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Limb-size proportions in Australopithecus afarensis and Australopithecus africanus.

Previous analyses have suggested that Australopithecus africanus possessed more apelike limb proportions than Australopithecus afarensis. However, due to the errors involved in estimating limb length and body size, support for this conclusion has been limited. In this study, we use a new Monte Carlo method to (1) test the hypothesis that A. africanus had greater upper:lower limb-size proportions than A. afarensis and (2) assess the statistical significance of interspecific differences among these taxa, extant apes, and humans. Our Monte Carlo method imposes sampling constraints that reduce extant ape and human postcranial measurements to sample sizes comparable to the fossil samples. Next, composite ratios of fore- and hindlimb geometric means are calculated for resampled measurements from the fossils and comparative taxa. Mean composite ratios are statistically indistinguishable (alpha=0.05) from the actual ratios of extant individuals, indicating that this method conserves each sample's central tendency. When applied to the fossil samples, upper:lower limb-size proportions in A. afarensis are similar to those of humans (p=0.878) and are significantly different from all great ape proportions (p< or =0.034), while Australopithecus africanus is more similar to the apes (p> or =0.180) and significantly different from humans and A. afarensis (p< or =0.031). These results strongly support the hypothesis that A. africanus possessed more apelike limb-size proportions than A. afarensis, suggesting that A. africanus either evolved from a more postcranially primitive ancestor than A. afarensis or that the more apelike limb-size proportions of A. africanus were secondarily derived from an A. afarensis-like ancestor. Among the extant taxa, limb-size proportions correspond with observed levels of forelimb- and hindlimb-dominated positional behaviors. In conjunction with detailed anatomical features linked to arboreality, these results suggest that arboreal posture and locomotion may have been more important components of the A. africanus behavioral repertoire relative to that of A. afarensis.

Animals↗

Nematode (Nematoda: Mermithidae) and hairworm (Nematomorpha: Chordodidae) parasites in Early Cretaceous amber.

The present report describes a mermithid nematode (Nematoda: Mermithidae) and a gordiid hairworm (Nematomorpha: Chordodidae) from Early Cretaceous Burmese amber dated at 100-110 million years. The mermithid, Cretacimermis protus sp. n., is emerging from a biting midge (Diptera: Ceratopogonidae) while the hairworm, Cretachordodes burmitis, gen. n., sp. n. had already emerged from its host. These rare specimens represent the first fossil mermithid parasite of a ceratopogonid midge and second oldest described nematode and the earliest known and only Mesozoic fossil of the phylum Nematomorpha. A list of previously described fossil mermithids is included.

Amber↗

Molecular systematics of the Hyaenidae: relationships of a relictual lineage resolved by a molecular supermatrix.

The four extant species of hyenas (Hyaenidae; Carnivora) form a morphologically and ecologically heterogeneous group of feliform carnivorans that are remnants of a formerly diverse group of mammalian predators. They include the aardwolf (Proteles cristatus), a termite-feeding specialist, and three species with a craniodental morphology adapted to cracking the bones of prey and/or carcasses, the spotted hyena (Crocuta crocuta), brown hyena (Parahyaena brunnea), and striped hyena (Hyaena hyaena). Hyenas have been the subject of a number of systematic studies during the last two centuries, due in large part to the extensive fossil record of the group, with nearly 70 described fossil species. Morphological studies incorporating both fossil and living taxa have yielded different conclusions regarding the evolutionary relationships among living hyenas. We used a molecular supermatrix comprised of seven nuclear gene segments and the complete mitochondrial cytochrome b gene to evaluate phylogenetic relationships among the four extant hyaenid species. We also obtained sequence data from representative species of all the main families of the Feliformia (Felidae, Herpestidae, and Viverridae) to estimate the sister group of the Hyaenidae. Maximum parsimony and maximum likelihood analyses of the supermatrix recovered identical topologies. Furthermore, Bayesian phylogenetic analyses of the supermatrix, with among-site rate variation among data partitions parameterized in three different ways, also yielded the same topology. For each phylogeny reconstruction method, all but two nodes received 100% bootstrap or 1.00 posterior probability nodal support. Within the monophyletic Hyaenidae, Parahyaena and Hyaena were joined together, with Crocuta as the sister to this clade, and Proteles forming the most basal lineage. A clade containing two species of mongoose (core Herpestidae) plus Cryptoprocta ferox (currently classified in Viverridae) was resolved as the sister group of Hyaenidae. The pattern of relationships among the three bone-cracking hyaenids (Crocuta, Hyaena, and Parahyaena) is incongruent with recent cladistic assessments based on morphology and suggests the need to reevaluate some of the morphological characters that have been traditionally used to evaluate relationships among hyenas. Divergence time estimates based on a Bayesian relaxed molecular clock indicates that hyaenids diverged from their feliform sister group 29.2 MYA, in the Middle Oligocene. Molecular clock estimates also suggest that the origin of the aardwolf is much more recent (10.6 MYA) than that implied by a cladistic analysis of morphology ( approximately 20 MYA) and suggests that the aardwolf is possibly derived from a bone and meat eating lineage of hyaenids that were present in the Late Miocene. [Hyaenidae; phylogeny; cytochrome b; nuclear gene segments; Proteles; Crocuta; Hyaena; Parahyaena.].

Animals↗

Age at first molar emergence in early Miocene Afropithecus turkanensis and life-history evolution in the Hominoidea.

Among primates, age at first molar emergence is correlated with a variety of life history traits. Age at first molar emergence can therefore be used to broadly infer the life histories of fossil primate species. One method of determining age at first molar emergence is to determine the age at death of fossil individuals that were in the process of erupting their first molars. This was done for an infant partial mandible of Afropithecus turkanensis (KNM-MO 26) from the approximately 17.5 Ma site of Moruorot in Kenya. A range of estimates of age at death was calculated for this individual using the permanent lateral incisor germ preserved in its crypt, by combining the number and periodicity of lateral enamel perikymata with estimates of the duration of cuspal enamel formation and the duration of the postnatal delay in the inception of crown mineralization. Perikymata periodicity was determined using daily cross striations between adjacent Retzius lines in thin sections of two A. turkanensis molars from the nearby site of Kalodirr. Based on the position of the KNM-MO 26 M(1)in relation to the mandibular alveolar margin, it had not yet undergone gingival emergence. The projected time to gingival emergence was estimated based on radiographic studies of M(1)eruption in extant baboons and chimpanzees. The estimates of age at M(1)emergence in KNM-MO 26 range from 28.2 to 43.5 months, using minimum and average values from extant great apes and humans for the estimated growth parameters. Even the absolute minimum value is well outside the ranges of extant large Old World monkeys for which there are data (12.5 to <25 months), but is within the range of chimpanzees (25.7 to 48.0 months). It is inferred, therefore, that A. turkanensis had a life history profile broadly like that of Pan. This is additional evidence to that provided by Sivapithecus parvada (Function, Phylogeny, and Fossils: Miocene Hominoid Evolution and Adaptations, 1997, 173) that the prolonged life histories characteristic of extant apes were achieved early in the evolutionary history of the group. However, it is unclear at present whether life-history prolongation in apes represents the primitive catarrhine pace of life history extended through phyletic increase in body mass, or whether it is derived with respect to a primitive, size-adjusted life history that was broadly intermediate between those of extant hominoids and cercopithecoids. Life history evolution in primates as a whole may have occurred largely through a series of grade-shifts, with the establishment of fundamental life-history profiles early in the histories of major higher taxa. These may have included shifts that were largely body mass dependent, as well as those that occurred in the absence of significant changes in body mass.

Adaptation, Physiological↗

Really old-palaeoimmunology: immunohistochemical analysis of extracellular matrix proteins in historic and pre-historic material.

In this review, we summarize data concerning the respective preservation and deterioration of antigenic determinants of various collagenous and non-collagenous extracellular matrix (ECM) proteins in palaeontologic material of different ages. ECM proteins are the major quantitative constituents of mammalian organisms and were, therefore, selected as important representative proteins for these analyses. The specimens, studied by immunofluorescence and immunohistochemical techniques, included the skin of 500-1500 year-old human mummies from Peru, skin and striated muscle from the 5300-year-old glacier mummy ("Iceman") from Tyrol, Austria, and a 50-million-year-old bat with preserved soft body parts from the fossil excavation site of Messel, Germany. In frozen sections of the former two sources, epitopes recognized by specific antibodies for triple-helical antigenic determinants of different types of collagen resistant against conventional proteases were preserved, while non-helical domains, as well as the non-collagenous ECM proteins, could no longer be demonstrated. The fossil bat, although showing evidence of fibrous, collagen-like structures in conventional histology, revealed no collagenous or non-collagenous ECM proteins by any technique. It later turned out that this was due to the replacement of the original soft parts in these fossils by lawns of bacteria. These studies introduced immunological techniques into palaeontology and opened new approaches for studying physiologically- and pathologically-altered structures in tissues of animals and humans of considerable historical age.

Allergy and Immunology↗

Hominid thumb strength predicted by high resolution magnetic resonance imaging and force measurements in living subjects.

This study investigates whether in-vivo measurements of the morphology of the pollical distal phalanx (PDP) can be used to predict the maximum force that can be exerted by the thumb in flexion at the interphalangeal joint. A predictive equation was obtained via which measurements of fossil PDP's can be used to predict flexion force in hominids. A home-built insertable coil assembly, which increased the available magnetic field gradient strength from 10 to 40 (mT) m-1, was used to acquire high resolution T1-weighted 3D SPGR Magnetic Resonance (MR) images of the left and right thumbs of nine volunteers. The subjects' age, sex, weight, height and self-assessed hand-dominance were recorded. The MR images were transferred to a computer for reformatting of sections in the transverse and volar plane (VP). Measurements were made of the maximum length (ML), breadth (MB), tuft breadth (MT) and joint depth (JD) of the PDP. For each subject the maximum flexion force that could be exerted by each thumb was measured. The mean maximum flexion force generated was 8.21 kg (range 5.68-13.13 kg). Considerable inter-individual variability was observed in the magnitude of the force difference between the left and right thumbs and this is greatest in individuals with the strongest thumbs. The best model produced by multiple linear regression analysis of all the available data has a value of r = 0.77 and needs only the two covariates of sex and weight, i.e. FORCE = 2.217 + 0.6651 x WEIGHT + 2.488 x SEX. However, in order to be useful for predicting the maximum flexion force of the thumb in hominids the model should only contain those parameters which can be measured for fossil PDP's. Accordingly, the best predictive model has a value of r = 0.73 and needs only the two covariates of ML and JD, i.e. FORCE = -10.28 + 0.2053 x ML + 1.571 x JD. The maximum force predicted for any of the hominid fossil PDP's was not significantly different from those recorded for the volunteers in the present study, although they were on average somewhat lower than those obtained for modern humans. This may be a reflection of the hominid's overall smaller body weights.

Animals↗

Metazoan phylogenies: falling into place or falling to pieces? A palaeontological perspective.

Metazoan phylogeny is in a state of ferment, stirred by the addition of new molecular trees as well as controversial interpretations of molecular 'clocks'. Concerning the latter topic, the clocks recurrently point to divergence times substantially older than the known fossil record. Some attempt reconciliation by appealing to a conveniently cryptic interval prior to the first fossils. This effectively reduces the fossil record to an erratic search-light giving only glimpses into the true evolutionary history. Other options, however, remain open. Molecular clocks may themselves run erratically and what happens in molecular history may not coincide with the emergence of body plans.

Animals↗

The radiation of the Cape flora, southern Africa.

The flora of the south-western tip of southern Africa, the Cape flora, with some 9000 species in an area of 90,000 km2 is much more speciose than can be expected from its area or latitude, and is comparable to that expected from the most diverse equatorial areas. The endemism of almost 70%, on the other hand, is comparable to that found on islands. This high endemism is accounted for by the ecological and geographical isolation of the Cape Floristic Region, but explanations for the high species richness are not so easily found. The high species richness is accentuated when its taxonomic distribution is investigated: almost half of the total species richness of the area is accounted for by 33 'Cape floral clades'. These are clades which may have initially diversified in the region, and of which at least half the species are still found in the Cape Floristic Region. Such a high contribution by a very small number of clades is typical of island floras, not of mainland floras. The start of the radiation of these clades has been dated by molecular clock techniques to between 18 million years ago (Mya) (Pelargonium) and 8 Mya (Phylica), but only six radiations have been dated to date. The fossil evidence for the dating of the radiation is shown to be largely speculative. The Cenozoic environmental history of southern Africa is reviewed in search of possible triggers for the radiations, climatic changes emerge as the most likely candidate. Due to a very poor fossil record, the climatic history has to be inferred from larger scale patterns, these suggest large-scale fluctuations between summer wet (Palaeocene, Early Miocene) and summer dry climates (Oligocene, Middle Miocene to present). The massive speciation in the Cape flora might be accounted for by the diverse limitations to gene flow (dissected landscapes, pollinator specialisation, long flowering times allowing much phenological specialisation), as well as a richly complex environment providing a diversity of selective forces (geographically variable climate, much altitude variation, different soil types, rocky terrain providing many micro-niches, and regular fires providing both intermediate disturbances, as well as different ways of surviving the fires). However, much of this is based on correlation, and there is a great need for (a) experimental testing of the proposed speciation mechanisms, (b) more molecular clock estimates of the age and pattern of the radiations, and (c) more fossil evidence bearing on the past climates.

Africa, Southern↗

Radiocarbon as a tool to apportion the sources of polycyclic aromatic hydrocarbons and black carbon in environmental samples.

To determine the relative inputs of polycyclic aromatic hydrocarbons (PAHs) and black carbon (BC) in environmental samples from the combustion of fossil fuels and biomass, we have developed two independent analytical methods for determining the 14C abundance of PAHs and BC. The 5730 yr half-life of 14C makes it an ideal tracer for identifying combustion products derived from fossil fuels (14C-free) versus those stemming from modern biomass (contemporary 14C). The 14C abundance of PAHs in several environmental Standard Reference Materials was measured by accelerator mass spectrometry after extraction and then purification by high-performance liquid chromatography and preparative capillary gas chromatography. This method yields pure compounds that allow for a high degree of confidence in the 14C results. The PAHs data were then used to compare and evaluate results from an operationally defined thermal oxidation method used to isolate a BC fraction. The 14C compositions of PAHs and BC were very similar and suggest that the thermal oxidation method employed for isolating BC is robust and free from interferences by non-BC components. In addition, these data indicate that both the PAHs and the BC species derive mostly from fossil fuels and/or their combustion products.

Biomass↗

Compound class specific 14C analysis of polycyclic aromatic hydrocarbons associated with PM10 and PM1.1 aerosols from residential areas of suburban Tokyo.

Compound class specific radiocarbon analysis (CCSRA) was performed for polycyclic aromatic hydrocarbons (PAHs) associated with airborne particulate matter (APM) with diameter <10 microm (PM10) and <1.1 microm (PM1.1) collected from a residential area of suburban Tokyo, Japan, and seasonal and particle-size radiocarbon variations were investigated. Source diagnostic isomer pair ratios indicated mixed contributions from petroleum combustion and from biomass and coal combustion to the PAHs in APM. The delta14C- PAHs in APM, ranging from -787 to -514 per thousand, indicated dominance of fossil fuel combustion. The delta14C of 5-6 rings (HMW) PAHs were higher than the 3-4 rings (LMW) species in both PM10 and PM1.1 samples. The delta14C of HMW-PAHs indicated greater biomass-burning contributions in summer than in winter and no apparent particle-size variation. Conversely, the delta14C of LMW species showed a greater contribution from fossil sources in summer and in larger particles (PM10). This finding could be tentatively attributed to the recondensation of fossil-PAHs vaporized from petroleum sources. A 14C isotopic mass balance approach estimated that biomass burning contributes 17-45% of the PAH burden in suburban Tokyo, and that the increase in the biomass-PAH accounts for approximately 27% and 22% of winter-time elevation of LMW- and HMW-PAHs, respectively. These are far exceeding what is expected from the emission statistics for CO2 and combusted materials in Japan and emphasizing the importance of biomass-burning as a source of PAHs; which, in turn, demonstrates the utility and the significance of field-based source assessment by using CCSRA for an effective regulation of atmospheric pollution by PAHs.

Aerosols↗

A molecular timescale for vertebrate evolution.

A timescale is necessary for estimating rates of molecular and morphological change in organisms and for interpreting patterns of macroevolution and biogeography. Traditionally, these times have been obtained from the fossil record, where the earliest representatives of two lineages establish a minimum time of divergence of these lineages. The clock-like accumulation of sequence differences in some genes provides an alternative method by which the mean divergence time can be estimated. Estimates from single genes may have large statistical errors, but multiple genes can be studied to obtain a more reliable estimate of divergence time. However, until recently, the number of genes available for estimation of divergence time has been limited. Here we present divergence-time estimates for mammalian orders and major lineages of vertebrates, from an analysis of 658 nuclear genes. The molecular times agree with most early (Palaeozoic) and late (Cenozoic) fossil-based times, but indicate major gaps in the Mesozoic fossil record. At least five lineages of placental mammals arose more than 100 million years ago, and most of the modern orders seem to have diversified before the Cretaceous/Tertiary extinction of the dinosaurs.

Animals↗

Organically preserved microbial endoliths from the late Proterozoic of East Greenland.

Diverse microorganisms ranging from cyanobacteria to eukaryotic algae and fungi live endolithically within ooids, hardgrounds and invertebrate shells on the present-day sea floor. These organisms are involved in the mechanical destruction of carbonates, and are useful ecological indicators of water depth and pollution. The Phanerozoic history of microbial endoliths has been elucidated through the study of microborings (the trace fossils of endolithic microorganisms) and rare cellularly preserved individuals, but nothing was known of the possible Precambrian evolution of comparable microorganisms until Campbell documented the occurrence of microborings in late Proterozoic ooids from central East Greenland. We now report the discovery of large populations of organically preserved endolithic microorganisms in silicified pisolites from 700-800-Myr-old Limestone-Dolomite Series of East Greenland. This fossil assemblage is significant for three reasons: (1) It confirms the prediction that oolites, pisolites and hardgrounds--the substrates for pre-Phanerozoic endoliths--provide a hitherto poorly explored but rewarding set of environments into which the search for early microfossils must be broadened; (2) the assemblage is diverse, containing about 12 taxa of morphologically distinct and previously unknown endolithic cyanobacteria, plus associated epilithic and interstitial populations; and (3) at least six of the fossil populations are indistinguishable in morphology, pattern of development, reproductive biology and inferred ecology from distinctive cyanobacterial species that bore ooids today in the Bahama Banks.

Carbonates↗

Filamentous microfossils in a 3,235-million-year-old volcanogenic massive sulphide deposit.

The record of Archaean microfossils is sparse. Of the few bona fide fossil assemblages, most are from shallow-water settings, and they are typically associated with laminated, stromatolitic sedimentary rocks. Microfossils from deep-sea hydrothermal systems have not been reported in Precambrian rocks (> 544 million years old), although thermophilic microbes are ubiquitous in modern sea-floor hydrothermal settings, and apparently have the most ancient lineages. Here, I report the discovery of pyritic filaments, the probable fossil remains of thread-like microorganisms, in a 3,235-million-year-old deep-sea volcanogenic massive sulphide deposit from the Pilbara Craton of Australia. From their mode of occurrence, the micro-organisms were probably thermophilic chemotropic prokaryotes, which inhabited sub-sea-floor hydrothermal environments. They represent the first fossil evidence for microbial life in a Precambrian submarine thermal spring system, and extend the known range of submarine hydrothermal biota by more than 2,700 million years. Such environments may have hosted the first living systems on Earth, consistent with proposals for a thermophilic origin of life.

Archaea↗

An Early Cambrian tunicate from China.

Like the Burgess Shales of Canada, the Chengjiang Lagerstätte from the Lower Cambrian of China is renowned for the detailed preservation as fossils of delicate, soft-bodied creatures, providing an insight into the Cambrian explosion. The fossils of possible hemichordate chordates and vertebrates have attracted particular attention. Tunicates, or urochordates, comprise the most basal chordate clade, and details of their evolution could be important in understanding the sequence of character acquisition that led to the emergence of chordates and vertebrates. However, definitive fossils of tunicates from the Cambrian are scarce or debatable. Here we report a probable tunicate Cheungkongella ancestralis from the Chengjiang fauna. It resembles the extant ascidian tunicate genus Styela whose morphology could be useful in understanding the origin of the vertebrates.

Animals↗

Neanderthal cranial ontogeny and its implications for late hominid diversity.

Homo neanderthalensis has a unique combination of craniofacial features that are distinct from fossil and extant 'anatomically modern' Homo sapiens (modern humans). Morphological evidence, direct isotopic dates and fossil mitochondrial DNA from three Neanderthals indicate that the Neanderthals were a separate evolutionary lineage for at least 500,000 yr. However, it is unknown when and how Neanderthal craniofacial autapomorphies (unique, derived characters) emerged during ontogeny. Here we use computerized fossil reconstruction and geometric morphometrics to show that characteristic differences in cranial and mandibular shape between Neanderthals and modern humans arose very early during development, possibly prenatally, and were maintained throughout postnatal ontogeny. Postnatal differences in cranial ontogeny between the two taxa are characterized primarily by heterochronic modifications of a common spatial pattern of development. Evidence for early ontogenetic divergence together with evolutionary stasis of taxon-specific patterns of ontogeny is consistent with separation of Neanderthals and modern humans at the species level.

Adolescent↗

The earliest known fully quadrupedal sirenian.

Modern seacows (manatees and dugongs; Mammalia, Sirenia) are completely aquatic, with flipperlike forelimbs and no hindlimbs. Here I describe Eocene fossils from Jamaica that represent nearly the entire skeleton of a new genus and species of sirenian--the most primitive for which extensive postcranial remains are known. This animal was fully capable of locomotion on land, with four well-developed legs, a multivertebral sacrum, and a strong sacroiliac articulation that could support the weight of the body out of water as in land mammals. Aquatic adaptations show, however, that it probably spent most of its time in the water. Its intermediate form thus illustrates the evolutionary transition between terrestrial and aquatic life. Similar to contemporary primitive cetaceans, it probably swam by spinal extension with simultaneous pelvic paddling, unlike later sirenians and cetaceans, which lost the hindlimbs and enlarged the tail to serve as the main propulsive organ. Together with fossils of later sirenians elsewhere in the world, these new specimens document one of the most marked examples of morphological evolution in the vertebrate fossil record.

Animals↗

Primitive deuterostomes from the Chengjiang Lagerstätte (Lower Cambrian, China).

Cambrian fossil-Lagerstätten (sites of exceptional fossil preservation), such as those from Chengjiang (Lower Cambrian) and the Burgess Shale (Middle Cambrian), provide our best window into the Cambrian 'explosion'. Such faunas are known from about 40 localities, and have yielded a widely disparate series of taxa ranging from ctenophores to agnathan fish. Recent excavations of the Chengjiang fossil-Lagerstätte, known from a series of sites near Kunming in Yunnan, south China, have resulted in the discovery of several new forms. In conjunction with material described earlier, these provide evidence for a new group of metazoans, the vetulicolians. Several features, notably a series of gill slits, suggest that this group can throw light on an early stage of deuterostome diversification.

Animals↗

Primate origins: plugging the gaps.

Recent discoveries of fossil primate specimens have produced several surprises and challenged prevailing views of early primate evolution. Plesiadapiformes, long regarded as 'archaic primates', may perhaps be linked to the peculiar colugos instead. Inferred relationships of the earliest known undoubted primates (adapids and omomyids) are in turmoil. Both groups have been proposed as sources for the simian primates. Although the origin of the simian primates is obscure, new fossil evidence could push it further back by at least 10 million years. Such uncertainties reflect the low sampling level of the primate fossil record, which can potentially also lead to underestimation of times of origin within the primate tree.

Animals↗