属11种。这种全部由当地或邻近地区现生种类构成的组合显然比当地早更新世的三合大洞和川黔地区中更新世歌乐山期的鼠类组合进步, 而与广西田东雾云洞的鼠类组合相似, 其动物群的时代为晚更新世早期, 与不平衡铀系法得到的测年结果(距今约11万年)相吻合。智人洞鼠科动物的组合明显具有东洋界热带-亚热带动物群的特点, 其中林灌和草地型所占比例较大表明当时的森林面积可能减少、林灌和草地面积可能增加, 反映出当时的气候相对干旱。     

The Upper Cave at Zhoukoudian and the origins of the Mongoloids

The best evidence for identifying the inhabitants of northeast Asia in the terminal Pleistocene or early Holocene periods is provided by the human burials from the Upper Cave at Zhoukoudian, and in particular the “Old Man”. Apart from the Minatogawa finds on Okinawa, all Late Pleistocene human remains from East Asia that are reasonably well published are poorly preserved and often have equivocal dates. Since the time the Upper Cave was excavated it has been supposed that the human remains were the link between “Peking Man” and the modern Mongoloid complex. We have carried out multivariate analyses of the “Old Man” cranium employing new measurements of Weidenreich's cast of the skull and comparative data from Howells' survey of modern human groups. Despite our expectations the analyses have not shown the “Old Man” to be closely linked with the Mongoloids. Considering all the evidence available we conclude that the common belief of a close biological relationship between the people buried in the Upper Cave and the modern Mongoloids is not yet adequately demonstrated. Of crucial importance in interpreting the Upper Cave burials is their antiquity, which is still commonly thought to be in the order of at least 18 000 years. We believe that recent C-14 dates of about 11 000 years, determined from animal bones, indicate the earliest possible date for the burials. The time span between the Upper Cave burials and the earliest known modern Mongoloids in north China is in the order of about 4–5000 years. It is possible that a major population shift has occurred in north China between the terminal Pleistocene and the mid Holocene, when farming first appears. If this is so, the Upper Cave people may not have been closely allied to the Mongoloid groups that now inhabit East Asia and the Americas.     

Cladistic analysis of early Homo crania from Swartkrans and Sterkfontein, South Africa

The phylogenetic relationships of early Pleistocene Homo crania from the South African sites of Swartkrans and Sterkfontein were investigated through cladistic analyses of 99 morphological characters. The Swartkrans Member 1 specimen SK 847 and the Stw 53 cranium from Sterkfontein Member 5A were treated as separate operational taxonomic units (OTUs), distinct from the three species of early Homo-H. erectus, H. habilis, and H. rudolfensis-that are recognized from the Plio-Pleistocene deposits of East Africa. The cladistic analyses differed in the treatment of the South African OTUs (separate Swartkrans and Sterkfontein OTUs vs. a single Swartkrans+Sterkfontein OTU). PAUP 4.0 was used to construct cladograms and address hypotheses about relationships. In the analysis that treated the South African specimens as a single OTU, the position of that OTU was stable as a separate branch on the Homo clade between H. rudolfensis and [H. habilis+(H. erectus+H. sapiens)]. When SK 847 and Stw 53 were treated as separate OTUs, the majority of most parsimonious trees indicated that they were positioned in similar positions as the combined South African Homo OTU; that is, as separate branches between H. rudolfensis and [H. habilis+(H. erectus+H. sapiens)], with the Swartkrans OTU generally occupying a more derived position. The position of the Sterkfontein OTU was more stable than that of the Swartkrans OTU, which was found in several other positions among the minimum length trees. Running the analyses with only those characters preserved by SK 847 and Stw 53 resulted in similar topologies for minimum length trees, although the positions of Stw 53, SK 847, and H. habilis exchanged places in some trees. In no case was an exclusive sister relationship between either South African OTU and a particular species of Homo supported statistically. Both South African OTUs differ from H. habilis in the fewest number of cladistic characters.     

贵州毕节麻窝口洞与智人伴生的长鼻类化石

最近在贵州毕节麻窝口洞发现了3枚古人类牙齿化石和伴生的哺乳动物群。其中,古人类牙齿经初步研究可归入解剖学上的现代人,而与古人类相伴的大、小哺乳动物化石,经初步鉴定共计8目20科43属53种。本文系统记述了该动物群中大哺乳动物的典型代表——长鼻类化石,共2属2种:东方剑齿象(Stegodon orientalis)和亚洲象(Elephas maximus)。麻窝口洞的长鼻类缺失我国南方早更新世的典型种类——中华乳齿象(Sinomastodon)和华南剑齿象(Stegodon huananensis),具有从典型的中更新世大熊猫-剑齿象动物群(Ailuropoda-Stegodon fauna)向晚更新世亚洲象动物群(Asian elephant fauna)过渡的特征。依动物群的性质和地貌地层的特征,毕节麻窝口洞的智人及伴生动物群的地质时代很可能为中更新世晚期或晚更新世早期,这与堆积物的光释光年代测定的初步结果(距今约11.2-17.8万年)基本吻合。麻窝口洞东方剑齿象与亚洲象的组合明显具有东洋界亚热带动物群的特点,指示温暖潮湿的气候,这些长鼻类与智人等生存于近水的森林和灌丛中,并镶嵌了一些草地。     

Gamma-ray spectrometric dating of late Homo erectus skulls from Ngandong and Sambungmacan, Central Java, Indonesia

Hominid fossils from Ngandong and Sambungmacan, Central Java, Indonesia, are considered to be the most anatomically derived and youngest representatives of Homo erectus. Nondestructive gamma-ray spectrometric dating of three of these Homo erectus skulls showed that all samples underwent uranium leaching. Nevertheless, we could establish minimum age estimates of around 40ka, with an upper age limit of around 60 to 70ka. This means that the Homo erectus of Java very likely survived the Toba eruption and may have been contemporaneous with the earliest Homo sapiens in Southeast Asia and Australasia.     

On the paleopathologic findings exhibited by the late Homo erectus of Ceprano, Italy

In March 1994, during excavation work for the construction of a motorway, a fragmented calvaria was discovered in the “Campo Grande” area near Ceprano, a town in southern Latium (Central Italy) about 55 miles from Rome. After reconstruction, the remain was recognized as belonging to aHomo erectus; it has been estimated that it goes back to the lowest middle Pleistocene. The calvaria exhibits two pathologic findings. The first is a congenital malformation of the sphenoidal sinus consisting in a deep, wide recess penetrating into the left greater wing as far as the sphenotemporal suture. The lesion was examined with the help of a cast whose features were compared with those described in some early and recent cases reported in the literature. The second finding is a healed, depressed fracture of the right brow ridge. The cause and mechanism of this lesion are discussed, and a mechanical approach has been used to provide information pertinent to the specific reason for the injury. The possibility that the hominid was butted by a large animal appears to be the most likely cause of the fracture.     

A closer look at the “Protopithecus” fossil assemblages: new genus and species from Bahia,Brazil

The recently extinct large-bodied New World monkey Protopithecus brasiliensis Lund 1836 was named based on a distal humerus and proximal femur found in the Lagoa Santa cave system in the southeastern Brazilian state of Minas Gerais. These bones are from an animal about twice the size of the largest extant platyrrhines. One hundred and seventy-five years later, a nearly complete skeleton was discovered in the Toca da Boa Vista caves in the neighboring state of Bahia and was allocated to the same taxon as it was the first platyrrhine fossil of comparable size found since the originals. Our detailed study of the equivalent elements, however, reveals important morphological differences that do not correspond to intraspecific variation as we know it in related platyrrhine taxa. The presence of both an expanded brachioradialis flange on the humerus and gluteal tuberosity on the femur of the Bahian skeleton distinguishes it from the Lagoa Santa fossil as well as from all other platyrrhines. Further cranial and postcranial evidence suggests a closer relationship of the former with the alouattine Alouatta, while the limited Lund material fits more comfortably with the ateline clade. Therefore, we propose to limit P. brasiliensis Lund to the distal humerus and proximal femur from Lagoa Santa and erect a new genus and species for the skeleton from Toca da Boa Vista. Cartelles coimbrafilhoi was a large-bodied frugivore with a relatively small brain and diverse locomotor repertoire including both suspension and climbing that expands the range of platyrrhine biodiversity beyond the dimensions of the living neotropical primates.