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Discoveries of archaic Mesozoic fossil birds ('opposite' birds, or enantiornithines) during the past decade have revolutionized our understanding of early avian evolution, but the rarity of Early Cretaceous ornithurines - birds that are closely related to the modern avian radiation - has meant that information about these species has lagged behind.
Plants respond to changes in atmospheric carbon dioxide levels by regulating the number of stomata in their leaves. In his reconstruction of a continuous, 300-million-year record of atmospheric CO2, Retallack bases his curve on stomatal counts of fossil plant cuticles taken from published micrographs. However, the preservation of cuticles from Permian times is generally too fragmentary for the stomatal index to be reliably determined, the micrographs used could have biased the results, and there are important errors in the supplementary data - all of which cast doubt on the Permian part of Retallack's record.
The dynamics of extinction and diversification determine the long-term effects of extinction episodes. If rapid bursts of extinction are offset by equally rapid bursts of diversification, their biodiversity consequences will be transient. But if diversification rates cannot accelerate rapidly enough, pulses of extinction will lead to long-lasting depletion of biodiversity. Here I use spectral analysis of the fossil record to test whether diversification rates can accelerate as much as extinction rates, over both short and long spans of geological time. I show that although the long-wavelength variability of diversification rates equals or exceeds that of extinctions, diversification rates are markedly less variable than extinction rates at wavelengths shorter than roughly 25 million years. This implies that there are intrinsic speed limits that constrain how rapidly diversification rates can accelerate in response to pulses of extinction.
The causes of mass extinctions and the nature of biological selectivity at extinction events are central questions in palaeobiology. It has long been recognized, however, that the amount of sedimentary rock available for sampling may bias perceptions of biodiversity and estimates of taxonomic rates of evolution. This problem has been particularly noted with respect to the principal mass extinctions. Here we use a new compilation of the amount of exposed marine sedimentary rock to predict how the observed fossil record of extinction would appear if the time series of true extinction rates were in fact smooth. Many features of the highly variable record of apparent extinction rates within marine animals can be predicted on the basis of temporal variation in the amount of exposed rock. Although this result is consistent with the possibility that a common geological cause determines both true extinction rates and the amount of exposed rock, it also supports the hypothesis that much of the observed short-term volatility in extinction rates is an artefact of variability in the stratigraphic record.
The chordates, hemichordates (such as acorn worms) and echinoderms (such as starfish) comprise the group Deuterostomia, well established as monophyletic. Among extant deuterostomes, a skeleton in which each plate has the crystallographic structure of a single crystal of calcite is characteristic of echinoderms and is always associated with radial symmetry and never with gill slits. Among fossils, however, such a skeleton sometimes occurs without radial symmetry. This is true of Jaekelocarpus oklahomensis, from the Upper Carboniferous of Oklahoma, USA, which, being externally almost bilaterally symmetrical, is traditionally placed in the group Mitrata (Ordovician to Carboniferous periods, 530-280 million years ago), by contrast with the bizarrely asymmetrical Cornuta (Cambrian to Ordovician periods, 540 to 440 million years ago). Using computer X-ray microtomography, we describe the anatomy of Jaekelocarpus in greater detail than formerly possible, reveal evidence of paired gill slits internally and interpret its functional anatomy. On this basis we suggest its phylogenetic position within the deuterostomes.
Morphological, molecular, and biogeographic data bearing on early primate evolution suggest that the clade containing extant (or 'crown') strepsirrhine primates (lemurs, lorises and galagos) arose in Afro-Arabia during the early Palaeogene, but over a century of palaeontological exploration on that landmass has failed to uncover any conclusive support for that hypothesis. Here we describe the first demonstrable crown strepsirrhines from the Afro-Arabian Palaeogene--a galagid and a possible lorisid from the late middle Eocene of Egypt, the latter of which provides the earliest fossil evidence for the distinctive strepsirrhine toothcomb. These discoveries approximately double the previous temporal range of undoubted lorisiforms and lend the first strong palaeontological support to the hypothesis of an ancient Afro-Arabian origin for crown Strepsirrhini and an Eocene divergence of extant lorisiform families.
The long-term carbon cycle operates over millions of years and involves the exchange of carbon between rocks and the Earth's surface. There are many complex feedback pathways between carbon burial, nutrient cycling, atmospheric carbon dioxide and oxygen, and climate. New calculations of carbon fluxes during the Phanerozoic eon (the past 550 million years) illustrate how the long-term carbon cycle has affected the burial of organic matter and fossil-fuel formation, as well as the evolution of atmospheric composition.
Comparative embryology is integral to uncovering the pattern and process of metazoan phylogeny, but it relies on the assumption that life histories of living taxa are representative of their antecedents. Fossil embryos provide a crucial test of this assumption and, potentially, insight into the evolution of development, but because discoveries so far lack phylogenetic constraint, their significance is moot. Here we describe a collection of embryos from the Middle and Late Cambrian period (500 million years ago) of Hunan, south China, that preserves stages of development from cleavage to the pre-hatching embryo of a direct-developing animal comparable to living Scalidophora (phyla Priapulida, Kinorhyncha, Loricifera). The latest-stage embryos show affinity to the Lower Cambrian embryo Markuelia, whose life-history strategy contrasts both with the primitive condition inferred for metazoan phyla and with many proposed hypotheses of affinity, all of which prescribe indirect development. Phylogenetic tests based on these embryological data suggest a stem Scalidophora affinity. These discoveries corroborate, rather than contradict, the predictions of comparative embryology, providing direct historical support for the view that the life-history strategies of living taxa are representative of their stem lineages.
Marine sediments act as the ultimate sink for organic carbon, sequestering otherwise rapidly cycling carbon for geologic timescales. Sedimentary organic carbon burial appears to be controlled by oxygen exposure time in situ, and much research has focused on understanding the mechanisms of preservation of organic carbon. In this context, combustion-derived black carbon has received attention as a form of refractory organic carbon that may be preferentially preserved in soils and sediments. However, little is understood about the environmental roles, transport and distribution of black carbon. Here we apply isotopic analyses to graphitic black carbon samples isolated from pre-industrial marine and terrestrial sediments. We find that this material is terrestrially derived and almost entirely depleted of radiocarbon, suggesting that it is graphite weathered from rocks, rather than a combustion product. The widespread presence of fossil graphitic black carbon in sediments has therefore probably led to significant overestimates of burial of combustion-derived black carbon in marine sediments. It could be responsible for biasing radiocarbon dating of sedimentary organic carbon, and also reveals a closed loop in the carbon cycle. Depending on its susceptibility to oxidation, this recycled carbon may be locked away from the biologically mediated carbon cycle for many geologic cycles.
Modern chitons (Mollusca: Polyplacophora) possess a highly conserved skeleton of eight shell plates (valves) surrounded by spicules or scales, and fossil evidence suggests that the chiton skeleton has changed little since the first appearance of the class in the Late Cambrian period (about 500 million years before present, Myr bp). However, the Palaeozoic problematic taxon Multiplacophora, in spite of having a more complex skeleton, shares several derived characters with chitons. The enigmatic status of the Multiplacophora is due in part to the fact that its members had an exoskeleton of numerous calcium carbonate valves that usually separated after death. A new articulated specimen from the Carboniferous period (about 335 Myr bp) of Indiana reveals that multiplacophorans had a dorsal protective surface composed of head and tail valves, left and right columns of overlapping valves (five on each side), and a central zone of five smaller valves, all surrounded by an annulus of large spines. Here we describe and name the articulated specimen and present evidence that multiplacophorans were chitons. Thus the highly conserved body plan of living chitons belies the broad disparity of this clade during the Palaeozoic era.
It is well known that the diversity of life appears to fluctuate during the course of the Phanerozoic, the eon during which hard shells and skeletons left abundant fossils (0-542 million years ago). Here we show, using Sepkoski's compendium of the first and last stratigraphic appearances of 36,380 marine genera, a strong 62 +/- 3-million-year cycle, which is particularly evident in the shorter-lived genera. The five great extinctions enumerated by Raup and Sepkoski may be an aspect of this cycle. Because of the high statistical significance we also consider the contributions of environmental factors, and possible causes.
Elevated carbon monoxide (CO) mixing ratios in excess of those derived from emissions inventories have been observed in plumes from one gas- and coal-fired power plant and three of four lignite coal-fired electric utility power plants observed in east and central Texas. Observations of elevated CO on days characterized by differing wind directions show that CO emissions from the lignite plants were relatively constant over time and cannot be ascribed to separate sources adjacent to the power plants. These three plants were found to be emitting CO at rates 22 to 34 times those tabulated in State and Federal emissions inventories. Elevated CO emissions from the gas- and coal-fired plant were highly variable on time scales of hours to days, in one case changing by a factor of 8 within an hour. Three other fossil-fueled power plants, including one lignite-fired plant observed during this study, did not emit substantial amounts of CO, suggesting that a combination of plant operating conditions and the use of lignite coal may contribute to the enhanced emissions. Observed elevated CO emissions from the three lignite plants, if representative of average operating conditions, represent an additional 30% of the annual total CO emissions from point sources for the state of Texas.
We use space-based observations of NO2 columns from the Global Ozone Monitoring Experiment (GOME) to derive monthly top-down NOx emissions for 2000 via inverse modeling with the GEOS-CHEM chemical transport model. Top-down NOx sources are partitioned among fuel combustion (fossil fuel and biofuel), biomass burning and soils by exploiting the spatio-temporal distribution of remotely sensed fires and a priori information on the location of regions dominated by fuel combustion. The top-down inventory is combined with an a priori inventory to obtain an optimized a posteriori estimate of the relative roles of NOx sources. The resulting a posteriori fuel combustion inventory (25.6 TgN year(-1)) agrees closely with the a priori (25.4 TgN year(-1)), and errors are reduced by a factor of 2, from +/- 80% to +/- 40%. Regionally, the largest differences are found over Japan and South Africa, where a posteriori estimates are 25% larger than a priori. A posteriori fuel combustion emissions are aseasonal, with the exception of East Asia and Europe where winter emissions are 30-40% larger relative to summer emissions, consistent with increased energy use during winter for heating. Global a posteriori biomass burning emissions in 2000 resulted in 5.8 TgN (compared to 5.9 TgN year(-1) in the a priori), with Africa accounting for half of this total. A posteriori biomass burning emissions over Southeast Asia/India are decreased by 46% relative to a priori; but over North equatorial Africa they are increased by 50%. A posteriori estimates of soil emissions (8.9 TgN year(-1)) are 68% larger than a priori (5.3 TgN year(-1)). The a posteriori inventory displays the largest soil emissions over tropical savanna/woodland ecosystems (Africa), as well as over agricultural regions in the western U.S. (Great Plains), southern Europe (Spain, Greece, Turkey), and Asia (North China Plain and North India), consistent with field measurements. Emissions over these regions are highest during summer at mid-latitudes and during the rainy season in the Tropics. We estimate that 2.5-4.5 TgN year(-1) are emitted from N-fertilized soils, at the upper end of previous estimates. Soil and biomass burning emissions account for 22% and 14% of global surface NOx emissions, respectively. We infer a significant role for soil NOx emissions at northern mid-latitudes during summer, where they account for nearly half that of the fuel combustion source, a doubling relative to the a priori. The contribution of soil emissions to background ozone is thus likely to be underestimated by the current generation of chemical transport models.
Even though echinoderms are members of the Bilateria, the location of their anterior/posterior axis has remained enigmatic. Here we propose a novel solution to the problem employing three lines of evidence: the expression of a posterior class Hox gene in the coeloms of the nascent adult body plan within the larva; the anatomy of certain early fossil echinoderms; and finally the relation between endoskeletal plate morphology and the associated coelomic tissues. All three lines of evidence converge on the same answer, namely that the location of the adult mouth is anterior, and the anterior/posterior axis runs from the mouth through the adult coelomic compartments. This axis then orients the animal such that there is but a single plane of symmetry dividing the animal into left and right halves. We tentatively hypothesize that this plane of symmetry is positioned along the dorsal/ventral axis. These axis identifications lead to the conclusion that the five ambulacra are not primary body axes, but instead are outgrowths from the central anterior/posterior axis. These identifications also shed insight into several other evolutionary mysteries of various echinoderm clades such as the independent evolution of bilateral symmetry in irregular echinoids, but do not elucidate the underlying mechanisms of the adult coelomic architecture.
The molecular tool kit for producing flat-bladed photosynthetic structures evolved in marine and terrestrial plants during the middle Paleozoic, but it took a further 20 million years before leaves suddenly spread throughout land floras. This delay has long been difficult to explain, given the apparent advantage of leaves for photosynthetic primary production. Theory and experiments predict that exceptionally high atmospheric CO2 levels in the middle Paleozoic delayed the origin of leaves by restricting stomatal development. This would have limited evaporative cooling, leading to lethal overheating of leaves absorbing large quantities of solar energy. Here we test the central prediction of this argument with a morphometric analysis of 300 plant fossils from major European collections. We show a 25-fold enlargement of leaf blades in two phylogenetically independent clades as atmospheric CO2 levels fell during the late Paleozoic. Furthermore, preliminary data suggest that the first abrupt increase in leaf size was accompanied by an 8-fold rise in stomatal density. These evolutionary patterns support the relaxation of biophysical constraints on leaf area predicted by theory and point to a significant role for CO2 in plant evolution.
Molecular clocks have the potential to shed light on the timing of early metazoan divergences, but differing algorithms and calibration points yield conspicuously discordant results. We argue here that competing molecular clock hypotheses should be testable in the fossil record, on the principle that fundamentally new grades of animal organization will have ecosystem-wide impacts. Using a set of seven nuclear-encoded protein sequences, we demonstrate the paraphyly of Porifera and calculate sponge/eumetazoan and cnidarian/bilaterian divergence times by using both distance [minimum evolution (ME)] and maximum likelihood (ML) molecular clocks; ME brackets the appearance of Eumetazoa between 634 and 604 Ma, whereas ML suggests it was between 867 and 748 Ma. Significantly, the ME, but not the ML, estimate is coincident with a major regime change in the Proterozoic acritarch record, including: (i) disappearance of low-diversity, evolutionarily static, pre-Ediacaran acanthomorphs; (ii) radiation of the high-diversity, short-lived Doushantuo-Pertatataka microbiota; and (iii) an order-of-magnitude increase in evolutionary turnover rate. We interpret this turnover as a consequence of the novel ecological challenges accompanying the evolution of the eumetazoan nervous system and gut. Thus, the more readily preserved microfossil record provides positive evidence for the absence of pre-Ediacaran eumetazoans and strongly supports the veracity, and therefore more general application, of the ME molecular clock.
The "false thumb" of pandas is a carpal bone, the radial sesamoid, which has been enlarged and functions as an opposable thumb. If the giant panda (Ailuropoda melanoleuca) and the red panda (Ailurus fulgens) are not closely related, their sharing of this adaptation implies a remarkable convergence. The discovery of previously unknown postcranial remains of a Miocene red panda relative, Simocyon batalleri, from the Spanish site of Batallones-1 (Madrid), now shows that this animal had a false thumb. The radial sesamoid of S. batalleri shows similarities with that of the red panda, which supports a sister-group relationship and indicates independent evolution in both pandas. The fossils from Batallones-1 reveal S. batalleri as a puma-sized, semiarboreal carnivore with a moderately hypercarnivore diet. These data suggest that the false thumbs of S. batalleri and Ailurus fulgens were probably inherited from a primitive member of the red panda family (Ailuridae), which lacked the red panda's specializations for herbivory but shared its arboreal adaptations. Thus, it seems that, whereas the false thumb of the giant panda probably evolved for manipulating bamboo, the false thumbs of the red panda and of S. batalleri more likely evolved as an aid for arboreal locomotion, with the red panda secondarily developing its ability for item manipulation and thus producing one of the most dramatic cases of convergence among vertebrates.
Earth history was punctuated during the Permo-Carboniferous [300-250 million years (Myr) ago] by the longest and most severe glaciation of the entire Phanerozoic Eon. But significant uncertainty surrounds the concentration of CO(2) in the atmosphere through this time interval and therefore its role in the evolution of this major prePleistocene glaciation. Here, I derive 24 Late Paleozoic CO(2) estimates from the fossil cuticle record of arborsecent lycopsids of the equatorial Carboniferous and Permian swamp communities. Quantitative calibration of Late Carboniferous (330-300 Myr ago) and Permian (270-260 Myr ago) lycopsid stomatal indices yield average atmospheric CO(2) concentrations of 344 ppm and 313 ppm, respectively. The reconstructions show a high degree of self-consistency and a degree of precision an order of magnitude greater than other approaches. Low CO(2) levels during the Permo-Carboniferous glaciation are in agreement with glaciological evidence for the presence of continental ice and coupled models of climate and ice-sheet growth on Pangea. Moreover, the Permian data indicate atmospheric CO(2) levels were low 260 Myr ago, by which time continental deglaciation was already underway. Positive biotic feedbacks on climate, and geotectonic events, therefore are implicated as mechanisms underlying deglaciation.