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Limits to randomness in paleobiologic models: the case of Phanerozoic species diversity.

The question of how random, or unconstrained, paleobiologic models should be is examined with a case study: Signor's (1982, 1985) inverse calculation of levels of marine species diversity through the Phanerozoic. His calculation involved an ingenious model that estimated species numbers and species abundances in the world oceans of the past by correcting known numbers of fossil species for variations in sedimentary rocks available for sampling and in effort paleontologists might devote to sampling. The model proves robust to changes in possible shapes of species-abundance distributions, but it is sensitive to alterations in the assumption that paleontologists collect fossils at random. If it is assumed that ease of collecting varies with age of sediment (with the Cenozoic offering easy sampling) or that paleontologists tend to seek out rarer fossils, results of the inverse calculation change. In particular, the magnitude of the calculated Cenozoic diversity increase always declines from the factor of about seven as originally reported to something considerably smaller. This leaves open the problem of the magnitude of Cenozoic increase in marine species diversity, awaiting better empirical data and, perhaps, more exacting models, random or otherwise.

Algorithms↗

On morphological clocks and paleophylogeography: towards a timescale for Sorex hybrid zones.

Phylogeography--the study of within-species phylogenetic and geographic divergence--has been primarily the domain of molecular evolutionists because molecular markers record population structure on smaller scales than do traditional morphological traits. But when geometric morphometrics are combined with distance-based phylogenetics molar shape divergence appears to record population-level phylogeny, a fact that allows extant and fossil populations to be combined in a single phylogeographic study. The European Sorex araneus complex--a genetically complicated group composed of multiple karyotypic races and species--illustrates the principle. The phylogeographic patterns revealed by molar shape broadly agree with scenarios based on molecular data and circumstantial evidence. Importantly, the inclusion of fossil samples of known age allows minimum divergence times to be inferred. Some races of S. araneus may have diverged more than 120,000 years ago, but others may have diverged less than 14,000. Supporting evidence that molar shape can be used to reconstruct phylogeographic relationships comes from strong correlations between molar shape distances and both phylogenetic divergence time and cytochrome b sequence divergence in datasets where these variables are known independently (fossil carnivorans from a well-constrained stratigraphic setting and shrew species of the genus Sorex, respectively). However, molar shape may have a 'saturation point' beyond which it is not applicable.

Animals↗

[Proterozoic history and present state of cyanobacteria].

The paper delves into the main regularities of the distribution of fossil microorganisms in Precambrian rocks, beginning from the Archean Eon about 3.5 billion years ago and ending in the Cambrian Period about 0.5 billion years ago. The paper analyzes facial peculiarities in the lateral differentiation of microfossils in Proterozoic basins and the main stages of temporal changes in fossil cyanobacterial communities, which are based on the irreversible succession of physicochemical conditions on the Earth and the evolution of eukaryotic microorganisms and their incorporation into prokaryotic ecosystems. To gain insight into Proterozoic fossil records, modern stratified cyanobacterial mats built up from layers of prokaryotes are considered. The analysis of phosphatization, carbonatization, and silification processes in modern algal-bacterial communities suggests that analogous processes took place in Proterozoic microbiotas. A comparison of modern and Precambrian living forms confirms the inference that cyanobacterial communities are very conservative and have changed insignificantly both morphologically and physiologically during the past two billion years.

Biological Evolution↗

Greenhouse gas production in wastewater treatment: process selection is the major factor.

Many practical design and operating decisions on wastewater treatment plants can have significant impacts on the overall environmental performance, in particular the greenhouse gas (GHG) emissions. The main factor in this regard is the use of aerobic or anaerobic treatment technology. This paper compares the GHG production of a number of case studies with aerobic or anaerobic main and sludge treatment of domestic wastewater and also looks at the energy balances and economics. This comparison demonstrates that major advantages can be gained by using primarily anaerobic processes as it is possible to largely eliminate any net energy input to the process, and therefore the production of GHG from fossil fuels. This is achieved by converting the energy of the incoming wastewater pollutants to methane which is then used to generate electricity. This is sufficient to power the aerobic processes as well as the mixing etc. of the anaerobic stages. In terms of GHG production, the total output (in CO2 equivalents) can be reduced from 2.4 kg CO2/kg COD(removed) for fully aerobic treatment to 1.0 kg CO2/kg COD(removed) for primarily anaerobic processes. All of the CO2 produced in the anaerobic processes comes from the wastewater pollutants and is therefore greenhouse gas neutral, whereas up to 1.4 kg CO2/kg COD(removed) originates from power generation for the fully aerobic process. This means that considerably more CO2 is produced in power generation than in the actual treatment process, and all of this is typically from fossil fuels, whereas the energy from the wastewater pollutants comes primarily from renewable energy sources, namely agricultural products. Even a change from anaerobic to aerobic sludge treatment processes (for the same aerobic main process) has a massive impact on the CO2 production from fossil fuels. An additional 0.8 kg CO2/kg COD(removed) is produced by changing to aerobic sludge digestion, which equates for a typical 100,000 EP plant to an additional production of over 10 t CO2 per day. Preliminary cost estimates confirm that the largely anaerobic process option is a fully competitive alternative to the mainly aerobic processes used, while achieving the same effluent quality.

Bacteria, Aerobic↗

Experimental taphonomy of velvet worms (Onychophora) and implications for the Cambrian "explosion, disparity and decimation" model.

Experimental preservation of velvet worms (phylum Onychophora), a very rare but evolutionarily important group that has existed for more than 500 million years, showed that the absence of bucal parts, adhesive-expelling organs, gonopore, eyes, legs, claws, annulation and papillation in fossils may not represent absence in the living animals. In fossils, leg thickness and claw orientation can be unreliable. The experiments indicate that not only absence, but even presence of certain structures can simply be the result of tissue decomposition. Computer-aided photorealistic reconstructions of fossi onychophorans are presented. We recommend future researchers to conduct taphonomy experiments specially before analysing unusual fossils.

Animals↗

Time flies, a new molecular time-scale for brachyceran fly evolution without a clock.

The insect order Diptera, the true flies, contains one of the four largest Mesozoic insect radiations within its suborder Brachycera. Estimates of phylogenetic relationships and divergence dates among the major brachyceran lineages have been problematic or vague because of a lack of consistent evidence and the rarity of well-preserved fossils. Here, we combine new evidence from nucleotide sequence data, morphological reinterpretations, and fossils to improve estimates of brachyceran evolutionary relationships and ages. The 28S ribosomal DNA (rDNA) gene was sequenced for a broad diversity of taxa, and the data were combined with recently published morphological scorings for a parsimony-based phylogenetic analysis. The phylogenetic topology inferred from the combined 28S rDNA and morphology data set supports brachyceran monophyly and the monophyly of the four major brachyceran infraorders and suggests relationships largely consistent with previous classifications. Weak support was found for a basal brachyceran clade comprising the infraorders Stratiomyomorpha (soldier flies and relatives), Xylophagomorpha (xylophagid flies), and Tabanomorpha (horse flies, snipe flies, and relatives). This topology and similar alternative arrangements were used to obtain Bayesian estimates of divergence times, both with and without the assumption of a constant evolutionary rate. The estimated times were relatively robust to the choice of prior distributions. Divergence times based on the 28S rDNA and several fossil constraints indicate that the Brachycera originated in the late Triassic or earliest Mesozoic and that all major lower brachyceran fly lineages had near contemporaneous origins in the mid-Jurassic prior to the origin of flowering plants (angiosperms). This study provides increased resolution of brachyceran phylogeny, and our revised estimates of fly ages should improve the temporal context of evolutionary inferences and genomic comparisons between fly model organisms.

Animals↗

Hominin brain evolution--new century, new directions.

The study of hominin brain evolution focuses on the interiors of fossilized braincases. Applications of recent three-dimensional computed tomography (CT) and magnetic resonance imaging (MRI) techniques for visualizing and measuring >>virtual endocasts<< from braincases in combination with advances in computer graphics and software for acquiring relevant data are transforming the way in which fossil skulls are analyzed, and improving the quality of paleoneurological investigations. Although CT imaging is preferred for fossil skulls, a novel method that combines high-resolution MRI of physical endocasts, electronic reconstruction of their missing parts, and warping of the resulting virtual endocasts is currently being developed and has great potential for future studies of hominin brain evolution. Applications of CT and MR techniques have already resulted in surprising new findings, which are briefly outlined. Exciting revelations about hominin brain evolution are expected as the 21st century unfolds.

Animals↗

Phylogeny, biogeography, and molecular dating of cornelian cherries (Cornus, Cornaceae): tracking Tertiary plant migration.

Data from four DNA regions (rbcL, matK, 26S rDNA, and ITS) as well as extant and fossil morphology were used to reconstruct the phylogeny and biogeographic history of an intercontinentally disjunct plant group, the cornelian cherries of Cornus (dogwoods). The study tests previous hypotheses on the relative roles of two Tertiary land bridges, the North Atlantic land bridge (NALB) and the Bering land bridge (BLB), in plant migration across continents. Three approaches, the Bayesian, nonparametric rate smoothing (NPRS), and penalized likelihood (PL) methods, were employed to estimate the times of geographic isolations of species. Dispersal and vicariance analysis (DIVA) was performed to infer the sequence and directionality of biogeographic pathways. Results of phylogenetic analyses suggest that among the six living species, C. sessilis from western North America represents the oldest lineage, followed by C. volkensii from Africa. The four Eurasian species form a clade consisting of two sister pairs, C. mas-C. officinalis and C. chinensis-C. eydeana. Results of DIVA and data from fossils and molecular dating indicate that the cornelian cherry subgroup arose in Europe as early as the Paleocene. Fossils confirm that the group was present in North America by the late Paleocene, consistent with the DIVA predictions that, by the end of the Eocene, it had diversified into several species and expanded its distribution to North America via the NALB and to Africa via the last direct connection between Eurasia and Africa prior to the Miocene, or via long-distance dispersal. The cornelian cherries in eastern Asia appear to be derived from two independent dispersal events from Europe. These events are inferred to have occurred during the Oligocene and Miocene. This study supports the hypothesis that the NALB served as an important land bridge connecting the North American and European floras, as well as connecting American and African floras via Europe during the early Tertiary.

Base Sequence↗

Isotopic biogeochemistry as a marker of Neandertal diet.

Natural abundances in 13C and 15N of bone collagen are linked to those of the diet. This isotopic signal can thus be linked to the dietary parameters of a given individual, such as the plants at the beginning of his food web and his position in the trophic web. In order to use this approach to study the diet of ancient humans, it is crucial to be sure that the original isotopic abundances of fossil collagen are preserved. This is done by controlling the biochemical purity of the organic matter extracted from fossil bones, and by checking that the isotopic differences observed in modern environments between herbivorous and carnivorous species are indeed measured in the fossil samples. Upper Pleistocene sites with a good isotopic preservation of collagen have been recognized in temperate and arctic environments. The isotopic signatures measured in such sites highlight particularities of the "mammoth steppe" fauna, and improve our knowledge of the diet of Neandertals.

Animals↗

Sexual dimorphic features within extant great ape faciodental skeletal anatomy and testing the single species hypothesis.

This paper examines sexually dimorphic skeletal characters within the face and upper dentition of extant hominids (great ape), not including members of the Hominini. Specimens of Pan paniscus, Pan troglodytes, Gorilla gorilla, and Pongo pygmaeus are used to help identify likely sex specific characters for the Hominidae. The aim of this paper is to identify extant hominid faciodental sexual features which can be used to help sex fossil specimens. A morphometric and skeletal study of sexual variability demonstrates relatively diverse patterns of sexual variability within the extant hominids. In terms of morphometrics, P. paniscus is relatively non-dimorphic, while P. troglodytes, Gorilla and Pongo display a large degree of sexual dimorphism. In their respective skeletal anatomies, however, each has specific characters which tend to differentiate between the sexes. Some faciodental sex features are shown to be common amongst all four taxa and as such are likely to be important criteria for determining the sex of Miocene and Plio-Pleistocene fossil hominid specimens. The construction of extant great ape sexual ranges of variability are also important in helping to test the fossil ape single species hypotheses. The testing of sex and species ranges of variability should employ range based statistics not only because they are sample size independent, (relative to C.V.) but also because they are of low power.

Animals↗

CO2 and the greenhouse effect: present assessment and perspectives.

Our present knowledge on the increasing greenhouse effect is based on the 1990 assessment of the Intergovernmental Panel on Climate Change (IPCC) and its 1992 supplement. Model predictions suggest that a doubling of atmospheric CO2 concentration would increase global temperature by 2-4 degrees C. Because time is needed for the upper layers of the oceans to warm up, there is a delay between the realized increase and the estimated equilibrium increase. The 0.5 degrees C increase in global temperature over the past 100 years is in accordance with model predictions but also within the range of natural variability. The IPCC's assessment suggests that if fossil fuel consumption continues according to a 'business as usual' scenario, global temperature will increase at 0.3 degrees C per decade; droughts, flooding and storms would become more frequent and more severe, and sea-level would continue to rise. Analysis of gas in ice cores from Antarctica and Greenland provides estimates of CO2 concentrations in pre-industrial ages; accurate measurement of atmospheric CO2 began in 1958. Atmospheric CO2 concentration has increased from 280 p.p.m. in AD 1800 to 355 p.p.m. at present. Between 1945 and 1973 global emissions of CO2 increased at a rate of 4.4% per year; after 1973 the rate of increase decreased to 1.6% per year. This change permits a test of the CO2 model. Reconstructed CO2 emission agrees within less than 10% with estimated fossil fuel-generated CO2 emission; contributions to CO2 emission from non-fossil fuel sources must be smaller than assumed previously. There are strong indications that in the past changes in atmospheric greenhouse gas concentrations inherent to the glacial-interglacial cycles did play an important climatic role. For example, they were probably responsible for interhemispheric coupling during the major climatic changes. Measures which might stabilize the greenhouse effect include energy conservation and improved energy efficiency, a transition to hydrogen rather than carbon as a source of fuel, and reforestation.

Air Pollutants↗

Functional anatomy of the olecranon process in hominoids and plio-pleistocene hominins.

This study examines the functional morphology of the olecranon process in hominoids and fossil hominins. The length of the bony lever of the triceps brachii muscle (TBM) is measured as the distance between the trochlear articular center and the most distant insertion site of the TBM, and olecranon orientation is measured as the angle that this bony lever makes with the long axis of the ulna. Results show that Homo, Pan, Gorilla, most monkeys, and the Australopithecus fossils studied have similar relative olecranon lengths. Suspensory hominoids and Ateles have shorter olecranons, suggesting, in some instances, selection for greater speed in extension. The orientation that the lever arm of the TBM makes with the long axis of the ulna varies with preferred locomotor mode. Terrestrial primates have olecranons that are more posteriorly oriented as body size increases, fitting general models of terrestrial mammalian posture. Arboreal quadrupeds have more proximally oriented lever arms than any terrestrial quadrupeds, which suggests use of the TBM with the elbow in a more flexed position. Olecranon orientation is not consistent in suspensory hominoids, although they are all characterized by orientations that are either similar or more posterior than those observed in quadrupeds. Homo and the fossils have olecranons that are clearly more proximally oriented than expected for a quadruped of their size. This suggests that Homo and Australopithecus used their TBM in a flexed position, a position most consistent with manipulatory activities.

Animals↗

Perikymata spacing and distribution on hominid anterior teeth.

We documented the spacing and distribution of perikymata on the buccal enamel surface of fossil hominin anterior teeth with reference to a sample of modern human and modern great ape teeth. A sample of 27 anterior teeth attributed to Australopithecus (5 to A. afarensis, 22 to A. africanus) and of 33 attributed to Paranthropus (6 to P. boisei, and 27 to P. robustus) were replicated and sputter-coated with gold to enable reflected light microscopy of their surface topography. Anterior teeth were then divided into 10 equal divisions of buccal crown height. The total perikymata count in each division of crown height was recorded using a binocular microscope fitted with a vernier micrometer eyepiece. Then the mean number of perikymata per millimeter was calculated for each division. Similar comparative data for a modern sample of 115 unworn human anterior teeth and 30 African great ape anterior teeth were collected from ground sections. Perikymata counts in each taxon (together with either known or presumed periodicities of perikymata) were then used to estimate enamel formation times in each division of crown height, for all anterior tooth types combined. The distributions of these estimates of time taken to form each division of crown height follow the same trends as the actual perikymata counts and differ between taxa in the same basic way. The distinction between modern African great apes and fossil hominins is particularly clear. Finally, we calculated crown formation times for each anterior tooth type by summing cuspal and lateral enamel formation times. Estimates of average crown formation times in australopiths are shorter than those calculated for both modern human and African great ape anterior teeth. The data presented here provide a better basis for exploring differences in perikymata spacing and distribution among fossil hominins, and provide the first opportunity to describe four specimens attributed to Homo in this context. Preliminary data indicate that differences may exist among the species attributed to early Homo, especially between Homo ergaster and Homo rudolfensis on the one hand, and Homo habilis sensu strico on the other.

Animals↗

Stereophotogrammetric analysis of occlusal morphology of extant hominoid molars: phenetics and function.

Because teeth are commonly preserved in the fossil record, dental remains have often been employed in estimating evolutionary relationships among fossil hominoids. This is appropriate, however, only to the extent that dental morphology is phylogenetically informative. I have used phenetic analytic techniques to assess whether hominoid molars are likely to be useful for phylogenetic inference. Thirty-four occlusal landmarks for first and second molars were chosen; seven on each maxillary and ten on each mandibular tooth. Three-dimensional locations of these points were determined from stereophotographs of dental arcades of more than 260 specimens from six taxa (gorilla, chimpanzee, human, orangutan, siamang, and gibbon). Analytic emphasis was on canonical variates analyses of landmark coordinates for mandibular and maxillary second molars, adjusted for intergroup size differences. There is little correspondence between the systematic implications of hominoid molar morphometrics and reliable estimates of evolutionary propinquity based on interhominoid biomolecular similarities. The former seem to have been determined largely by dietary constraints. Although this suggests the possibility of using the protocol employed here to infer diets of fossil hominoids, molar crown measurements seem unlikely to serve well as phylogenetic indicators in the Hominoidea.

Animals↗

Phylogenetic, paleodemographic, and taphonomic implications of Victoriapithecus deciduous teeth from Maboko, Kenya.

More than 240 milk teeth of Victoriapithecus macinnesi, representing all elements of the deciduous dentition, are described. Their morphology is intermediate between that of hominoids and cercopithecids. Unlike colobine and cercopithecine milk teeth, Victoriapithecus dp3s, dp4s, and at least 7 (14%) dp4s lack transverse distal loph(id)s and are not bilophodont. Victoriapithecus also differs from cercopithecids in having less elongated deciduous premolars, a dp3 metacone set mesial to a very small hypocone, a dp4 crista obliqua, and a dp4 hypoconulid. The deciduous canines and incisors of Victoriapithecus are like those of other cercopithecids in having an uneven distribution of enamel around a compressed (rather than cylindrical) root, but differ from cercopithecids and resemble hominoids in being more elongated. Since Colobinae and Cercopithecinae share features of the deciduous dentition that are derived relative to Victoriapithecus and hominoids, extant Old World monkeys are interpreted as representing the sister-taxon of the middle Miocene cercopithecoids. Due to a dramatic increase in the number of deciduous teeth found at Maboko Island, juvenile individuals represent a much larger proportion of recently excavated Victoriapithecus collections than is true of pre-1987 assemblages which mainly derive from sediment first excavated during the 1930s and 1940s. Age distribution differences between pre- and post-1987 samples indicate that paleontological collection procedures were more important than taphonomic biases in determining the paleodemographic profile of the Maboko fossils. Since the Victoriapithecus assemblage from Maboko is strikingly similar to that of fossil Theropithecus oswaldi from Olorgesailie in terms of the large number of specimens and high representation of juvenile and infant individuals, the latter can no longer be viewed as unique among cercopithecoid fossil assemblages. Rather than being related to a specific cause of death, such as selective hunting of T. oswaldi by Homo (Shipman et al. [1981] Curr. Anthropol. 22:257-268), the large number of cercopithecoids at both sites is attributed to the fact that both assemblages represent excavated samples and that the preferred habitats of the extinct monkeys were probably at or near the site of deposition. The greater number of young adult male than female canines in the apparently attritional Maboko Bed 5 assemblage, indicate that the social organization of V. macinnesi may have been similar to that of modern macaques, with males migrating out of their natal group and suffering higher death rates than females at puberty.

Animals↗

Brief communication: a new Pleistocene hominid-bearing locality at Hoedjiespunt, South Africa.

HDP1 is an archaeological and faunal site located on the Hoedjiespunt peninsula at Saldanha Bay, South Africa, that has recently yielded fossil human remains. Artefacts from the associated archaeological deposits are identified as being Middle Stone Age. U series analysis of capping calcretes and analysis of the foraminifera and fauna associated with the human fossils indicate an age for the deposit in excess of 74,000 years before present, and it most probably dates to around 300,000 years before present. The fossil human teeth from in situ deposits at Hoedjiespunt are described and found to be large by comparison with modern humans but smaller than the known upper dentitions of southern African "archaic" Homo sapiens. The Hoedjiespunt molars are found to be morphologically within the range of variation observed in the teeth of modern Homo sapiens.

Animals↗

Carnivoran paleoguilds of Africa: implications for hominid food procurement strategies.

Tool-using hominids, as carnivorous animals, would have been part of the various carnivore guilds present in Plio-Pleistocene Africa. Hominid dietary strategies must be understood within the larger context of carnivore behavior and ecology, as carnivorans could have affected the abilities of hominids to procure meat and/or marrow. The functional anatomy of extant and fossil carnivorans was examined to infer behaviors in fossil carnivorans that would have impacted on hominid dietary strategies in terms of carcass availability. Comparisons of guild structure were carried out to examine changes in carnivoran interactions and their implications for hominid behavior. Plio-Pleistocene carnivorans engaged in a wider range of behaviors than modern carnivorans. The sabertoothed felids Dinofelis and Megantereon probably did not provide much larger carcasses than modern species. Another sabertooth, Homotherium generated larger carcasses, but may have disarticulated and transported these carcasses. Fossil representatives of modern taxa may not have been equivalent ecologically within the carnivoran guild. Overall, hominids in eastern Africa probably had a greater range of scavenging opportunities than did those of southern Africa during the Plio-Pleistocene. Local and continent-wide extinction events in large-bodied carnivoran guilds from 1 to 2 Ma had a substantial effect on carcass availability and the risk to hominid scavengers. These structural changes in the carnivore guild may have provided an opportunity for hominids to widen their niche with respect to dietary behavior.

Africa↗

Combining isotopic and ecomorphological data to refine bovid paleodietary reconstruction: a case study from the Makapansgat Limeworks hominin locality.

The relationship between environmental change and hominin evolution remains obscure. For the most part, this stems from the difficulty of reconstructing ancient hominin habitats. Bovids are among the most frequently utilized paleoenvironmental indicators, but little is known about the habitat preferences of extinct taxa. It is generally assumed that fossil bovids both ate the same things and occupied the same habitats as their closest extant relatives. We test the first part of this assumption by reconstructing the diets of seven bovids from Makapansgat Limeworks, South Africa. Since diet and habitat are linked, these reconstructions have implications for our understanding of fossil bovid habitat tolerances. Ecomorphological and stable carbon isotope analyses are employed, allowing us to take advantage of the strengths and overcome the weaknesses of both. In most cases, fossil bovids did have similar diets to their extant relatives, and probably occupied similar habitats. Gazella vanhoepeni and Aepyceros sp., however, were almost exclusive browsers, and not mixed feeders like their living counterparts.

Animals↗