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Paranthropus boisei: an example of evolutionary stasis?

Of the presently recognised early hominid species, Paranthropus boisei is one of the better known from the fossil record and arguably the most distinctive; the latter interpretation rests on the numbers of apparently derived characters it incorporates. The species as traditionally diagnosed is distributed across approximately one million years and is presently confined to samples from East African sites. The hypodigm has been examined for evidence of intraspecific phyletic evolution, with particular emphasis on the areas best represented in the fossil record, namely the teeth and mandible. The results of this examination of 55 mandibular and dental variables which uses the gamma test statistic for the detection of trend, and nonparametric spline regression (Loess regression) for investigating pattern and rate of temporal change, show that within Paranthropus boisei sensu stricto most evidence of temporally related morphological trends relates to the morphology of the P4 crown. There is little or no evidence of any tendency to increase in overall size through time. Fossils from the Omo Shungura Formation and West Turkana which have been recovered from a time period earlier than the P. boisei sensu stricto hypodigm resemble the latter taxon in some features, but differ from it in aspects of cranial morphology. There is insufficient fossil evidence of the earlier taxon to tell whether it changes through time, but when trends of 47 mandibular and dental variables are assessed across the combined East African "robust" australopithecine sample, there is evidence for a relatively abrupt change around 2.2-2.3 Myr in approximately 25% of the dental and mandibular remains. Some of these changes correspond with the temporal trends within P. boisei sensu stricto, but others, such as mandible height, do not. If the earlier material is ancestral to P. boisei sensu stricto, evidence from the teeth and jaws is consistent with a punctuated origin for the latter taxon.

Africa, Eastern↗

Endocast of Sambungmacan 3 (Sm 3): a new Homo erectus from Indonesia.

A new fossil calvaria, Sambungmacan 3 (Sm 3), described in New Fossil Hominid Calvaria From Indonesia--Sambungmacan 3 by Márquez et al., this volume, yields one of the most advanced and complete endocasts yet recovered from Java. This communication provides a thorough interpretation of the external anatomical landmarks observable on Sm 3. Using computer tomography (CT) and traditional morphological measurements, our comparative paleoneurological analyses show that while Sm 3 has a mosaic of features that are similar to both Indonesian and Chinese H. erectus, it also possesses significant characters reminiscent of later hominins. These include a greater degree of asymmetry characterized by a possible left-occipital, right-frontal petalial pattern, left-right volumetric cerebral asymmetry, and marked asymmetry in Broca's cap. Moreover, the frontal lobe offers a more rounded, shortened appearance in contrast to the flat, elongated appearance of other Indonesian fossils (e.g., Sangiran 17). Another unique trait is exhibited in the transverse plane where the widest breadth of Sm 3 occurs more superiorly than in other Indonesian H. erectus. Thus, the endocast of Sm 3 presents a unique morphology not seen previously in the hominin fossil record. While the strong modern human characteristics in this endocast may not represent a particular ancestry, they do allow us to recognize a new dimension of the remarkable variation in Indonesian Homo erectus.

Animals↗

Reconstructing cetacean brain evolution using computed tomography.

Until recently, there have been relatively few studies of brain mass and morphology in fossil cetaceans (dolphins, whales, and porpoises) because of difficulty accessing the matrix that fills the endocranial cavity of fossil cetacean skulls. As a result, our knowledge about cetacean brain evolution has been quite limited. By applying the noninvasive technique of computed tomography (CT) to visualize, measure, and reconstruct the endocranial morphology of fossil cetacean skulls, we can gain vastly more information at an unprecedented rate about cetacean brain evolution. Here, we discuss our method and demonstrate it with several examples from our fossil cetacean database. This approach will provide new insights into the little-known evolutionary history of cetacean brain evolution.

Animals↗

Race and the odd history of human paleontology.

Although the late 17th century witnessed the recognition of fossils as the remains of extinct organisms-because they could be incorporated into the creation story embodied in the Great Chain of Being-acceptance of human antiquity through the indisputable demonstration of the contemporaneity of human bones, stone tools, and accepted fossils was not forthcoming for nearly 2 centuries thereafter. When it did occur, however, ancient humans were not seen as presenting a pattern of diversity similar to that seen in the fossil records of nonhuman organisms. Instead, human evolution then, as now, has typically been interpreted as being unilinear. This belief can be traced to Huxley (1863), who argued that the Feldhofer Grotto Neanderthal skullcap was merely an extension into the past of morphology seen in the Australian Aborigine, whom he took to represent the primitive end of an extreme range of variation he thought characterized Homo sapiens. During the mid-20th century, Mayr and Dobzhansky (mis)used their clout as founders of the evolutionary synthesis to cement in paleoanthropology the idea that human evolutionary history was characterized by nonspeciation. As such, anything that could be interpreted as potentially representing taxic diversity was relegated to the status of individual variation. Lack of understanding of the history of human paleontology, and the biases that constrained its perspective on human evolution, continue to affect the ways in which most paleoanthropologists pigeonhole human fossils.

Animals↗

Precambrian animal life: probable developmental and adult cnidarian forms from Southwest China.

The evolutionary divergence of cnidarian and bilaterian lineages from their remote metazoan ancestor occurred at an unknown depth in time before the Cambrian, since crown group representatives of each are found in Lower Cambrian fossil assemblages. We report here a variety of putative embryonic, larval, and adult microfossils deriving from Precambrian phosphorite deposits of Southwest China, which may predate the Cambrian radiation by 25-45 million years. These are most probably of cnidarian affinity. Large numbers of fossilized early planula-like larvae were observed under the microscope in sections. Though several forms are represented, the majority display remarkable conformity, which is inconsistent with the alternative that they are artifactual mineral inclusions. Some of these fossils are preserved in such high resolution that individual cells can be discerned. We confirm in detail an earlier report of the presence in the same deposits of tabulates, an extinct crown group anthozoan form. Other sections reveal structures that most closely resemble sections of basal modern corals. A large number of fossils similar to modern hydrozoan gastrulae were also observed. These again displayed great morphological consistency. Though only a single example is available, a microscopic animal remarkably similar to a modern adult hydrozoan is also presented. Taken together, the new observations reported in this paper indicate the existence of a diverse and already differentiated cnidarian fauna, long before the Cambrian evolutionary event. It follows that at least stem group bilaterians must also have been present at this time.

Animals↗

The temporal bones from Sima de los Huesos Middle Pleistocene site (Sierra de Atapuerca, Spain). A phylogenetic approach.

Three well-preserved crania and 22 temporal bones were recovered from the Sima de los Huesos Middle Pleistocene site up to and including the 1994 field season. This is the largest sample of hominid temporal bones known from a single Middle Pleistocene site and it offers the chance to characterize the temporal bone morphology of an European Middle Pleistocene population and to study the phylogenetic relationships of the SH sample with other Upper and Middle Pleistocene hominids. We have carried out a cladistic analysis based on nine traits commonly used in phylogenetic analysis of Middle and Late Pleistocene hominids: shape of the temporal squama superior border, articular eminence morphology, contribution of the sphenoid bone to the median glenoid wall, postglenoid process projection, tympanic plate orientation, presence of the styloid process, mastoid process projection, digastric groove morphology and anterior mastoid tubercle. We have found two autapomorphies on the Home erectus temporal bone: strong reduction of the postglenoid process and absence of the styloid process. Modern humans, Neandertals and the Middle Pleistocene fossils from Europe and Africa constitute a clade characterized by a convex superior border of the temporal squama. The European Middle Pleistocene fossils from Sima de los Huesos, Petralona, Steinheim, Bilzingsleben and Castel di Guido share a Neandertal apomorphy: a relatively flat articular eminence. The fossils from Ehringsdorf, La Chaise Suardi and Biache-Saint-Vaast also display another Neandertal derived trait: an anteriorly obliterated digastric groove. Modern humans and the African Middle Pleistocene fossils share a synapomorphy: a sagittally orientated tympanic plate.

Adult↗

The current state of korean paleoanthropology.

The hominid fossil and Paleolithic archaeology records from the Korean Peninsula are extensive, but relatively little is known about the Korean human evolutionary record outside this region. The Korean paleoanthropological record is reviewed here in light of major research issues, including the hominid fossil record, relative and chronometric dating, lithic analysis, hominid subsistence, and the presence of bone tools, art and symbolism. Some of the major conclusions drawn from this review include: (1) hominid fossils have been found in nine separate sites on the Korean Peninsula; (2) possible Homo erectus fossils are present in North Korea; (3) Ryonggok Cave, in North Korea, has exposed the remains of at least five archaic Homo sapiens individuals; (4) a possible burial of an anatomically modern Homo sapiens child, discovered in Hungsu Cave in South Korea, has been tentatively dated to roughly 40,000 years ago; (5) handaxes and cleavers have been found at a number of sites near Chongokni and they appear to date to at least 100,000 years ago; and (6) taphonomic studies are necessary for addressing issues related to determining the nature of hominid-carnivore interaction over similar resources (e.g. carcasses and shelter); and the presence/absence of Early Paleolithic bone tools, art, and symbolism in Korea.

Animals↗

The taxonomic status of the Chemeron temporal (KNM-BC 1).

Temporal bone morphology, as part of the basicranium, is commonly used in systematic evaluation of early hominid fossils. When an isolated right temporal bone, KNM-BC 1 (the Chemeron temporal) was discovered in the Baringo Basin, Kenya, Tobias (1967 a, Nature215, 476-480), citing ambiguity of characters, hesitated to place the specimen generically, attributing the fossil only to Hominidae gen. et sp. indet. Since that discovery, the early hominid sample has grown considerably and comparisons with this expanded dataset led Hill et al. (1992 a, Nature355, 719-722) to revise the placement of KNM-BC 1 including it within the genus Homo. This revision was possible due to the increased number of hominid fossil specimens from the late Pliocene/early Pleistocene, most notably members of the genus Homo. A thorough investigation into the utility of the temporal bone in hominid systematics shows that many features, as currently used in the literature, demonstrate high levels of variation thus questioning their phyletic valence. It is shown, however, that the temporal bone still contains useful systematic information. A detailed anatomical description of KNM-BC 1 is provided and, when discussed in the context of temporal bone features provided, affirms the conclusion of Hill et al. (1992 a) and places the fossil within the genus Homo.

Adult↗

New cercopithecoids and a hominoid from 12.5 Ma in the Tugen Hills succession, Kenya.

The early evolutionary history of the cercopithecoids is poorly understood, primarily due to a lack of fossil material from between 15 and about 9 Ma. Cercopithecoid primate specimens from a fossil site in the Ngorora Formation of the Tugen Hills, Kenya, belong to the genus Victoriapithecus, possibly a new species. These fossils are associated with a hominoid specimen that resembles Proconsul, and another tooth of a catarrhine, also probably hominoid. The locality is BPRP#38, in the Kabasero type section of the Ngorora Formation, and well dated at 12.5 Ma. If the hominoid specimen is confirmed as Proconsul, it would be one of the most recent recorded. The relatively diverse mammal fauna from the site in some ways resembles that of Fort Ternan. The site is underlain, and not far removed in time, from one of the best fossil macro-floras in Africa, which indicates lowland rainforest conditions in this portion of the Rift Valley at 12.6 Ma.

Animals↗

Primate phylogeny: morphological vs. molecular results.

Our comparative study of morphological (our data on selected living primates) and molecular characters (from the literature) confirms that, overall, phylogenetic reconstructions of Primates, and consequently their classifications, are more similar than dissimilar. When data from fossil Primates are incorporated, there may be several possible relationships among living Primates; the difference between most of them hinges mainly on the position of Tarsius. In one hypothesis, tarsiers are closely related to lemurs and lorises, and thus Primates is divided into Prosimii [lorises, lemurs, and tarsiers] and Anthropoidea [Platyrrhini and Catarrhini, i.e., monkeys, apes, and humans]. Two additional alternatives are that Tarsius is a sister group to the clade embracing lorises + lemurs and Anthropoidea and that in which all three lineages (Tarsius, lorises + lemurs, and Anthropoidea) form a polychotomy. In another hypothesis, tarsiers are closely related to anthropoids, giving these two branches: Strepsirhini [lemurs, lorises] and Haplorhini [tarsiers and Anthropoidea (Platyrrhini, the New World monkeys, and Catarrhini, Old World monkeys and Hominoidea)]. The first three alternatives gain some support from the fossil record, and the fourth from morphology of the living Tarsius and molecular data. It is emphasized that the morphological characters employed in this study for Tarsius are based on the only surviving genus of once-diverse tarsiiform primates known from the Eocene, and, although considered a "living fossil," it cannot represent all of them. Furthermore, Tarsius embodies derived features of its own which may affect its systematic position, but not necessarily the position of Tarsiiformes. Although the early Tertiary adapoids might have more nearly resembled anthropoids in their biochemistry and placental developments, this hypothesis is not testable from fossils, and any inferred relationships here must be based on characters of skeletal anatomy. Alternatively, anthropoids may be derived from certain omomyids or from some as yet undiscovered Eocene African taxon. Close relationships among Homo, Pan, and Gorilla have been confirmed during recent decades; Pongo is the sister group to this trichotomy. With increasing molecular data, Homo and Pan appear to be closer to each other than to any other living hominid taxon. Gorilla is a sister group to the Homo-Pan clade and Pongo is a sister group to all of them. Morphologists have given limited evidence for such a dichotomous grouping. In this study, we support the Homo-Pan clade, although with characters not as strong as for other clades.

Animals↗

What does palaeontology contribute to systematics in a molecular world?

Palaeontology provides the only direct record for morphological and genetic change through time and uniquely contributes to systematics in two ways: by providing access to denser taxon sampling than is otherwise possible and by dating divergence times. Claims that ancient DNA has survived millions of years in certain fossils suggested the possibility that palaeontology could contribute directly to molecular systematic studies. Unfortunately, none of the supposed geologically ancient DNA records stands up to detailed scrutiny and fossils therefore contribute primarily through the morphological information they preserve. Denser taxon sampling can improve the accuracy of phylogenetic estimates primarily through allowing better discrimination of homoplasy from homology. This in turn leads to more accurate hypotheses of character transformation. Denser taxon sampling also offers the opportunity for more accurate rooting, since more characters can be polarized by reference to a stem-group taxon than to an extant sister-group taxon. Missing data can be a problem for fossils, but is not crippling. Finally the temporal order of clade appearances in the fossil record can provide ancillary evidence for selecting a working phylogeny from among a number of equally most parsimonious cladograms.

Animals↗

A chicken-sized crane precursor from the early Oligocene of France.

A skeleton of a chicken-sized crane precursor is described from the Lower Oligocene of the Lubéron in Southern France. Parvigrus pohli gen. et sp. nov. is the most substantial Paleogene fossil record of the Grues (Aramidae [limpkin] + Gruidae [cranes]), and among its oldest representatives. The fossil species is classified in the new taxon Parvigruidae, which is shown to be the sister group of extant Grues. It is the first fossil record of a stem lineage representative of the Grues and, among others, differs from modern Grues in its smaller size, shorter beak, and rail-like limb proportions. Size increase in the stem lineage of the Gruidae may be related to the spread of grasslands during the Oligocene and Miocene. Occurrence of stem lineage Grues in the Lower Oligocene of Europe is in concordance with the fact that there is no evidence for the presence of crown group members of modern avian "families" in pre-Oligocene fossil deposits.

Animals↗

Fertilizers for food production vs energy needs and environmental quality.

The world is experiencing an energy crisis that is restrictive to agricultural requisites production at the same time that food is becoming increasingly short on a global basis. Fertilizers are the most energy demanding of these inputs and have become very expensive and intermittently short in supply with the reduced availability of fossil fuels. They have been indicted, furthermore, as environmental pollutants due to their presumed role in eutrophication and in being a source of excessive NO3-N that may accumulate in some leaf crops and in drinking waters. Exponential growth in fossil fuel consumption cannot continue. Economies can be made in the agricultural sector, which does indeed consume substantial quantities of energy. The energy consumed in this very essential food-producing process, however, is almost insignificant compared with that involved in transport and processing of food beyond the farm and with other energy expenditures in modern society. A shift in priorities will certainly be required in adapting to the real world of the 1970s if man's first need is to be met. Economies in fertilizer use can be made by adherence to known agronomic principles. Savings in fossil fuel energy can probably be effected also in the production of N fertilizer, by far the most fossil-energy-demanding process in the realm of agriculture. Considerable research remains to be done, however, under varied climatic conditions for understanding and controlling processes by which residuals from fertilizers may become environmental pollutants. The various issues in this paper must be resolved promptly in consideration of the now-existing energy crisis and the imminent world food crisis.

Air Pollution↗

The palaeoecology of the micromammals from the late middle Pleistocene site of Hoedjiespunt 1 (Cape Province, South Africa).

The palaeontological site of Hoedjiespunt 1 (HDP1) represents a fossilized hyaena lair. A rich mammalian fauna, including four hominid teeth, has been recovered from the site. Micromammals were recovered from the same sediments as the larger fauna. Taphonomic analysis suggests that the micromammal assemblages from HDP1 were accumulated by a barn owl. The barn owl produces micromammal assemblages that provide a broad sample of micromammals, within a certain size range, living in the hunting area of the owl. There are size-related and other biases inherent in the prey selection of this predator, and owls may roost in one area and hunt in another however, the barn owl has frequently been found to provide a better indication of micromammals living within an area than trapping. The micromammals from HDP1 were used to reconstruct the microhabitats in the vicinity of the site. Two taxonomic habitat indexes were used to assess the environment and dominant habitat types at Hoedjiespunt 1. The variability and adaptability of many of the southern African micromammals complicates interpretation of the results, however, it appears that the micromammals from the HDP1 fossil assemblages utilized habitats of open, scrub vegetation, and rocky and sandy areas. It is suggested that the environment was not markedly different from today, but it may have been relatively more arid. A comparison between HDP1 and other fossil sites in the area dating from the terminal Pleistocene to the Holocene indicates that HDP1 is lacking certain species that are common to all the other west coast fossil sites. There is some discrepancy in the environment indicated by the large mammals as compared that indicated by to the micromammals at the site. It is suggested that this discrepancy may reflect the fact that an owl is likely to have hunted in the vicinity of the hyaena den, probably in the more open areas around the roost site, whereas the macrofauna, accumulated by the further-ranging brown hyaena (Hyaena brunnea), represents environments from further afield.

Animals↗

Mandibular size and shape variation in the hominins at Dmanisi, Republic of Georgia.

The hominin fossils of Dmanisi, Republic of Georgia, present an ideal means of assessing levels of skeletal size and shape variation in a fossil hypodigm belonging to the genus Homo because they have been recovered from a spatially and temporally restricted context. We compare variation in mandible size and shape at Dmanisi to that of extant hominoids and extinct hominins. We use height and breadth measurements of the mandibular corpus at the first molar and the symphysis to assess size, and analyze shape based on size-adjusted (using a geometric mean) versions of these four variables. We compare size and shape variation at Dmanisi relative to all possible pairs of individuals within each comparative taxon using an exact resampling procedure of the ratio of D2600 to D211 and the average Euclidean distance (AED) between D2600 and D211, respectively. Comparisons to extant hominoids were conducted at both the specific and subspecific taxonomic levels and to extinct hominins by adopting both a more, and less speciose, hominin taxonomy. Results indicate that the pattern of variation for the Dmanisi hominins does not resemble that of any living species: they exhibit significantly more size variation when compared to modern humans, and they have significantly more corpus shape variation and size variation in corpus heights and overall mandible size than any extant ape species. When compared to fossil hominins they are also more dimorphic in size (although this result is influenced by the taxonomic hypothesis applied to the hominin fossil record). These results highlight the need to re-examine expectations of levels of sexual dimorphism in members of the genus Homo and to account for marked size and shape variation between D2600 and D211 under the prevailing view of a single hominin species at Dmanisi.

Animals↗

Cranial base morphology and temporal bone pneumatization in Asian Homo erectus.

The external morphological features of the temporal bone are used frequently to determine taxonomic affinities of fossils of the genus Homo. Temporal bone pneumatization has been widely studied in great apes and in early hominids. However, this feature is rarely examined in the later hominids, particularly in Asian Homo erectus. We provide a comparative morphological and quantitative analysis of Asian Homo erectus from the sites of Ngandong, Sambungmacan, and Zhoukoudian, and of Neandertals and anatomically modern Homo sapiens in order to discuss causes and modalities of temporal bone pneumatization during hominid evolution. The evolution of temporal bone pneumatization in the genus Homo is more complex than previously described. Indeed, the Zhoukoudian fossils have a unique pattern of temporal bone pneumatization, whereas Ngandong and Sambungmacan fossils, as well as the Neandertals, more closely resemble the modern human pattern. Moreover, these Chinese fossils are characterized by a wide midvault and a relatively narrow occipital bone. Our results support the point of view that cell development does not play an active role in determining cranial base morphology. Instead, pneumatization is related to available space and to temporal bone morphology, and its development is related to correlated morphology and the relative disposition of the bones and cerebral lobes. Because variation in pneumatization is extensive within the same species, the phyletic implications of pneumatization are limited in the taxa considered here.

Animals↗

Dental metric comparisons of Morotopithecus and Afropithecus: implications for the validity of the genus Morotopithecus.

Morotopithecus bishopi and Afropithecus turkanensis are two large-bodied hominoid primates from early Miocene deposits of eastern Africa. Researchers have used both cranial and postcranial characters to distinguish these two species. Unfortunately, of the fossil material attributed to each, only the face, palate, and upper dentition are preserved well enough in both species for direct comparisons. There are currently no known directly comparable postcranial elements. In this study, we reevaluated dental characters argued to distinguish the type specimens of Morotopithecus from Afropithecus: relative size of the upper premolars and M3. Exact randomization methods were used to address two questions. First, is it possible to find the degree of dental-size difference observed between Morotopithecus (UMP 62-11) and Afropithecus (KNM-WK 16999) within extant African hominoids? Second, what is the probability of observing the levels of difference found between the fossils among pairs of extant individuals? Metric differences in relative premolar and M3 size were calculated between all possible pairs within the extant sample and the observed difference of the fossil pair was then compared to the resulting distribution of extant pairs. The observed size differences for all comparisons in the fossil teeth were well within the variation observed in the extant African hominoid samples (p>0.05). In light of these results and other currently available cranial evidence, we suggest that the type specimens of Morotopithecus and Afropithecus are not different enough to support taxonomic distinction.

Africa, Eastern↗

Cercopithecoid cervical vertebral morphology and implications for the presence of Theropithecus in early Pleistocene Europe.

While it is recognized that the overall configuration of the vertebral column, as well as the size and shape of individual vertebrae, differ within and between primate taxa, relatively little is known about the degree to which vertebral morphology reflects a phylogenetic signal or the degree to which vertebral elements can be used in accurate taxonomic classification. Isolated vertebrae are occasionally found in fossil assemblages, and proper taxonomic identification is necessary to make inferences about the animal's biology and place it in a broader phylogenetic and evolutionary context. Recently, three large primate cervical vertebrae (C3, C5, and C6) from Pirro Nord, Italy (early Pleistocene, late Villafranchian) were attributed to the genus Theropithecus based on size comparisons with extant cercopithecoid primates (Rook et al., 2004, J. Hum. Evol. 47, 267-277). These fossils were suggested to indicate an early dispersal of this genus out of Africa around 1.6-1.3 Ma possibly co-incident with early Pleistocene dispersals of Homo. Because of the potential importance of these fossils for interpreting Theropithecus evolution and the relatively few morphological data on primate cervical vertebral morphology, we examined the size and shape of cervical vertebrae in a large sample of extant cercopithecoid taxa (n=106). Specifically, we evaluated whether subfamily and genus level assignments can be made on the basis of isolated cervical elements. Discriminant analyses reveal that scaled shape variables are good discriminators of taxonomic affinity at the subfamily level but are poor discriminators at the genus level. Least-squares regressions show that raw linear dimensions of cervical vertebral morphology are good predictors of body mass in the extant sample. Our regression results produce a likely body mass estimate of 22-38 kg for the Pirro Nord cervical vertebrae. Based on these regression estimates, the poor ability to discriminate cervical vertebrae at the genus level, and paleoenvironmental reconstructions of Pirro Nord, it is unlikely that the Pirro Nord fossils can be confidently attributed to the genus Theropithecus. These findings have important implications for recent interpretations of the nature of Theropithecus dispersal out of Africa.

Animals↗