Fossil Age Research Articles

The legacy of the mastodon: the golden age of fossils in America

The legacy of the mastodon: the golden age of fossils in America

青森県弘前市南方の付加体泥岩から産出した前期ジュラ紀放散虫化石

Radiolarian assemblages including Canoptum spp. in part with Natoba minuta or ?Parahsuum spp. were extracted from terrigenous mudstone within an accretionary complex located south of Hirosaki City, near the western margin of the North Kitakami Belt. These assemblages suggest sedimentation during the Early Jurassic, older than any known fossil ages (Middle Jurassic to latest Jurassic-earliest Cretaceous) obtained for mudstone in the North Kitakami Belt. This result indicates the occurrence of near-continuous accretionary tectonics in NE Japan from the Early Jurassic to earliest Cretaceous, comparable with age data obtained for SW Japan. Although the accretionary complex located south of Hirosaki occurs as an isolated inlier among younger cover sediments, its age and location suggest that it corresponds to one of the structurally uppermost units in the North Kitakami Belt.

New geochronological, paleoclimatological, and archaeological data from the Narmada Valley hominin locality, central India

The oldest known fossil hominin in southern Asia was recovered from Hathnora in the Narmada Basin, central India in the early 1980's. Its age and taxonomic affinities, however, have remained uncertain. Current estimates place its maximum age at >236 ka, but not likely older than the early middle Pleistocene. The calvaria, however, could be considerably younger. We report recent fieldwork at Hathnora and associated Quaternary type-sections that has provided new geological and archaeological insights. The portion of the exposed ‘Boulder Conglomerate’ within the Surajkund Formation, which forms a relict terrace and has yielded the hominin fossils, contains reworked and stylistically mixed lithic artifacts and temporally mixed fauna. Three mammalian teeth stratigraphically associated with the hominin calvaria were dated by standard electron spin resonance (ESR). Assuming an early uranium uptake (EU) model for the teeth, two samples collected from the reworked surface deposit averaged 49 ± 1 ka (83 ± 2 ka, assuming linear uptake [LU]; 196 ± 7 ka assuming recent uptake [RU]). Another sample recovered from freshly exposed, crossbedded gravels averaged 93 ± 5 ka (EU), 162 ± 8 ka (LU) or 407 ± 21 ka (RU). While linear uptake models usually provide the most accurate ages for this environment and time range, the EU ages represent the minimum possible age for fossils in the deposit. Regardless, the fossils are clearly reworked and temporally mixed. Therefore, the current data constrains the minimum possible age for the calvaria to 49 ± 1 ka, although it could have been reworked and deposited into the Hathnora deposit any time after 160 ka (given the LU uptake ages) or earlier (given the RU ages). At Hathnora, carbonaceous clay, bivalve shells, and a bovid tooth recovered from layers belonging to the overlying Baneta Formation have yielded 14C ages of 35.66 ± 2.54 cal ky BP, 24.28 ± 0.39 cal ky BP, and 13.15 ± 0.34 ky BP, respectively. Additional surveys yielded numerous lithics and fossils on the surface and within the stratigraphic sequence. At the foot of the Vindhyan Hills 2 km from the river, we recovered a typologically Early Acheulean assemblage comprised of asymmetrical bifaces, large cleavers with minimal working, trihedral picks, and flake tools in fresh condition. These tools may be the oldest Acheulean in the Narmada Valley. Several lithics recovered from the Dhansi Formation may represent the first unequivocal evidence for an early Pleistocene hominin presence in India. In situ invertebrate and vertebrate fossils, pollen, and spores indicate a warm, humid climate during the late middle Pleistocene. High uranium concentrations in the mammalian teeth indicate exposure to saline water, suggesting highly evaporative conditions in the past. Late Pleistocene sediment dated between 24.28 ± 0.39 cal ky BP and 13.15 ± 340 ky BP has yielded pollen and spores indicating cool, dry climatic conditions corresponding to Oxygen Isotope Stage 2 (OIS 2). An early Holocene palynological assemblage from the type locality at Baneta shows evidence for relatively dry conditions and a deciduous forest within the region. The Dhansi Formation provisionally replaces the Pilikarar Formation as the oldest Quaternary formation within the central Narmada Basin. The Baneta Formation, previously dated at 70 ka to 128 ka, correlates with the late Pleistocene and early Holocene. Our results highlight the need for further Quaternary geological and paleoanthropological research within the Narmada Basin, especially because dam construction threatens these deposits.

Using fossils and molecular data to reveal the origins of the Cape proteas (subfamily Proteoideae)

The angiosperm family Proteaceae is a distinct component of the Cape Floristic Region biodiversity hotspot with 330 endemic species. Phylogenetic analyses of subfamily Proteoideae using sequence data from one nuclear and six plastid loci show that most of this diversity is contained in two distinct Cape floral clades. Molecular dating analyses, using Bayesian and penalized likelihood methods and four phylogenetically supported fossil age constraints, reveal contrasting histories for these two clades. The genus Protea belongs to a lineage that may have been in Africa since the Late Cretaceous but began to diversify in the Cape only 5–18 Myr ago. In contrast, the Leucadendrinae clade presumably arrived in the region no earlier than 46 Myr ago by long-distance dispersal from an Australian ancestor and the extant members of this clade began to diversify in the Cape 22–39 Myr ago. These results join a growing number of case studies that challenge the commonly accepted view that most of the Cape flora radiated synchronously in the Late Miocene and Early Pliocene when a Mediterranean climate settled in the region.

A new Cretaceous age for the Saiqu “mélange,” southern Tibet: evidence from Radiolaria

This study concerns the mélange that crops out in the north of Saiqu, Sagya County, southern Tibet. The age of the mélange has been debated for years, because the ages of fossils from various blocks conflicted with each other, and no macrofossils had been discovered from the matrix. Furthermore, no one had looked at the matrix for microfossils, mainly because of their mistaking this suite of mélange as straightforward sedimentary strata. However, abundant microfossils are present in the matrix of the mélange, and four radiolarian assemblages have been recognized: (1) Holocryptocanium aff. barbui- Dictyomitra turris; (2) Acanthocircus dicrancanthos- Thanarla pulchra; (3) Thanarla veneta-Pseudodictyomitra pseudomacrocephala; and (4) Dictyomitra magnifica- Dictyomitra turritum. The matrix-derived radiolarian assemblages indicate a Late Cretaceous age. The mélange in Saiqu can be correlated with the Zongzhuo Formation of neighbouring regions by either lithological associations or fossil assemblages. Consequently, the mélange has been referred to the Upper Cretaceous Zongzhuo Formation, instead of the Triassic Xiukang Group, with which it was previously associated. The mélange of the Zongzhuo Formation resulted from the subduction of the Indian Plate beneath the Asian Plate during the Late Cretaceous.

Ichnotaxonomy of bird-like footprints: an example from the Late Triassic-Early Jurassic of northwest Argentina

ABSTRACT The ichnotaxobases previously used to classify avian-like footprints, at the ichnogeneric, ichnospecific, and ichnofamily level, are varied and contrasting. Consequently, an agreement on the most adequate taxobases to use for classifying these vertebrate trace fossils is necessary. The authors follow an ichnotaxonomy treatment independent to the age, locality provenance, and possible tracemaker of the trace fossils. The ichnotaxobases used to classify tracks with avian affinities at ichnogeneric and ichnospecific levels are evaluated and a proposal is made for useful and appropriate ichnotaxonomic criteria, considering those currently in use. Previous criteria used to distinguish avian footprints from non-avian theropod or ornithischian tracks are discussed. These concepts are applied to the avian footprints from the upper part of the Late Triassic-Early Jurassic Santo Domingo Formation from La Rioja province, northwest Argentina, which has yielded a diverse assemblage of trace fossils. The most conspicuous avian footprint is Gruipeda dominguensis isp. nov. The ichnogenus Gruipeda Panin and Avram, 1962 is revised and an emendation of its diagnosis is suggested. Trisauropodiscus Ellenberger, 1972, from South Africa and Antarctichnus Covacevich and Lamperein, 1970 from Antarctica are considered as junior synonyms of Gruipeda. Three other morphotypes of avian footprints are left under open nomenclature: one is assigned as cf. Alaripeda isp., other as bird-like footprints type C, and the third bird-like footprint with elongated drag marks. These specimens could be related to avian origin, but the possibility of a case of convergence with birds is not discarded. The presence of tracks with a wide total divarication produced in ephemeral fluvial systems with shallow ponds and mudflats suggest that it is likely the attainment of a convergent avian-form feet to improve controlled movements.

Landscape ecology and sustainability

One of the major themes of the recent 7th IALE World Congress was ‘‘Landscape Ecology and Sustainability.’’ However, in spite of the many planning, management, conservation, and restoration projects presented at the congress and published in ‘‘Landscape Ecology’’ and other journals, the real impact of landscape ecology (LE) on decision making on sustainable land use is still very limited. With exception of the Netherlands, Slovakia, and UK, the ‘‘landscape’’ as the most suitable, integrative conceptual, and practical tool for sustainable development has not yet reached nationand worldwide recognition. This IALE World Congress took place in the crucial transitional period from the industrial age to the global information age. This ‘‘Macroshift’’ (sensu Laszlo 2001) is marked by a severe ecological, cultural, and socio-economic crisis, in which human society has little time left for the choice of navigating this transformation either to a breakdown or to breakthrough towards a sustainable world. Such a breakthrough is a ‘‘chaos point’’ (Laszlo 2006), during which any input or influence on the system, however small, can replace existing trends by new trends and processes. It can be achieved only by an urgently needed, ecological, socio-economic and cultural and technological ‘‘sustainability revolution’’, leading to the sustainable future of nature and human life on Earth. In this sustainability revolution full use of scientific and technological potentials should be made for a post-industrial symbiosis between human society and nature. This requires above all a shift from the ‘‘fossil age’’ to the ‘‘solar age’’ of a new world economy, based on the limitless power of the sun as the non-polluting and renewable energy source. It requires a shift from depletion of natural resources to their more efficient and wiser sustainable use, by recycling and reducing through-flows of material and energy and their adverse impacts on human and landscape health. It will be driven not only by the widespread adoption of technological innovations of regenerative and recycling methods, but also the efficient utilization of solar, wind, water and other non-polluting and renewable sources of energy. As a cultural evolutionary process it must be coupled with more sustainable lifestyles and consumption patterns, caring for nature and even investing in nature. We will only be able to gain a significant influence in the decision process towards sustainability if we will take an active role in steering this Macroshift towards such an all-embracing sustainability revolution as concerned landscape scientists. Our main challenge is to respond together with all those dealing Z. Naveh (&) Faculty of Civil and Environmental Engineering, Technion Israel Institute of Technology, Haifa 32 000, Israel e-mail: znave@technion.ac.il

Deformation process and kinematics of mélange in the Early Cretaceous accretionary complex of the Mino‐Tamba Belt, eastern southwest Japan

The Kanayama unit of the Mesozoic Mino‐Tamba accretionary complex in eastern southwest Japan is a thick mélange unit composed mainly of abundant muddy matrix and variously sized clasts of chert, hemipelagic siliceous mudstone, and sandstone. Two episodes of deformation formed the mélange fabrics: the first involved fragmentation of sandstone layers in response to mud injections; the second involved layer‐parallel, noncoaxial shear that resulted in the mixing of pelagic to terrigenous clasts and the formation of S‐C‐like asymmetric fabrics with scaly foliation. The S‐C fabrics exhibit a predominantly sinistral shear sense in plan view. After fold and tilt corrections, the restored slip motion inferred from the S‐C fabrics indicates a bulk top‐to‐the‐south shear. Combining the unit's terminal fossil age and radiometric age with a regional plate reconstruction model, these data suggest that the Kanayama mélange formed along a décollement during oblique subduction of the Izanagi plate in the Early Cretaceous.

Geological and geochemical characteristics of the Sawaya'erdun gold deposit, southwestern Chinese Tianshan

The Sawaya'erdun gold deposit in the western segment of the South Tianshan, Xinjiang, China, is a large, low-grade orogenic gold deposit. An abundance of index fossils in the ore-hosting strata, including brachiopoda, corals and Schwagerina, show that the fossil age of the country rocks ranges from Late Carboniferous to Early Permian. Whole rock Rb–Sr and Sm–Nd isochron ages, ranging from 292.43 ± 0.38 to 304.7 ± 11.6 Ma, are concordant with a Late Carboniferous age. Gold mineralization is strictly controlled by a fault zone. Ore minerals are dominated by sulfides, with gold inhomogeneously distributed within them; gangue minerals are quartz, siderite or calcite. Data suggest that the ore-forming fluid was derived predominantly from an active meteoric groundwater system. Ore-forming temperatures are estimated to have been within the range 110 to 220 °C. New 40Ar/ 39Ar isotope age determinations of auriferous quartz are in excellent agreement with Rb–Sr ages obtained from fluid inclusions in quartz. They suggest gold mineralization at 206 to 213 Ma (Late Indosinian). Such an age is consistent with the age of formation given for other gold deposits in the region, such as Bulong and Dashankou. Combined with K–Ar geochronological data and observed geological features, it is concluded that the Indosinian metallogentic epoch represents a previously unrecognized, major period of gold metallogeny in the southwestern Chinese Tianshan. In some ways, the Sawaya'erdun gold deposit can be compared with the Muruntau gold deposit in Uzbekistan with respect to geological setting, host lithology, mineralization style, mineral assemblages, geochemical association and metallogenic processes involved. Despite this, the Sawaya'erdun gold deposit possesses a number of characteristics typical of only low-temperature metallogenesis.

Origin and Cretaceous tectonic history of the coastal Ecuadorian forearc between 1°N and 3°S: Paleomagnetic, radiometric and fossil evidence

The presence of at least one Block (Pallatanga Block) derived from the Caribbean–Colombian Oceanic Plateau has been proposed in the Western Cordillera of Ecuador. New 40Ar/ 39Ar dating of the basement of the coastal blocks (90–87 Ma) and fossil ages in the overlying Calentura Fm. (89–84 Ma) argue for late Turonian–Coniacian extrusion of the basement of the Piñon block. Paleomagnetic inclinations indicate that the mafic basement of the Piñon and San Lorenzo blocks extruded at equatorial, low southern latitudes. No subsequent significant latitudinal drift can be observed during the Cretaceous. Rapid changes in paleomagnetic declination between 73 and 70 Ma in both the Piñon and the San Lorenzo blocks indicate that the large igneous province, which originally hosted the coastal blocks experienced a major tectonic event during Late Campanian times. Similar Late Turonian–Coniacian basement ages and Late Campanian tectonic events are reported in the Pallatanga Block of the Western Cordillera. We propose that the Piñon, San Lorenzo and Pallatanga blocks all derive from a common large igneous province, the Caribbean–Colombian Oceanic Plateau, and were incorporated into the South American margin by collision during the Late Campanian (73–70 Ma).

Beginsels en metodes vir die bepaling van die ouderdom van hominied fossiele en hulle vindplekke

Principles and methods used for the determination of the age of hominid fossils and their discovery sites Paleoanthropologists have an array of physical dating methods developed by researchers in the physical sciences. These techniques permit students of hominid evolution and associated culture to have increasing confidence that the evidence of past hominid morphology as expressed by the fossil finds is being placed in the correct chronological order. This article explains in a simple and straightforward way the evidence and logic that have led scientists to conclude that hominid fossils are not a few thousand years but at least several million years old. The main focus is on the 40Ar/ 39Ar technique, but other absolute dating techniques such as carbon-14, uranium-thorium-lead and radiocalcium are also reviewed. Relative dating methods are briefly discussed together with paleomagnetism (geomagnetism), optic stimulated luminescence and amino acids.

Uncertainty in the Age of Fossils and the Stratigraphic Fit to Phylogenies

The ages of first appearance of fossil taxa in the stratigraphic record are inherently associated to an interval of error or uncertainty, rather than being precise point estimates. Contrasting this temporal information with topologies of phylogenetic relationships is relevant to many aspects of evolutionary studies. Several indices have been proposed to compare the ages of first appearance of fossil taxa and phylogenies. For computing most of these indices, the ages of first appearance of fossil taxa are currently used as point estimates, ignoring their associated errors or uncertainties. The effect of age uncertainty on measures of stratigraphic fit to phylogenies is explored here for two indices based on the extension of ghost lineages (MSM* and GER). A solution based on randomization of the ages of terminal taxa is implemented, resulting in a range of possible values for measures of stratigraphic fit to phylogenies, rather than in a precise but arbitrary stratigraphic fit value. Sample cases show that ignoring the age uncertainty of fossil taxa can produce misleading results when comparing the stratigraphic fit of competing phylogenetic hypotheses. Empirical test cases of alternative phylogenies of two dinosaur groups are analyzed through the randomization procedure proposed here.

The making of geology

In the late eighteenth century, ideas about the age of rocks and fossils gave rise to a new science.

Dating Paleosol and Animal Remains in Loess Deposits

Accurate and reliable dating of paleosols, animal remains, and artifacts is of crucial importance in reconstructing environmental change and understanding the interrelationship between human activities and natural environments. Dating different materials in the same sample can help resolve problems such as soil carbon sources and carbon storage state. Conventional radiocarbon dating of soil (inorganic and organic matter) and accelerator mass spectrometry (AMS) dating of animal remains (fossil bones and teeth) result in different ages for materials from the same sample position in a typical loess section at Xinglong Mountain, Yuzhong County, Gansu Province in NW China. Inorganic matter is ∼3400 yr older than organic matter, 4175 ± 175 cal BP to 3808 ± 90 cal BP. A 1610-yr difference between the 14C ages of fossils (animal bones and teeth) and soil organic matter suggests that a depositional hiatus exists in the studied profile. The varying 14C ages of fossils and soil organic and inorganic matter have important implications for paleoclimate reconstructions from loess sections. It is critical to consider the meaning of the variable 14C ages from different material components from the same sample position in terms of soil organic and inorganic carbon storage, vegetation history reconstruction, archaeology, and the study of ancient civilizations.

Historical biogeography of Australian Rhamnaceae, tribe Pomaderreae

AbstractAim To discover the pattern of relationships of areas of endemism for Australian genera in the plant family Rhamnaceae tribe Pomaderreae for comparison with other taxa and interpretation of biogeographical history.Location Australian mainland, Tasmania and New Zealand.Methods A molecular phylogeny and geographic distribution of species within four clades of Pomaderreae are used as a basis for recognition of areas of endemism and analysis of area relationships using paralogy‐free subtrees. The taxon phylogeny is the strict consensus tree from a parsimony analysis of 54 taxa, in four clades, and sequence data for the internal transcribed spacer regions of ribosomal DNA (ITS1‐5.8S‐ITS2) and the plastid DNA region trnL‐F.Results The biogeographical analysis identified five subtrees, which, after parsimony analysis, resulted in a minimal tree with 100% consistency and seven resolved nodes. Three sets of area relationships were identified: the areas of Arnhem and Kimberley in tropical north Australia are related based on the phylogeny of taxa within Cryptandra; the moister South‐west of Western Australia, its sister area the coastal Geraldton Sandplains, the semi‐arid Interzone region and arid Western Desert are related, based on taxa within Cryptandra, Spyridium, Trymalium and Pomaderris; and the eastern regions of Queensland, McPherson‐Macleay, south‐eastern New South Wales (NSW), Victoria, southern Australia, Tasmania and New Zealand are related based on Cryptandra, Pomaderris and Spyridium. Tasmania and NSW are related based entirely on Cryptandra, but the position of New Zealand relative to the other south‐eastern Australian regions is unresolved.Main conclusions The method of paralogy‐free subtrees identified a general pattern of geographic area relationships based on Australian Pomaderreae. The widespread distribution of clades, the high level of endemicity and the age of fossils for the family, suggest that the Pomaderreae are an old group among the Australian flora. Their biogeographical history may date to the early Palaeogene with subsequent changes through to the Pleistocene.

FOSSIL CALIBRATIONS AND MOLECULAR DIVERGENCE TIME ESTIMATES IN CENTRARCHID FISHES (TELEOSTEI: CENTRARCHIDAE)

Molecular clock methods allow biologists to estimate divergence times, which in turn play an important role in comparative studies of many evolutionary processes. It is well known that molecular age estimates can be biased by heterogeneity in rates of molecular evolution, but less attention has been paid to the issue of potentially erroneous fossil calibrations. In this study we estimate the timing of diversification in Centrarchidae, an endemic major lineage of the diverse North American freshwater fish fauna, through a new approach to fossil calibration and molecular evolutionary model selection. Given a completely resolved multi-gene molecular phylogeny and a set of multiple fossil-inferred age estimates, we tested for potentially erroneous fossil calibrations using a recently developed fossil cross-validation. We also used fossil information to guide the selection of the optimal molecular evolutionary model with a new fossil jackknife method in a fossil-based model cross-validation. The centrarchid phylogeny resulted from a mixed-model Bayesian strategy that included 14 separate data partitions sampled from three mtDNA and four nuclear genes. Ten of the 31 interspecific nodes in the centrarchid phylogeny were assigned a minimal age estimate from the centrarchid fossil record. Our analyses identified four fossil dates that were inconsistent with the other fossils, and we removed them from the molecular dating analysis. Using fossil-based model cross-validation to determine the optimal smoothing value in penalized likelihood analysis, and six mutually consistent fossil calibrations, the age of the most recent common ancestor of Centrarchidae was 33.59 million years ago (mya). Penalized likelihood analyses of individual data partitions all converged on a very similar age estimate for this node, indicating that rate heterogeneity among data partitions is not confounding our analyses. These results place the origin of the centrarchid radiation at a time of major faunal turnover as the fossil record indicates that the most diverse lineages of the North American freshwater fish fauna originated at the Eocene-Oligocene boundary, approximately 34 mya. This time coincided with major global climate change from warm to cool temperatures and a signature of elevated lineage extinction and origination in the fossil record across the tree of life. Our analyses demonstrate the utility of fossil cross-validation to critically assess individual fossil calibration points, providing the ability to discriminate between consistent and inconsistent fossil age estimates that are used for calibrating molecular phylogenies.

ANGIOSPERM DIVERGENCE TIMES: THE EFFECT OF GENES, CODON POSITIONS, AND TIME CONSTRAINTS

Abstract An understanding of the evolution of modern terrestrial ecosystems requires an understanding of the dynamics associated with angiosperm evolution, including the timing of their origin and diversification into their extraordinary present-day diversity. Molecular estimates of angiosperm age have varied widely, and many substantially predate the Early Cretaceous fossil appearance of the group. In this study, the effect of different genes, codon positions, and chronological constraints on node ages are examined on divergence time estimates across seed plants, with a special focus on angiosperms. Penalized likelihood was used to estimate divergence times on a phylogenetic hypothesis for seed plants derived from Bayesian analysis, with branch lengths estimated with maximum likelihood. The plastid genes atpB, psaA, psbB, and rbcL were used individually and in combination, using first and second, third, and the three codon positions, including and excluding age constraints on 20 nodes derived from a critical examination of the land-plant fossil record. The optimal level of rate smoothing according to each unconstrained and constrained dataset was obtained with penalized likelihood. Tests for a molecular clock revealed significantly unclocklike rates in all datasets. Addition of fossil constraints resulted in even greater departures from constancy. Consistently with significant deviations from a clock, estimated optimal smoothing values were low, but a strict correlation between rate heterogeneity and optimal smoothing value was not found. Age estimates for nodes across the phylogeny varied, sometimes substantially, with gene and codon position. Nevertheless, estimates based on the four concatenated genes are very similar to the mean of the four individual gene estimates. For any given node, unconstrained age estimates are more variable than constrained estimates and are frequently younger than well-substantiated fossil members of the clade. Constrained estimates of ages of clades are older than unconstrained estimates and oldest fossil representatives, sometimes substantially so. Angiosperm age estimates decreased as rate smoothing increased. Whereas the range of unconstrained angiosperm age estimates spans the fossil age of the clade, the range of constrained estimates is narrower (and older) than the earliest angiosperm fossils. Results unambiguously indicate the relevance of constraints in reducing the variability of ages derived from different partitions of the data and diminishing the effect of the smoothing parameter. Constrained optimizations of divergence times and substitution rates across the phylogeny suggest appreciably different evolutionary dynamics for angiosperms and for gymnosperms. Whereas the gymnosperm crown group originated shortly after the origin of seed plants, a long time elapsed before the origin of crown group angiosperms. Although absolute age estimates of angiosperms and angiosperm clades are older than their earliest fossils, the estimated pace of phylogenetic diversification largely agrees with the rapid appearance of angiosperm lineages in stratigraphic sequences.

FOSSIL CALIBRATIONS AND MOLECULAR DIVERGENCE TIME ESTIMATES IN CENTRARCHID FISHES (TELEOSTEI: CENTRARCHIDAE)

Molecular clock methods allow biologists to estimate divergence times, which in turn play an important role in comparative studies of many evolutionary processes. It is well known that molecular age estimates can be biased by heterogeneity in rates of molecular evolution, but less attention has been paid to the issue of potentially erroneous fossil calibrations. In this study we estimate the timing of diversification in Centrarchidae, an endemic major lineage of the diverse North American freshwater fish fauna, through a new approach to fossil calibration and molecular evolutionary model selection. Given a completely resolved multi-gene molecular phylogeny and a set of multiple fossil-inferred age estimates, we tested for potentially erroneous fossil calibrations using a recently developed fossil cross-validation. We also used fossil information to guide the selection of the optimal molecular evolutionary model with a new fossil jackknife method in a fossil-based model cross-validation. The centrarchid phylogeny resulted from a mixed-model Bayesian strategy that included 14 separate data partitions sampled from three mtDNA and four nuclear genes. Ten of the 31 interspecific nodes in the centrarchid phylogeny were assigned a minimal age estimate from the centrarchid fossil record. Our analyses identified four fossil dates that were inconsistent with the other fossils, and we removed them from the molecular dating analysis. Using fossil-based model cross-validation to determine the optimal smoothing value in penalized likelihood analysis, and six mutually consistent fossil calibrations, the age of the most recent common ancestor of Centrarchidae was 33.59 million years ago (mya). Penalized likelihood analyses of individual data partitions all converged on a very similar age estimate for this node, indicating that rate heterogeneity among data partitions is not confounding our analyses. These results place the origin of the centrarchid radiation at a time of major faunal turnover as the fossil record indicates that the most diverse lineages of the North American freshwater fish fauna originated at the Eocene-Oligocene boundary, approximately 34 mya. This time coincided with major global climate change from warm to cool temperatures and a signature of elevated lineage extinction and origination in the fossil record across the tree of life. Our analyses demonstrate the utility of fossil cross-validation to critically assess individual fossil calibration points, providing the ability to discriminate between consistent and inconsistent fossil age estimates that are used for calibrating molecular phylogenies.

ANGIOSPERM DIVERGENCE TIMES: THE EFFECT OF GENES, CODON POSITIONS, AND TIME CONSTRAINTS

An understanding of the evolution of modern terrestrial ecosystems requires an understanding of the dynamics associated with angiosperm evolution, including the timing of their origin and diversification into their extraordinary present-day diversity. Molecular estimates of angiosperm age have varied widely, and many substantially predate the Early Cretaceous fossil appearance of the group. In this study, the effect of different genes, codon positions, and chronological constraints on node ages are examined on divergence time estimates across seed plants, with a special focus on angiosperms. Penalized likelihood was used to estimate divergence times on a phylogenetic hypothesis for seed plants derived from Bayesian analysis, with branch lengths estimated with maximum likelihood. The plastid genes atpB, psaA, psbB, and rbcL were used individually and in combination, using first and second, third, and the three codon positions, including and excluding age constraints on 20 nodes derived from a critical examination of the land-plant fossil record. The optimal level of rate smoothing according to each unconstrained and constrained dataset was obtained with penalized likelihood. Tests for a molecular clock revealed significantly unclocklike rates in all datasets. Addition of fossil constraints resulted in even greater departures from constancy. Consistently with significant deviations from a clock, estimated optimal smoothing values were low, but a strict correlation between rate heterogeneity and optimal smoothing value was not found. Age estimates for nodes across the phylogeny varied, sometimes substantially, with gene and codon position. Nevertheless, estimates based on the four concatenated genes are very similar to the mean of the four individual gene estimates. For any given node, unconstrained age estimates are more variable than constrained estimates and are frequently younger than well-substantiated fossil members of the clade. Constrained estimates of ages of clades are older than unconstrained estimates and oldest fossil representatives, sometimes substantially so. Angiosperm age estimates decreased as rate smoothing increased. Whereas the range of unconstrained angiosperm age estimates spans the fossil age of the clade, the range of constrained estimates is narrower (and older) than the earliest angiosperm fossils. Results unambiguously indicate the relevance of constraints in reducing the variability of ages derived from different partitions of the data and diminishing the effect of the smoothing parameter. Constrained optimizations of divergence times and substitution rates across the phylogeny suggest appreciably different evolutionary dynamics for angiosperms and for gymnosperms. Whereas the gymnosperm crown group originated shortly after the origin of seed plants, a long time elapsed before the origin of crown group angiosperms. Although absolute age estimates of angiosperms and angiosperm clades are older than their earliest fossils, the estimated pace of phylogenetic diversification largely agrees with the rapid appearance of angiosperm lineages in stratigraphic sequences.

Early Carboniferous radiolarians discovered from the Hayachine Terrane, Northeast Japan: the oldest fossil age for clastic rocks of accretionary complex in Japan

Early Carboniferous radiolarians were newly discovered from siltstone of the Nedamo Complex in the Hayachine Terrane, Northeast Japan. This siltstone and other clastic rocks along with greenstone and chert are the components of an accretionary complex. Radiolarian fauna contains Palaeoscenidium cladophorum Deflandre that ranges in age from Early or Middle Devonian to Early Carboniferous. Since the Fe-Mn chert intercalated in a MORB-type basalt of the Nedamo Complex was assigned an age of Late Devonian (Hamano et al., 2002), the accretionary age of the Hayachine Terrane is no older than Late Devonian, most probable Early Carboniferous. This is the first report of an Early Carboni-ferous accretionary complex recognized by biostratigraphic data in Japan.