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Re-creating ancient hominid virtual endocasts by CT-scanning.

Probably the first radiographic study of human fossils, that by D. Gorganovic-Kramberger on Neandertal remains from Krapina, Croatia, was published in 1906, only 11 years after Röntgen announced the discovery of X-rays. Many subsequent studies on fossil hominids used regular clinical diagnostic radiological apparatus, as depicted in Atlas of Radiographs of Early Man by M.F. Skinner and G.H. Sperber (1982). Some specimens such as crania filled with heavily calcified matrix proved intractable. Ordinary radiographs of such specimens usually failed to reveal endocranial structure, as fossilized bone and calcified endocast were approximately equally radio-opaque. Thus, neither endocranial volume nor structural details were detectable. The only invasive method that could have been employed involved mechanical removal of the solid matrix, but this entailed hazards to the cranial vault and the destruction of the natural endocranial cast. In 1983--1984, G.C. Conroy and M. Vannier utilized recent advances in high-resolution computed tomography to produce non-invasive, intracranial capacity measurements of matrix-filled fossil skulls. They tried the method on two fossil mammal skulls filled with hard sandstone matrix (1984, Science 26:456-458), and then successfully applied it to a South African, matrix-filled cranium of the ancient hominid (hominin) species, Australopithecus africanus from Makapansgat (Conroy et al. 1990, Science 247:838-841). Details of the morphology of the endocranial surface of the braincase were revealed, including the pattern of venous sinus drainage in the posterior cranial fossa. A group based in St. Louis, Vienna, Paris, Rome, and Johannesburg has taken such studies further. Beautiful "virtual endocasts" have been produced on a large male specimen of A. africanus from Sterkfontein, South Africa, and the endocranial capacity has been determined (1998). The methods make it possible to re-create "virtual endocasts" of ancient hominids.

Cephalometry↗

Human taxonomic diversity in the pleistocene: does Homo erectus represent multiple hominid species?

Recently, nomina such as "Homo heidelbergensis" and "H. ergaster" have been resurrected to refer to fossil hominids that are perceived to be specifically distinct from Homo sapiens and Homo erectus. This results in a later human fossil record that is nearly as speciose as that documenting the earlier history of the family Hominidae. However, it is agreed that there remains only one extant hominid species: H. sapiens. Has human taxonomic diversity been significantly pruned over the last few hundred millennia, or have the number of taxa been seriously overestimated? To answer this question, the following null hypothesis is tested: polytypism was established relatively early and the species H. erectus can accommodate all spatio-temporal variation from ca. 1.7 to 0.5 Ma. A disproof of this hypothesis would suggest that modern human polytypism is a very recent phenomenon and that speciation throughout the course of human evolution was the norm and not the exception. Cranial variation in a taxonomically mixed sample of fossil hominids, and in a modern human sample, is analyzed with regard to the variation present in the fossils attributed to H. erectus. The data are examined using both univariate (coefficient of variation) and multivariate (determinant) analyses. Employing randomization methodology to offset the small size and non-normal distribution of the fossil samples, the CV and determinant results reveal a pattern and degree of variation in H. erectus that most closely approximates that of the single species H. sapiens. It is therefore concluded that the null hypothesis cannot be rejected.

Analysis of Variance↗

Squamotympanic fissure in the Ngandong and Sambungmacan hominids: a reply to Delson et al.

In the present study the course of the squamotympanic fissure of the mandibular fossa is examined in several fossil hominid specimens. While previous work by several authors had found that the course of this fissure was potentially autapomorphic for the Ngandong and Sambungmacan fossil samples from Indonesia, a more recent study performed by Delson et al. (Anat Rec 2001; 262:380--397) reported that a number of other fossils, including OH 9, Sangiran 4 and 17, and KNM WT 15000 were similar to the Ngandong and Sambungmacan specimens in the expression of this feature. To test these findings, those specimens purported to share similarities with Ngandong and Sambungmacan were examined. The results of this study indicate clearly differing morphologies between the Ngandong and Sambungmacan fossils, on the one hand, and the four African and Indonesian fossils named by Delson et al. (2001). Hence, the course of the squamotympanic fissure remains a potentially autapomorphic feature separating the Ngandong and Sambungmacan samples from the remaining specimens allocated to Homo erectus.

Animals↗

Assessment of the accuracy of dental enamel thickness measurements using microfocal X-ray computed tomography.

Tooth enamel thickness has long been an important character in studies of primate and especially hominin phylogeny, taxonomy, and adaptation. Current methods for accurately assessing enamel thickness involve the physical sectioning of teeth, because measurements of enamel thickness using some radiographic techniques are unreliable. However, because destructive methods limit sample sizes and access to important fossil specimens, it is desirable that they be replaced with nondestructive techniques. Although microfocal X-ray computed tomography (mCT) has been used recently in studies of enamel thickness, the accuracy of this technique has yet to be established. The present research compares physical sections to computer-generated mCT sections of teeth from a variety of primate and nonprimate, recent and fossil taxa to examine whether enamel thickness, tooth size, and diagenetic remineralization (fossilization) impact the ability of mCT to measure enamel thickness accurately. Results indicate that recent teeth of varying size and thickness are clearly and accurately depicted in mCT scans, with measurements from nearly identical planes in physical and mCT sections differing by 3-5%. A fossil papionin molar (ca. 2 Myr) was also accurately measured using mCT scans, although thinner enamel in much older therapsid (ca. 263-241 Myr) teeth could not be distinguished from dentine. mCT is thus an accurate technique for measuring enamel thickness in recent taxa, although heavily mineralized teeth pose an obstacle to the ability of mCT to distinguish dental tissues. Moreover, absolutely thin enamel (less than approximately 0.10 mm) is difficult to resolve adequately in raw mCT images based on pixel values alone. Therefore, caution must be exercised in the application of mCT to the study of fossilized teeth.

Animals↗

Human evolution.

The common ancestor of modern humans and the great apes is estimated to have lived between 5 and 8 Myrs ago, but the earliest evidence in the human, or hominid, fossil record is Ardipithecus ramidus, from a 4.5 Myr Ethiopian site. This genus was succeeded by Australopithecus, within which four species are presently recognised. All combine a relatively primitive postcranial skeleton, a dentition with expanded chewing teeth and a small brain. The most primitive species in our own genus, Homo habilis and Homo rudolfensis, are little advanced over the australopithecines and with hindsight their inclusion in Homo may not be appropriate. The first species to share a substantial number of features with later Homo is Homo ergaster, or 'early African Homo erectus', which appears in the fossil record around 2.0 Myr. Outside Africa, fossil hominids appear as Homo erectus-like hominids, in mainland Asia and in Indonesia close to 2 Myr ago; the earliest good evidence of 'archaic Homo' in Europe is dated at between 600-700 Kyr before the present. Anatomically modern human, or Homo sapiens, fossils are seen first in the fossil record in Africa around 150 Kyr ago. Taken together with molecular evidence on the extent of DNA variation, this suggests that the transition from 'archaic' to 'modern' Homo may have taken place in Africa.

Animals↗

Possible ctenophoran affinities of the Precambrian "sea-pen" Rangea.

The Namibian Kuibis Quartzite fossils of Rangea are preserved three-dimensionally owing to incomplete collapse of the soft tissues under the load of instantaneously deposited sand. The process of fossilization did not reproduce the original external morphology of the organism but rather the inner surface of collapsed organs, presumably a system of sacs connected by a medial canal. The body of Rangea had tetraradial symmetry, a body plan shared also by the White Sea Russian fossil Bomakellia and possibly some other Precambrian frond-like fossils. They all had a complex internal anatomy, smooth surface of the body, and radial membranes, making their alleged colonial nature unlikely. Despite a different style of preservation, the Middle Cambrian Burgess Shale frond-like Thaumaptilon shows several anatomical similarities to Rangea. The body plan of the Burgess Shale ctenophore Fasciculus, with its numerous, pinnately arranged comb organs, is in many respects transitional between Thaumaptilon and the Early Cambrian ctenophore Maotianoascus from the Chengjiang fauna of South China. It is proposed that the irregularly distributed dark spots on the fusiform units of the petaloid of Thaumaptilon represent a kind of macrocilia and that the units are homologous with the ctenophoran comb organs. These superficial structures were underlain by the complex serial organs, well represented in the fossils of Rangea. The Precambrian "sea-pens" were thus probably sedentary ancestors of the ctenophores.

Animals↗

Current research on the late Pliocene and Pleistocene deposits north of Homa Mountain, southwestern Kenya.

The late Pliocene and Pleistocene sediments of the Homa Peninsula in southwestern Kenya are richly fossiliferous, preserve Early Stone Age archaeological traces and provide one of the few paleoanthropological data sets for the region between the branches of the East African Rift Valley. This paper presents preliminary results of our ongoing investigation of late Pliocene and Pleistocene deposits at the localities of Rawi, Kanam East, Kanam Central and Kanjera. While fossils have been collected from the peninsula since 1911, little systematic effort has been made to place them into a broader litho-and chronostratigraphic framework. This project has conclusively demonstrated that fossils occur in good stratigraphic context at all of the study localities and that claims of sediment slumping (Boswell, 1935) have been greatly overstated (Behrensmeyer et al., 1995; Plummer & Potts, 1989). A provisional chronostratigraphic framework based on magneto- and biostratigraphy is presented here. We have revised the Plio-Pleistocene stratigraphy of the Rawi and Kanam gullies to include three formations: the Rawi, Abundu and Kasibos Formations. Based on magneto- and biostratigraphy, these formations are dated between approximately three and one m.y.a. (Gauss Chron-Jaramillo Subchron) (Cande & Kent, 1995). The Apoko Formation unconformably overlies the others and may be middle to late Pleistocene in age. All formations contain rich patches of fossils, and Acheulean artifacts have been surface collected from the Abundu and Kasibos Formations. Deposition of the fossil- and artefact-bearing sediments at Kanjera North began in the early Pleistocene and continued into the middle Pleistocene. Deposition at Kanjera South began over one million years earlier than previously thought, at approximately 2.2 m.y.a., and continued into the Olduvai Subchron (1.770-1.950 m.y.a.; Cande & Kent, 1995). Excavations have recovered Oldowan artefacts in association with well-preserved fossil fauna near the base of the sequence, the oldest archaeological traces yet known from southwestern Kenya.

Animals↗

Paleoenvironmental reconstruction of middle Miocene paleosols bearing Kenyapithecus and Victoriapithecus, Nyakach Formation, southwestern Kenya.

Paleosols in the middle Miocene (15 Ma) Nyakach Formation at Kaimogool, near Sondu, southwestern Kenya have yielded specimens of the early cercopithecoid Victoriapithecus macinnesi and the early kenyapithecine Kenyapithecus africanus, and can be used as evidence for the environmental mosaic occupied by these primates. Five distinct types of paleosols (pedotypes) are recognized in the Nyakach Formation section at Kaimogool South. The most common paleosols are reddish brown, silty calcareous profiles with blocky structure and large root traces (Ratong pedotype) which are interpreted as soils of well-drained, dry bushland or thicket (nyika). Weakly developed paleosols associated with paleochannels (Dhero pedotype) represent wooded grassland early in the ecological succession from streamside flooding. One of these paleosols has yielded a fossil flora of grasses and small-leaved dicots like those of modern semi-arid wooded grassland. Crumb structured, calcareous paleosols with iron-manganese nodules (Yom pedotype) are interpreted to represent seasonally waterlogged, wooded grassland (dambo or vlei). Thick, red clayey, calcareous paleosols with blocky ped structure and large root traces (Tut pedotype) are interpreted as soils of well-drained dry woodland. Other blocky-structured, gray to brown calcareous paleosols with iron-manganese nodules (Chido pedotype) are interpreted as soils of seasonally waterlogged, riparian dry woodland. Fossil soils, plants and gastropods are evidence of an unusually dry (300-500 mm mean annual precipitation) habitat for apes, consisting of a vegetational mosaic dominated by dry woodland, bushland and thickets with few areas of seasonally waterlogged grassland. Fossils of V. macinnesi are rare from Nyakach, but were found in paleosols representative of bushland and thicket habitats (Ratong). Fossils of the ape K. africanus were found within paleosols indicative of dry woodland (Tut). Other paleosol types representative of seasonally dry dambo grassland (Yom), colonizing grassland (Dhero) or riparian woodland (Chido) are also represented, but have not yet produced primate fossils.

Animals↗

Molecular clock calibrations and metazoan divergence dates.

It has recently been argued that living metazoans diverged over 800 million years ago, based on evidence from 22 nuclear genes for such a deep divergence between vertebrates and arthropods (Gu 1998). Two "internal" calibration points were used. However, only one fossil divergence date (the mammal-bird split) was directly used to calibrate the molecular clock. The second calibration point (the primate-rodent split) was based on molecular estimates that were ultimately also calibrated by the same mammal-bird split. However, the first tetrapods that can be assigned with confidence to either the mammal (synapsid) lineage or the bird (diapsid) lineage are approximately 288 million years old, while the first mammals that can be assigned with confidence to either the primate or the rodent lineages are 65 million years old, or 85 million years old if ferungulates are part of the primate lineage and zhelestids are accepted as ferungulate relatives. Recalibration of the protein data using these fossil dates indicates that metazoans diverged between 791 and 528 million years ago, a result broadly consistent with the palaeontological documentation of the "Cambrian explosion." The third, "external" calibration point (the metazoan-fungal divergence) was similarly problematic, since it was based on a controversial molecular study (which in turn used fossil dates including the mammal-bird split); direct use of fossils for this calibration point gives the absurd dating of 455 million years for metazoan divergences. Similar calibration problems affect another recent study (Wang et al. 1999), which proposes divergences for metazoans of 1000 million years or more: recalibrations of their clock again yields much more recent dates, some consistent with a "Cambrian explosion" scenario. Molecular clock studies have persuasively argued for the imperfection of the fossil record but have rarely acknowledged that their inferences are also directly based on this same record.

Animals↗

Spine anatomy reveals the diversity of catfish through time: a case study of Synodontis (Siluriformes).

Synodontis (Mochokidae, Siluriformes) is a freshwater catfish endemic to Africa. The 118 extant species are present in almost all hydrographic basins. Some species are restricted to a single stream, whereas others have a vast distribution. Synodontis is known in the fossil record since the Miocene, and its history depends on the connections among African basins through time. The identification of species in the fossil record is essential to reconstruct this historical pattern. Catfish pectoral and dorsal spines are robust, they preserve well and they form most of the fossil remains for the genus Synodontis. Unfortunately, the criteria for the identification of extant Synodontis species are not applicable to fossil specimens. Here, we define 11 original morphological characters that permit to discriminate four extant species from the Chad-Chari hydrographic system. Six of these characters are defined on pectoral spines and five on dorsal spines. We then show that these characters can be used successfully for identifying fossil specimens. In particular, we present a case study in which we identify Synodontis cf. schall and Brachysynodontis cf. batensoda in the hominid-bearing sector Toros-Menalla (Late Miocene, northern Chad). We show that spine anatomy can be a powerful tool to recognise catfish species through time and thus to identify historical diversity pattern.

Africa↗

Mapping and taphonomic analysis of the Homo erectus loci at Locality 1 Zhoukoudian, China.

From a detailed analysis of published and unpublished sources, we constructed a digitized three-dimensional, stratigraphically-controlled excavation grid of Zhoukoudian Locality 1 in order to assess the spatial relationships of the excavated materials. All 15 fossil Homo erectus loci were mapped on the grid. Meter cubes were used in excavation starting in 1934, and Loci H through O, established between 1934 and 1937, were mapped to within 1 m(3)vertical and horizontal provenience. Loci A through G, established between 1921 and 1933, were excavated in the northernmost part of Locality 1 by unmapped quarrying, but their stratigraphic levels were recorded. We could localize Loci A through G on the grid system by utilizing locations of remaining walls, stratigraphic sections, excavation reports, excavation maps, and photographs. Loci contained skeletal elements of Homo erectus individuals scattered over areas of the cave floor of up to 9 m in diameter. Scoring of taphonomic damage on the Homo erectus sample, as observed on casts and originals, demonstrates that 67% of the hominid sample shows bite marks or other modifications ascribed to large mammalian carnivores, particularly the large Pleistocene cave hyena, Pachycrocuta brevirostris. Virtually all of the remaining Homo erectus skeletal assemblage shows breakage consistent with this taphonomic pattern of fragmentation. Bioturbation by digging carnivores is the most likely explanation for a fragment of Homo erectus Skull XI discovered 1 m below its other conjoined portions in Locus L. Carbon on all the Homo erectus fossils from Locus G, a circumscribed area of 1-meter diameter, earlier taken to indicate burning, cooking, and cannibalism, is here interpreted as detrital carbon deposited under water, perhaps the result of hyaenid caching behavior. Locus G records the close stratigraphic and horizontal association of stone artifacts with Homo erectus and other vertebrate skeletal elements, an association that is seen at other loci as well. Layer 4 of the excavation contains equid cranial bone previously interpreted to have been burned while fresh. We here document that Locus B Homo erectus, including Skull I, is stratigraphically associated with this evidence, but at some 10-12 m distance. Even though the presence of wood-stoked fires and hearths is not supported by geochemical results, evidence of fire at Locality 1 in the form of burned bone is confirmed. Contextual relationships of fossil skeletal elements, relationships of carnivore damage and stone tool cutmarks on bone, and evidence of the burning of fresh bone associated with Homo erectus and stone tools support a model of transient hominid scavenging aided by the use of fire at the large hyenid den that became Zhoukoudian Locality 1. Although the original excavation catalogue from Locality 1, as well as a significant number of fossils and stone artifacts, were lost during World War II, catalogue numbers on the many surviving specimens can be used to locate fossils and artifacts within the three-dimensional grid provided in this paper.

Animals↗

Enamel thickness, microstructure and development in Afropithecus turkanensis.

Afropithecus turkanensis, a 17-17.5 million year old large-bodied hominoid from Kenya, has previously been reported to be the oldest known thick-enamelled Miocene ape. Most investigations of enamel thickness in Miocene apes have been limited to opportunistic or destructive studies of small samples. Recently, more comprehensive studies of enamel thickness and microstructure in Proconsul, Lufengpithecus, and Dryopithecus, as well as extant apes and fossil humans, have provided information on rates and patterns of dental development, including crown formation time, and have begun to provide a comparative context for interpretation of the evolution of these characters throughout the past 20 million years of hominoid evolution. In this study, enamel thickness and aspects of the enamel microstructure in two A. turkanensis second molars were quantified and provide insight into rates of enamel apposition, numbers of cells actively secreting enamel, and the time required to form regions of the crown. The average value for relative enamel thickness in the two molars is 21.4, which is a lower value than a previous analysis of this species, but which is still relatively thick compared to extant apes. This value is similar to those of several Miocene hominoids, a fossil hominid, and modern humans. Certain aspects of the enamel microstructure are similar to Proconsul nyanzae, Dryopithecus laietanus, Lufengpithecus lufengensis, Graecopithecus freybergi and Pongo pygmaeus, while other features differ from extant and fossil hominoids. Crown formation times for the two teeth are 2.4-2.6 years and 2.9-3.1 years respectively. These times are similar to a number of extant and fossil hominoids, some of which appear to show additional developmental similarities, including thick enamel. Although thick enamel may be formed through several developmental pathways, most Miocene hominoids and fossil hominids with relatively thick enamel are characterized by a relatively long period of cuspal enamel formation and a rapid rate of enamel secretion throughout the whole cusp, but a shorter total crown formation time than thinner-enamelled extant apes.

Animals↗

A new hominid from the Upper Miocene of Chad, Central Africa.

The search for the earliest fossil evidence of the human lineage has been concentrated in East Africa. Here we report the discovery of six hominid specimens from Chad, central Africa, 2,500 km from the East African Rift Valley. The fossils include a nearly complete cranium and fragmentary lower jaws. The associated fauna suggest the fossils are between 6 and 7 million years old. The fossils display a unique mosaic of primitive and derived characters, and constitute a new genus and species of hominid. The distance from the Rift Valley, and the great antiquity of the fossils, suggest that the earliest members of the hominid clade were more widely distributed than has been thought, and that the divergence between the human and chimpanzee lineages was earlier than indicated by most molecular studies.

Animals↗

Cratonia cotyledon gen. et sp. nov: a unique Cretaceous seedling related to Welwitschia.

The fossil history of most extant seed plant groups is relatively well documented. Cycads, conifers and Ginkgo all have an extensive fossil record, and the understanding of early angiosperm diversity is increasing. The Gnetales are an exception. Few macrofossils have been described, and character evolution within the group is poorly known. Cratonia cotyledon is a new gnetalean fossil from the Early Cretaceous Crato Formation of Brazil. This well-preserved seedling consists of two cotyledons, a feeder and a root. The leaf surface shows polygonal epidermal cells and apparently paracytic or actinocytic stomata. The cotyledons have a very specific venation pattern, shared only by Cratonia and Welwitschia, with parallel primary veins and secondary veins fusing to form inverted 'Y's between the main veins. Based on the 'Y'-venation and the presence of a feeder, we assign Cratonia to the Gnetum-Welwitschia clade. Fossil seedlings are unusual and this complete specimen with unambiguously welwitschioid characters is spectacular. Cratonia indicates that the evolutionary split between Gnetum and Welwitschia had occurred in the Early Cretaceous. Further, the close relationship between a West African plant and an east South American Early Cretaceous fossil is consistent with a major geological event: the rifting of the Gondwana continent.

Cotyledon↗

The role of mid-palaeozoic mesofossils in the detection of early bryophytes.

Recently discovered Silurian and Devonian coalified mesofossils provide an additional source of data on early embryophytes. Those reviewed in this paper are considered of some relevance to understanding the early history of bryophytes while highlighting the difficulties of recognizing bryophytes in often very fragmentary fossils. The first group comprises sporophytes in which terminal sporangia contain permanent dyads and tetrads. Such spores (cryptospores) are similar to those found dispersed in older Ordovician and Silurian strata, when they are considered evidence for a land vegetation of embryophytes at a bryophyte grade. The phylogenetic significance of plants, where the axes associated with both dyad- and tetrad-containing sporangia are branching, a character state not found in extant bryophytes, is discussed. The second group comprises axial fossils, many with occasional stomata, in which central conducting strands include G-type tracheids and a number of novel types of elongate elements not readily compared with those of any tracheophyte. They include smooth-walled, evenly thickened elongate elements as well as those with numerous branching +/- anastomosing projections into the lumen. Some of the latter bear an additional microporate layer, but the homogenized lateral walls between adjacent cells are never perforate. Such cells, which occur in various combinations in central strands, are compared with the leptoids and hydroids of mosses, hydroids of liverworts and presumed water-conducting cells in coeval Lower Devonian plants such as Aglaophyton. It is concluded that lack of information on the chemistry of their walls hampers sensible assessment of their functions and the affinities of the plants. Finally, a minute fossil, comprising an elongate sporangium in which a central cylindrical cavity containing spores and possible elaters terminates in a complex poral dehiscence apparatus, is used to exemplify problems of identifying early bryophytes. It is concluded that further progress necessitates the discovery of pre-Upper Silurian fossils with well-preserved anatomy, as well as a re-evaluation of criteria used to assess existing and new Devonian fossils for bryophyte affinity.

Biological Evolution↗

Recent paleoanthropological excavations of in situ deposits at Makapansgat, South Africa--a first report.

The Makapansgat Limeworks is a significant Pliocene site both for its sample of 35 hominin fossils as well as its wealth of fossil fauna. The lithological and paleontological successions reveal local environmental changes that are important for understanding the context of hominin evolution in southern Africa. Yet most of the site's fossils were found in dumps left behind by quarry operations, and the paleoecological interpretations rest upon debatable assumptions about the original fossil provenience. We have recently initiated systematic paleoanthropological excavations at Makapansgat to recover well provenanced fossils in order to: 1) assess whether faunal successions are discernable in the Makapansgat sequence; 2) assist environmental interpretations of the site; 3) and potentially recover the oldest hominins in South Africa, roughly coincident with Australopithecus afarensis in East Africa. This paper presents a summary of our current paleoenvironmental research at the Limeworks and preliminary results of ongoing in situ excavations.

Animals↗

The age of the angiosperms: a molecular timescale without a clock.

The age of the angiosperms has long been of interest to botanists and evolutionary biologists. Many early efforts to date the age of the angiosperms and evolutionary divergences within the angiosperm clade using a molecular clock have yielded age estimates that are grossly inconsistent with the fossil record. We investigated the age of angiosperms using Bayesian relaxed clock (BRC) and penalized likelihood (PL) approaches. Both of these methods allow the incorporation of multiple fossil constraints into the optimization procedure. The BRC method allows a range of values for among-lineage rate of substitution, from a nearly clocklike behavior to a condition in which each branch is allowed an optimal substitution rate, and also accounts for variation in molecular evolution across multiple genes. A topology derived from an analysis of genes from all three plant genomes for 71 taxa was used as a backbone. The effects on age estimates of different genes, single-gene versus concatenated datasets, and the inclusion and assumptions of fossils as age constraints were examined. In addition, the influence of prior distributions on estimates of divergence times was also explored. These results indicate that widely divergent age estimates can result from the different methods (198-139 million years ago), different sources of data (275-122 million years ago), and the inclusion of temporal constraints to topologies. Most dates, however, are between 180-140 million years ago, suggesting a Middle Jurassic-Early Cretaceous origin of flowering plants, predating the oldest unequivocal fossil angiosperms by about 45-5 million years. Nonetheless, these dates are consistent with other recent studies that have used methods that relax the assumption of a strict molecular clock and also agree with the hypothesis that the angiosperms may be somewhat older than the fossil record indicates.

Bayes Theorem↗

Population systematics of chimpanzees using molar morphometrics.

When dental morphological variation within extant species is used as a guideline to partition variation within fossil samples into species, the underlying assumption is that fossil species are equivalent to extant species. This is the case despite the fact that dental morphology, which is commonly used to differentiate fossil species, is rarely used to differentiate extant species. Aspects of external morphology, ecology, behavior, breeding patterns, and molecular structure that are used to delineate living species are generally not available for fossils. In this paper, the utility of dental evidence for sorting fossil samples into species is evaluated by testing whether molar occlusal morphology is capable of sorting populations of Pan into the species and subspecies already well-established by nondental evidence. The dentitions of 341 chimpanzee individuals, sampled from regions throughout equatorial Africa, were sorted into 16 populations using rivers to demarcate the boundaries between populations. Digital-imaging software was used to measure 15 traits on the occlusal surface of each upper molar and 19 on each lower molar. After applying size adjustments, size-transformed and untransformed variables were subjected to discriminant analysis, with separate analyses carried out for each molar type. Results indicate that populations of Pan troglodytes and Pan paniscus are well differentiated at all molar positions. Populations of P. t. verus are distinct from other populations of P. troglodytes. Populations of P. t. troglodytes and P. t. schweinfurthii show close dental similarity. A distinct population is recognized at the Nigeria-Cameroon border, indicating the presence of P. t. vellerosus. The concordance between the patterns of diversity recognized by this study and other molecular and nonmolecular studies indicates that paleontological species that are similar to species of Pan in terms of size and patterns of diversification may be differentiated using molar morphology.

Africa↗