Asian Fossils Research Articles

Cranial and mandibular anatomy of Plastomenus thomasii and a new time-tree of trionychid evolution

Trionychid (softshell) turtles have a peculiar bauplan, which includes shell reductions and cranial elongation. Despite a rich fossil record dating back to the Early Cretaceous, the evolutionary origin of the trionychid bauplan is poorly understood, as even old fossils show great anatomical similarities to extant species. Documenting structural detail of fossil trionychids may help resolve the evolutionary history of the group. Here, we study the cranial and mandibular anatomy of Plastomenus thomasii using µCT scanning. Plastomenus thomasii belongs to the Plastomenidae, a long-lived (Santonian–Eocene) clade with uncertain affinities among trionychid subclades. The skulls of known plastomenids are characterized by unusual features otherwise not known among trionychids, such as extremely elongated, spatulate mandibular symphyses. We use anatomical observations for updated phylogenetic analyses using both parsimony and Bayesian methods. There is strong support across methods for stem-cyclanorbine affinities for plastomenids. The inclusion of stratigraphic data in our Bayesian analysis indicates that a range of Cretaceous Asian fossils including Perochelys lamadongensis may be stem-trionychids, suggesting that many features of trionychid anatomy evolved prior to the appearance of the crown group. Divergence time estimates from Bayesian tip-dating for the origin of crown Trionychia (134.0 Ma) and Pan-Trionychidae (123.8 Ma) constrain the evolutionary time span during which the trionychid bauplan has evolved to a range of < 11 million years. Bayesian rate estimation implies high morphological rates during early softshell turtle evolution. If correct, plastomenids partially fill the stratigraphic gap which results from shallow divergence times of crown cyclanorbines during the late Eocene.

A ‘large and valuable’ Siwalik fossil collection in the archives of the Oxford University Museum of Natural History

ABSTRACT This article describes results from a review of South Asian fossils in the Oxford University Museum of Natural History. These materials include two early 19th century collections of fossils from the Siwalik Hills in India. While this assemblage was summarised in 1837 by William Buckland as ‘large and valuable collections of fossil bones’, it has remained largely unstudied and unpublished in any detail since collection. Here, as a precursor to a comprehensive re-evaluation, we establish a chronological and geographical context of one collection event, provide details of its donor, and outline its history after arrival in Oxford. We then describe select taxa in the collection, including a well-preserved maxilla and toothrow of the large extinct giraffid, Sivatherium giganteum, as a basis to justify our current understanding of the biostratigraphic affinity of the assemblage. Conservatively, the collection is a ‘classic’ Upper Siwalik Plio-Pleistocene fauna, possibly the first to be transported to the UK. While further analyses will realise the scientific potential of the fossils, the narrative of their journey from India to Oxford remains incomplete. Further investigation of the hidden history of the collection is warranted.

Resolving the "muddle in the middle": The case for Homo bodoensis sp. nov.

Recent developments in the field of palaeoanthropology necessitate the suppression of two hominin taxa and the introduction of a new species of hominins to help resolve the current nebulous state of Middle Pleistocene (Chibanian) hominin taxonomy. In particular, the poorly defined and variably understood hominin taxa Homo heidelbergensis (both sensu stricto and sensu lato) and Homo rhodesiensis need to be abandoned as they fail to reflect the full range of hominin variability in the Middle Pleistocene. Instead, we propose: (1) introduction of a new taxon, Homo bodoensis sp. nov., as an early Middle Pleistocene ancestor of the Homo sapiens lineage, with a pan‐African distribution that extends into the eastern Mediterranean (Southeast Europe and the Levant); (2) that many of the fossils from Western Europe (e.g. Sima de los Huesos) currently assigned to H. heidelbergensis s.s. be reassigned to Homo neanderthalensis to reflect the early appearance of Neanderthal derived traits in the Middle Pleistocene in the region; and (3) that the Middle Pleistocene Asian fossils, particularly from China, likely represent a different lineage altogether.

A mandible from the Middle Pleistocene Hexian site and its significance in relation to the variability of Asian Homo erectus

This study presents the first detailed morphological description and comparison of a Middle Pleistocene hominin mandibular fragment (PA 831) and associated teeth from the Hexian site in Eastern China. We aim to investigate where the Hexian mandible fits within the genus Homo variability in the light of an increased and better characterized Asian fossils record. Comparative samples include Pleistocene Homo mandibles and teeth from Africa, Asia, and Europe, as well as earlier African hominins (Australopithecus and early Homo) and Holocene recent humans. Both conventional morphological description and metric analysis were used. In addition, virtual reconstructions of the enamel dentine junction (EDJ) surface, pulp cavity, and roots with micro-CT were used to the mandible and teeth. The Hexian mandible is characterized by a plesiomorphic structural pattern for the Homo clade, with strong corpus robustness and a subparallel and low-positioned mylohyoid line that differentiates the swollen subalveolar planum from the shallow subalveolar fossa. Features that are derived compared to early Homo include a moderately curved dental arcade, a well-developed lateral prominence placed at the M2 -M3 level, and multiple mental foramina. The Hexian mandible's complex enamel surface and strong, stout root structure are primitive traits for the Homo clade. Finally, the highly crenulated "dendrite-like" EDJ found in the molars may represent a dental feature specific to the continental Asian Homo erectus, but more data is needed to confirm this. Mandibular and dental features indicate that the Hexian mandible and teeth differ from northern Chinese H. erectus and European Middle Pleistocene hominins, but show some affinities with the Early Pleistocene specimens from Africa (Homo ergaster) and Java (H. erectus), as well as the Middle-Late Pleistocene mandible from Penghu, Taiwan. Compared to contemporaneous continental Asian hominin populations, the Hexian fossils may represent the survival of a primitive hominin, with more primitive morphologies than other contemporaneous or some chronologically older Asian hominin specimens.

Global and local perspectives on cranial shape variation in Indonesian &lt;i&gt;Homo erectus&lt;/i&gt;

Homo erectus is among the best-represented fossil hominin species, with a particularly rich record in Indonesia. Understanding variation within this sample and relative to other groups of H. erectus in China, Georgia, and Africa is crucial for answering questions about H. erectus migration, local adaptation, and evolutionary history. Neurocranial shape is analyzed within the Indonesian sample, including representatives from Sangiran, Ngandong, Sambungmacan, and Ngawi, as well as a comparative sample of H. erectus from outside of Java, using three-dimensional geometric morphometric techniques. This study includes several more recently described Indonesian fossils, including Sambungmacan 4 and Skull IX, producing a more complete view of Indonesian variation than seen in previous shape analyses. While Asian fossils can be distinguished from the African/Georgian ones, there is not a single cranial Bauplan that distinguishes all Indonesian fossils from those in other geographic areas. Nevertheless, late Indonesian H. erectus, from sites such as Ngandong, are quite distinct relative to all other H. erectus groups, including earlier fossils from the same region. It is possible that this pattern represents a loss of genetic diversity through time on the island of Java, coupled with genetic drift, although other interpretations are plausible. A temporal pattern of diachronic change was identified within Indonesia for the posterior neurocranium such that younger Sangiran fossils more closely approached the Ngandong/Sambungmacan/Ngawi pattern, but there was not a linear trend of shape change from Sangiran to Sambungmacan to Ngandong, as has been suggested previously. The Sambungmacan 3 fossil, which often appears as a morphological outlier, fits the general pattern of late Indonesian vault shape, but has a more extreme expression of the shape trends for this group than other individuals.

Variations and asymmetries in regional brain surface in the genus Homo

Paleoneurology is an important field of research within human evolution studies. Variations in size and shape of an endocast help to differentiate among fossil hominin species whereas endocranial asymmetries are related to behavior and cognitive function. Here we analyse variations of the surface of the frontal, parieto-temporal and occipital lobes among different species of Homo, including 39 fossil hominins, ten fossil anatomically modern Homo sapiens and 100 endocasts of extant modern humans. We also test for the possible asymmetries of these features in a large sample of modern humans and observe individual particularities in the fossil specimens.This study contributes important new information about the brain evolution in the genus Homo. Our results show that the general pattern of surface asymmetry for the different regional brain surfaces in fossil species of Homo does not seem to be different from the pattern described in a large sample of anatomically modern H. sapiens, i.e., the right hemisphere has a larger surface than the left, as do the right frontal, the right parieto-temporal and the left occipital lobes compared with the contra-lateral side. It also appears that Asian Homo erectus specimens are discriminated from all other samples of Homo, including African and Georgian specimens that are also sometimes included in that taxon. The Asian fossils show a significantly smaller relative size of the parietal and temporal lobes. Neandertals and anatomically modern H. sapiens, who share the largest endocranial volume of all hominins, show differences when considering the relative contribution of the frontal, parieto-temporal and occipital lobes. These results illustrate an original variation in the pattern of brain organization in hominins independent of variations in total size. The globularization of the brain and the enlargement of the parietal lobes could be considered derived features observed uniquely in anatomically modern H. sapiens.

Morphométrie géométrique et classification phylogénétique, application à la question de l’origine d’Homo sapiens en Asie

The objective of this study was to find out whether or not there is a morphological gap between the fossils living in the last million years discovered in Asia. The geometric morphometric methods were applied to the skulls of 18 specimen Asian fossils dated 6,000 to 1,150,000 years. Landmarks were analyzed by Generalized Procrustes Analysis. Principal component analysis (PCA) and hierarchical classification methods (UPGMA and NJ) were applied to those samples in order to identify homogenous subgroups. The Mahalanobis distances between those subgroups have been calculated. Two phylogenetic analyses have been applied to the quantitative characters: the maximum likelihood approach and a cladistic analysis. A sample of gorilla and chimpanzees was used as the out-group. Classification and PCA analysis identified 2 subgroups in the fossil group: one with an Homo sapiens affinity and the other with an Homo erectus affinity. Both phylogenetic analyses identified Homo erectus as a monophyletic subgroup, but the fossil subgroup with an Homo sapiens affinity are scattered among the extant Homo sapiens. We finally concluded that the fossils belong to 2 different taxa. The first (fossil Homo sapiens) belong to the same taxon than extant Homo sapiens, and the second is attributed to Homo erectus.

The late Middle Pleistocene hominin fossil record of eastern Asia: Synthesis and review

Traditionally, Middle Pleistocene hominin fossils that cannot be allocated to Homo erectus sensu lato or modern H. sapiens have been assigned to different specific taxa. For example, in eastern Asia, these hominin fossils have been classified as archaic, early, or premodern H. sapiens. An increasing number of Middle Pleistocene hominin fossils are currently being assigned to H. heidelbergensis. This is particularly the case for the African and European Middle Pleistocene hominin fossil record. There have been suggestions that perhaps the eastern Asian late Middle Pleistocene hominins can also be allocated to the H. heidelbergensis hypodigm. In this article, I review the current state of the late Middle Pleistocene hominin fossil record from eastern Asia and examine the various arguments for assigning these hominins to the different specific taxa. The two primary conclusions drawn from this review are as follows: 1) little evidence currently exists in the eastern Asian Middle Pleistocene hominin fossil record to support their assignment to H. heidelbergensis; and 2) rather than add to the growing list of hominin fossil taxa by using taxonomic names like H. daliensis for northeast Asian fossils and H. mabaensis for Southeast Asian fossils, it is better to err on the side of caution and continue to use the term archaic H. sapiens to represent all of these hominin fossils. What should be evident from this review is the need for an increase in the quality and quantity of the eastern Asian hominin fossil data set. Fortunately, with the increasing number of large-scale multidisciplinary paleoanthropological field and laboratory research projects in eastern Asia, the record is quickly becoming better understood.

Evolutionary significance of cranial variation in Asian Homo erectus.

Homo erectus inhabited a wide geographic area of Asia, ranging from 40 degrees north latitude in China to 8 degrees south latitude in island Southeast Asia. Yet variation within Asian H. erectus and its relation to ecological and temporal parameters have been little studied. I synthesize the revised radiometric chronologies for hominid sites in Asia and their relation to new oxygen isotope curves (proxies for climatic fluctuations and landbridge connections). These data suggest substantial opportunities in the later Pleistocene for both regional isolation and gene flow between hominids in mainland and Southeast Asia. They also suggest that the most northerly located Chinese sites (Zhoukoudian and Nanjing) may have been occupied during sequential, interglacial periods. Probably reflecting these periods of isolation, nonmetric features and principal components analysis (PCA) of calvarial shape suggest regional differentiation between northern Asian and Southeast Asian H. erectus. The most recent Southeast Asian fossils (e.g., Ngandong) conform to the Southeast Asian pattern. Except perhaps in brain size, there is no evidence that the temporally intermediate Chinese fossils are intermediate in morphology between older and younger Indonesian fossils. In fact, northern Chinese calvaria are easier to exclude from the larger Asian H. erectus hypodigm than are the Ngandong fossils. The Chinese specimens differ from the others based on their narrower occipitals and frontals for their cranial size. The Chinese sample from Zhoukoudian alone is thus not a good proxy for the morphology and variation seen within Asian H. erectus. Both the Chinese and late Indonesian samples exhibit less variation than does the early Indonesian sample; this along with their shared morphological bauplan suggests a common origin and no more than subspecific differentiation. This shared morphology, despite regional differences, was likely maintained by the increasing intensity of multiple glaciations (and longer-lasting land bridge connections) between mainland and island Southeast Asia during the last million years.

Evidence from facial morphology for similarity of Asian and African representatives of Homo erectus.

It has been argued that Homo erectus is a species confined to Asia. Specialized characters displayed by the Indonesian and Chinese skulls are said to be absent in material from eastern Africa, and individuals from Koobi Fora and Nariokotome are now referred by some workers to H. ergaster. This second species is held to be the ancestor from which later human populations are derived. The claim for two taxa is evaluated here with special reference to the facial skeleton. Asian fossils examined include Sangiran 4 and Sangiran 17, several of the Ngandong crania, Gongwangling, and of course the material from Zhoukoudian described by Weidenreich ([1943] Palaeontol. Sin. [New Ser. D] 10:1-484). African specimens compared are KNM-ER 3733 and KNM-ER 3883 from Koobi Fora and KNM-WT 15000 from Nariokotome. Hominid 9 from Olduvai is useful only insofar as the brows and interorbital pillar are preserved. Neither detailed anatomical comparisons nor measurements bring to light any consistent patterns in facial morphology which set the African hominids apart from Asian H. erectus. Faces of the African individuals do tend to be high and less broad across the orbits. Both of the Koobi Fora crania but not KNM-WT 15000 have nasal bones that are narrow superiorly, while the piriform aperture is relatively wide. In many other characters, including contour of the supraorbital torus, glabellar prominence, nasal bridge dimensions, internasal keeling, anatomy of the nasal sill and floor, development of the canine jugum, orientation of the zygomaticoalveolar pillar, rounding of the anterolateral surface of the cheek, formation of a malar tubercle, and palatal rugosity, there is variation among individuals from localities within the major geographic provinces. Here it is not possible to identify features that are unique to either the Asian or African assemblages. Additional traits such as a forward sloping "crista nasalis," presence of a "sulcus maxillaris," a high (and massive) cheek coupled with some flexion of the malar pillar, and a posterior position for the incisive canal are present in all groups. These characters seem to be plesiomorphic, in comparison to the derived states evolved in later humans. Much or all of the variation in facial form can be attributed to sex dimorphism and/or local differentiation of populations within the Asian and African geographic regions. Metric differences among the fossils are comparable to those documented in a subset of recent H. sapiens, and there is no evidence that the Pleistocene specimens show greater dispersion than expected within a single species. This finding is generally in keeping with observations made on other parts of the cranium, lower jaw, and teeth. All of the hominids can be placed in H. erectus. Although its phylogenetic origins remain obscure, this lineage must be rooted in Africa. The species flourished for a long time. At several sites in China, H. erectus is known from deposits of the later Middle Pleistocene, while at Ngandong in Indonesia, archaic people may have survived even into the Late Pleistocene (Swisher et al. [1996] Science 274:1870-1874). The Ngandong fossils may record the last appearance of the lineage.

Evidence from facial morphology for similarity of Asian and African representatives of Hhomo erectus

It has been argued that Homo erectus is a species confined to Asia. Specialized characters displayed by the Indonesian and Chinese skulls are said to be absent in material from eastern Africa, and individuals from Koobi Fora and Nariokotome are now referred by some workers to H. ergaster. This second species is held to be the ancestor from which later human populations are derived. The claim for two taxa is evaluated here with special reference to the facial skeleton. Asian fossils examined include Sangiran 4 and Sangiran 17, several of the Ngandong crania, Gongwangling, and of course the material from Zhoukoudian described by Weidenreich ([1943] Palaeontol. Sin. [New Ser. D] 10:1–484). African specimens compared are KNM-ER 3733 and KNM-ER 3883 from Koobi Fora and KNM-WT 15000 from Nariokotome. Hominid 9 from Olduvai is useful only insofar as the brows and interorbital pillar are preserved. Neither detailed anatomical comparisons nor measurements bring to light any consistent patterns in facial morphology which set the African hominids apart from Asian H. erectus. Faces of the African individuals do tend to be high and less broad across the orbits. Both of the Koobi Fora crania but not KNM-WT 15000 have nasal bones that are narrow superiorly, while the piriform aperture is relatively wide. In many other characters, including contour of the supraorbital torus, glabellar prominence, nasal bridge dimensions, internasal keeling, anatomy of the nasal sill and floor, development of the canine jugum, orientation of the zygomaticoalveolar pillar, rounding of the anterolateral surface of the cheek, formation of a malar tubercle, and palatal rugosity, there is variation among individuals from localities within the major geographic provinces. Here it is not possible to identify features that are unique to either the Asian or African assemblages. Additional traits such as a forward sloping “crista nasalis,” presence of a “sulcus maxillaris,” a high (and massive) cheek coupled with some flexion of the malar pillar, and a posterior position for the incisive canal are present in all groups. These characters seem to be plesiomorphic, in comparison to the derived states evolved in later humans. Much or all of the variation in facial form can be attributed to sex dimorphism and/or local differentiation of populations within the Asian and African geographic regions. Metric differences among the fossils are comparable to those documented in a subset of recent H. sapiens, and there is no evidence that the Pleistocene specimens show greater dispersion than expected within a single species. This finding is generally in keeping with observations made on other parts of the cranium, lower jaw, and teeth. All of the hominids can be placed in H. erectus. Although its phylogenetic origins remain obscure, this lineage must be rooted in Africa. The species flourished for a long time. At several sites in China, H. erectus is known from deposits of the later Middle Pleistocene, while at Ngandong in Indonesia, archaic people may have survived even into the Late Pleistocene (Swisher et al. [1996] Science 274:1870–1874). The Ngandong fossils may record the last appearance of the lineage. Am J Phys Anthropol 106:61–85, 1998. © 1998 Wiley-Liss, Inc.

Dental remains from Atapuerca (Spain) I. Metrics

The metrical features (tooth diameters, indices and the premolar and molar size sequences) of 55 teeth from the “Sima de los Huesos” (Sierra de Atapuerca, Burgos, Spain) and belonging to at least nine individuals are analysed and compared with those of other fossil hominids. Metric analysis indicates that the traits of the Atapuerca dental sample correspond to a pattern of evolution, with size reduction of the permanent postcanine teeth and expansion in permanent anterior tooth size, characterizing the Middle Pleistocene populations. The tooth dimensions of Atapuerca are closer to those of European fossils such as Mauer, Montmaurin and Arago II, than to those of Middle Pleistocene North African and Asian fossils. The different tooth sizes which are observed in the sample could be attributed to sexual dimorphism, perhaps greater than that of living populations.