Low atmospheric CO(2) levels during the Permo- Carboniferous glaciation inferred from fossil lycopsids

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.     

Continued warming could transform Greater Yellowstone fire regimes by mid-21st century

Climate change is likely to alter wildfire regimes, but the magnitude and timing of potential climate-driven changes in regional fire regimes are not well understood. We considered how the occurrence, size, and spatial location of large fires might respond to climate projections in the Greater Yellowstone ecosystem (GYE) (Wyoming), a large wildland ecosystem dominated by conifer forests and characterized by infrequent, high-severity fire. We developed a suite of statistical models that related monthly climate data (1972-1999) to the occurrence and size of fires >200 ha in the northern Rocky Mountains; these models were cross-validated and then used with downscaled (~12 km × 12 km) climate projections from three global climate models to predict fire occurrence and area burned in the GYE through 2099. All models predicted substantial increases in fire by midcentury, with fire rotation (the time to burn an area equal to the landscape area) reduced to <30 y from the historical 100-300 y for most of the GYE. Years without large fires were common historically but are expected to become rare as annual area burned and the frequency of regionally synchronous fires increase. Our findings suggest a shift to novel fire-climate-vegetation relationships in Greater Yellowstone by midcentury because fire frequency and extent would be inconsistent with persistence of the current suite of conifer species. The predicted new fire regime would transform the flora, fauna, and ecosystem processes in this landscape and may indicate similar changes for other subalpine forests.     

Highly episodic fire and erosion regime over the past 2,000 y in the Siskiyou Mountains,Oregon

Fire is a primary mode of natural disturbance in the forests of the Pacific Northwest. Increased fuel loads following fire suppression and the occurrence of several large and severe fires have led to the perception that in many areas there is a greatly increased risk of high-severity fire compared with presettlement forests. To reconstruct the variability of the fire regime in the Siskiyou Mountains, Oregon, we analyzed a 10-m, 2,000-y sediment core for charcoal, pollen, and sedimentological data. The record reveals a highly episodic pattern of fire in which 77% of the 68 charcoal peaks before Euro-American settlement cluster within nine distinct periods marked by a 15-y mean interval. The 11 largest charcoal peaks are significantly related to decadal-scale drought periods and are followed by pulses of minerogenic sediment suggestive of rapid sediment delivery. After logging in the 1950s, sediment load was increased fourfold compared with that from the most severe presettlement fire. Less severe fires, marked by smaller charcoal peaks and no sediment pulses, are not correlated significantly with drought periods. Pollen indicators of closed forests are consistent with fire-free periods of sufficient length to maintain dense forest and indicate a fire-triggered switch to more open conditions during the Medieval Climatic Anomaly. Our results indicate that over millennia fire was more episodic than revealed by nearby shorter tree-ring records and that recent severe fires have precedents during earlier drought episodes but also that sediment loads resulting from logging and road building have no precedent in earlier fire events.     

Environmental and biotic controls on the evolutionary history of insect body size

Giant insects, with wingspans as large as 70 cm, ruled the Carboniferous and Permian skies. Gigantism has been linked to hyperoxic conditions because oxygen concentration is a key physiological control on body size, particularly in groups like flying insects that have high metabolic oxygen demands. Here we show, using a dataset of more than 10,500 fossil insect wing lengths, that size tracked atmospheric oxygen concentrations only for the first 150 Myr of insect evolution. The data are best explained by a model relating maximum size to atmospheric environmental oxygen concentration (pO(2)) until the end of the Jurassic, and then at constant sizes, independent of oxygen fluctuations, during the Cretaceous and, at a smaller size, the Cenozoic. Maximum insect size decreased even as atmospheric pO(2) rose in the Early Cretaceous following the evolution and radiation of early birds, particularly as birds acquired adaptations that allowed more agile flight. A further decrease in maximum size during the Cenozoic may relate to the evolution of bats, the Cretaceous mass extinction, or further specialization of flying birds. The decoupling of insect size and atmospheric pO(2) coincident with the radiation of birds suggests that biotic interactions, such as predation and competition, superseded oxygen as the most important constraint on maximum body size of the largest insects.     

Prolonged Permian Triassic ecological crisis recorded by molluscan dominance in Late Permian offshore assemblages

The end-Permian mass extinction was the largest biotic crisis in the history of animal life, eliminating as many as 95% of all species and dramatically altering the ecological structure of marine communities. Although the causes of this pronounced ecosystem shift have been widely debated, the broad consensus based on inferences from global taxonomic diversity patterns suggests that the shift from abundant brachiopods to dominant molluscs was abrupt and largely driven by the catastrophic effects of the end-Permian mass extinction. Here we analyze relative abundance counts of >33,000 fossil individuals from 24 silicified Middle and Late Permian paleocommunities, documenting a substantial ecological shift to numerical dominance by molluscs in the Late Permian, before the major taxonomic shift at the end-Permian mass extinction. This ecological change was coincident with the development of fluctuating anoxic conditions in deep marine basins, suggesting that numerical dominance by more tolerant molluscs may have been driven by variably stressful environmental conditions. Recognition of substantial ecological deterioration in the Late Permian also implies that the end-Permian extinction was the climax of a protracted environmental crisis. Although the Late Permian shift to molluscan dominance was a pronounced ecological change, quantitative counts of 847 Carboniferous-Cretaceous collections from the Paleobiology Database indicate that it was only the first stage in a stepwise transition that culminated with the final shift to molluscan dominance in the Late Jurassic. Therefore, the ecological transition from brachiopods to bivalves was more protracted and complex than their simple Permian-Triassic switch in diversity.     

Baseline intrinsic flammability of Earth's ecosystems estimated from paleoatmospheric oxygen over the past 350 million years

Atmospheric oxygen (O(2)) is estimated to have varied greatly throughout Earth's history and has been capable of influencing wildfire activity wherever fuel and ignition sources were present. Fires consume huge quantities of biomass in all ecosystems and play an important role in biogeochemical cycles. This means that understanding the influence of O(2) on past fire activity has far-reaching consequences for the evolution of life and Earth's biodiversity over geological timescales. We have used a strong electrical ignition source to ignite smoldering fires, and we measured their self-sustaining propagation in atmospheres of different oxygen concentrations. These data have been used to build a model that we use to estimate the baseline intrinsic flammability of Earth's ecosystems according to variations in O(2) over the past 350 million years (Ma). Our aim is to highlight times in Earth's history when fire has been capable of influencing the Earth system. We reveal that fire activity would be greatly suppressed below 18.5% O(2), entirely switched off below 16% O(2), and rapidly enhanced between 19-22% O(2). We show that fire activity and, therefore, its influence on the Earth system would have been high during the Carboniferous (350-300 Ma) and Cretaceous (145-65 Ma) periods; intermediate in the Permian (299-251 Ma), Late Triassic (285-201 Ma), and Jurassic (201-145 Ma) periods; and surprisingly low to lacking in the Early-Middle Triassic period between 250-240 Ma. These baseline variations in Earth's flammability must be factored into our understanding of past vegetation, biodiversity, evolution, and biogeochemical cycles.     

Date of the monocot-dicot divergence estimated from chloroplast DNA sequence data.

The divergence between monocots and dicots represents a major event in higher plant evolution, yet the date of its occurrence remains unknown because of the scarcity of relevant fossils. We have estimated this date by reconstructing phylogenetic trees from chloroplast DNA sequences, using two independent approaches: the rate of synonymous nucleotide substitution was calibrated from the divergence of maize, wheat, and rice, whereas the rate of nonsynonymous substitution was calibrated from the divergence of angiosperms and bryophytes. Both methods lead to an estimate of the monocot-dicot divergence at 200 million years (Myr) ago (with an uncertainty of about 40 Myr). This estimate is also supported by analyses of the nuclear genes encoding large and small subunit ribosomal RNAs. These results imply that the angiosperm lineage emerged in Jurassic-Triassic time, which considerably predates its appearance in the fossil record (approximately 120 Myr ago). We estimate the divergence between cycads and angiosperms to be approximately 340 Myr, which can be taken as an upper bound for the age of angiosperms.     

Climate regulation of fire emissions and deforestation in equatorial Asia

Drainage of peatlands and deforestation have led to large-scale fires in equatorial Asia, affecting regional air quality and global concentrations of greenhouse gases. Here we used several sources of satellite data with biogeochemical and atmospheric modeling to better understand and constrain fire emissions from Indonesia, Malaysia, and Papua New Guinea during 2000–2006. We found that average fire emissions from this region [128 ± 51 (1σ) Tg carbon (C) year⁻¹, T = 10¹²] were comparable to fossil fuel emissions. In Borneo, carbon emissions from fires were highly variable, fluxes during the moderate 2006 El Niño more than 30 times greater than those during the 2000 La Niña (and with a 2000–2006 mean of 74 ± 33 Tg C yr⁻¹). Higher rates of forest loss and larger areas of peatland becoming vulnerable to fire in drought years caused a strong nonlinear relation between drought and fire emissions in southern Borneo. Fire emissions from Sumatra showed a positive linear trend, increasing at a rate of 8 Tg C year⁻² (approximately doubling during 2000–2006). These results highlight the importance of including deforestation in future climate agreements. They also imply that land manager responses to expected shifts in tropical precipitation may critically determine the strength of climate–carbon cycle feedbacks during the 21st century.     

Two-phase increase in the maximum size of life over 3.5 billion years reflects biological innovation and environmental opportunity

The maximum size of organisms has increased enormously since the initial appearance of life >3.5 billion years ago (Gya), but the pattern and timing of this size increase is poorly known. Consequently, controls underlying the size spectrum of the global biota have been difficult to evaluate. Our period-level compilation of the largest known fossil organisms demonstrates that maximum size increased by 16 orders of magnitude since life first appeared in the fossil record. The great majority of the increase is accounted for by 2 discrete steps of approximately equal magnitude: the first in the middle of the Paleoproterozoic Era (≈1.9 Gya) and the second during the late Neoproterozoic and early Paleozoic eras (0.6–0.45 Gya). Each size step required a major innovation in organismal complexity—first the eukaryotic cell and later eukaryotic multicellularity. These size steps coincide with, or slightly postdate, increases in the concentration of atmospheric oxygen, suggesting latent evolutionary potential was realized soon after environmental limitations were removed.     

Response of C3 and C4 plants to middle-Holocene climatic variation near the prairie-forest ecotone of Minnesota

Paleorecords of the middle Holocene (MH) from the North American mid-continent can offer insights into ecological responses to pervasive drought that may accompany future climatic warming. We analyzed MH sediments from West Olaf Lake (WOL) and Steel Lake (SL) in Minnesota to examine the effects of warm/dry climatic conditions on prairie-woodland ecosystems. Mineral composition and carbonate delta(18)O were used to determine climatic variations, whereas pollen assemblages, charcoal delta(13)C, and charcoal accumulation rates were used to reconstruct vegetation composition, C(3) and C(4) plant abundance, and fire. The ratio of aragonite/calcite at WOL and delta(18)O at SL suggest that pronounced droughts occurred during the MH but that drought severity decreased with time. From charcoal delta(13)C data we estimated that the MH abundance of C(4) plants averaged 50% at WOL and 43% at SL. At WOL C(4) abundance was negatively correlated with aragonite/calcite, suggesting that severe moisture deficits suppressed C(4) plants in favor of weedy C(3) plants (e.g., Ambrosia). As climate ameliorated C(4) abundance increased (from approximately 33 to 66%) at the expense of weedy species, enhancing fuel availability and fire occurrence. In contrast, farther east at SL where climate was cooler and wetter, C(4) abundance showed no correlation with delta(18)O-inferred aridity. Woody C(3) plants (e.g., Quercus) were more abundant, biomass flammability was lower, and fires were less important at SL than at WOL. Our results suggest that C(4) plants are adapted to warm/dry climatic conditions, but not to extreme droughts, and that the fire regime is controlled by biomass-climate interactions.     

On the evolutionary history of Ephedra: Cretaceous fossils and extant molecules

Gnetales comprise three unusual genera of seed plants, Ephedra, Gnetum, and Welwitschia. Their extraordinary morphological diversity suggests that they are survivors of an ancient, more diverse group. Gnetalean antiquity is also supported by fossil data. Dispersed "ephedroid" (polyplicate) pollen first appeared in the Permian >250 million years ago (Myr), and a few megafossils document the presence of gnetalean features in the early Cretaceous. The Cretaceous welwitschioid seedling Cratonia cotyledon dates the split between Gnetum and Welwitschia to before 110 Myr. Ages and character evolution of modern diversity are, however, controversial, and, based on molecular data, it has recently been suggested that Ephedra is very young, only 8-32 Myr. Here, we present data on the evolutionary history of Ephedra. Fossil seeds from Buarcos, Portugal, unequivocally link one type of Cretaceous polyplicate pollen to Ephedra and document that plants with unique characters, including the peculiar naked male gametophyte, were established already in the Early Cretaceous. Clades in our molecular phylogeny of extant species correspond to geographical regions, with African species in a basal grade/clade. The study demonstrates extremely low divergence in both molecular and morphological characters in Ephedra. Features observed in the fossils are present in all major extant clades, showing that modern species have retained unique reproductive characters for >110 million years. A recent origin of modern species of Ephedra would imply that the Cretaceous Ephedra fossils discussed here were members of widespread, now extinct sister lineage(s), and that no morphological innovations characterized the second diversification.     

Effects of sampling standardization on estimates of Phanerozoic marine diversification

Global diversity curves reflect more than just the number of taxa that have existed through time: they also mirror variation in the nature of the fossil record and the way the record is reported. These sampling effects are best quantified by assembling and analyzing large numbers of locality-specific biotic inventories. Here, we introduce a new database of this kind for the Phanerozoic fossil record of marine invertebrates. We apply four substantially distinct analytical methods that estimate taxonomic diversity by quantifying and correcting for variation through time in the number and nature of inventories. Variation introduced by the use of two dramatically different counting protocols also is explored. We present sampling-standardized diversity estimates for two long intervals that sum to 300 Myr (Middle Ordovician-Carboniferous; Late Jurassic-Paleogene). Our new curves differ considerably from traditional, synoptic curves. For example, some of them imply unexpectedly low late Cretaceous and early Tertiary diversity levels. However, such factors as the current emphasis in the database on North America and Europe still obscure our view of the global history of marine biodiversity. These limitations will be addressed as the database and methods are refined.     

Oxygen-Collagen Priority and the Early Metazoan Fossil Record

The thesis is developed that a low oxygen level Precambrian atmosphere presented early-evolving metazoan organisms with physiological connective tissue priorities resulting from the important molecular oxygen requirements in the biosynthesis of collagen hydroxyproline. Shells, cuticles, and carapaces which are not mandatory metazoan prerequisites but which directly or indirectly demand substantial connective tissue collagen are oxygen expensive, low priority features. A marked increase in atmospheric oxygen level near the beginning of the Paleozoic would eliminate oxygen-collagen priorities simultaneously and on a world-wide basis in all metazoan stocks providing evolutionary pressure for enlarged musculatures and associated "hard parts." This could explain the sudden presence in the fossil record of the early Cambrian of advanced and diversified metazoans, the earlier forms of which were essentially unpreservable.     

Earliest Carboniferous tetrapod and arthropod faunas from Scotland populate Romer's Gap

Devonian tetrapods (limbed vertebrates), known from an increasingly large number of localities, have been shown to be mainly aquatic with many primitive features. In contrast, the post-Devonian record is marked by an Early Mississippian temporal gap ranging from the earliest Carboniferous (Tournaisian and early Viséan) to the mid-Viséan. By the mid-Viséan, tetrapods had become effectively terrestrial as attested by the presence of stem amniotes, developed an essentially modern aspect, and given rise to the crown group. Up to now, only two localities have yielded tetrapod specimens from the Tournaisian stage: one in Scotland with a single articulated skeleton and one in Nova Scotia with isolated bones, many of uncertain identity. We announce a series of discoveries of Tournaisian-age localities in Scotland that have yielded a wealth of new tetrapod and arthropod fossils. These include both terrestrial and aquatic forms and new taxa. We conclude that the gap in the fossil record has been an artifact of collection failure.     

Cretaceous flowers of Nymphaeaceae and implications for complex insect entrapment pollination mechanisms in early angiosperms

Based on recent molecular systematics studies, the water lily lineage (Nymphaeales) provides an important key to understanding ancestral angiosperm morphology and is of considerable interest in the context of angiosperm origins. Therefore, the fossil record of Nymphaeales potentially provides evidence on both the timing and nature of diversification of one of the earliest clades of flowering plants. Recent fossil evidence of Turonian age (approximately 90 million years B.P.) includes fossil flowers with characters that, upon rigorous analysis, firmly place them within Nymphaeaceae. Unequivocally the oldest floral record of the Nymphaeales, these fossils are closely related to the modern Nymphaealean genera Victoria (the giant Amazon water lily) and Euryale. Although the fossils are much smaller than their modern relatives, the precise and dramatic correspondence between the fossil floral morphology and that of modern Victoria flowers suggests that beetle entrapment pollination was present in the earliest part of the Late Cretaceous.     

Exceptionally preserved crustaceans from western Canada reveal a cryptic Cambrian radiation

The early history of crustaceans is obscured by strong biases in fossil preservation, but a previously overlooked taphonomic mode yields important complementary insights. Here we describe diverse crustacean appendages of Middle and Late Cambrian age from shallow-marine mudstones of the Deadwood Formation in western Canada. The fossils occur as flattened and fragmentary carbonaceous cuticles but provide a suite of phylogenetic and ecological data by virtue of their detailed preservation. In addition to an unprecedented range of complex, largely articulated filtering limbs, we identify at least four distinct types of mandible. Together, these fossils provide the earliest evidence for crown-group branchiopods and total-group copepods and ostracods, extending the respective ranges of these clades back from the Devonian, Pennsylvanian, and Ordovician. Detailed similarities with living forms demonstrate the early origins and subsequent conservation of various complex food-handling adaptations, including a directional mandibular asymmetry that has persisted through half a billion years of evolution. At the same time, the Deadwood fossils indicate profound secular changes in crustacean ecology in terms of body size and environmental distribution. The earliest radiation of crustaceans is largely cryptic in the fossil record, but "small carbonaceous fossils" reveal organisms of surprisingly modern aspect operating in an unfamiliar biosphere.     

Life in the end-Permian dead zone

The fossil record of land plants is an obvious source of information on the dynamics of mass extinctions in the geological past. In conjunction with the end-Permian ecological crisis, approximately 250 million years ago, palynological data from East Greenland reveal some unanticipated patterns. We document the significant time lag between terrestrial ecosystem collapse and selective extinction among characteristic Late Permian plants. Furthermore, ecological crisis resulted in an initial increase in plant diversity, instead of a decrease. Paradoxically, these floral patterns correspond to a "dead zone" in the end-Permian faunal record, characterized by a paucity of marine invertebrate megafossils. The time-delayed, end-Permian plant extinctions resemble modeled "extinction debt" responses of multispecies metapopulations to progressive habitat destruction.     

Rapid landscape transformation in South Island, New Zealand, following initial Polynesian settlement

Humans have altered natural patterns of fire for millennia, but the impact of human-set fires is thought to have been slight in wet closed-canopy forests. In the South Island of New Zealand, Polynesians (Māori), who arrived 700-800 calibrated years (cal y) ago, and then Europeans, who settled ～150 cal y ago, used fire as a tool for forest clearance, but the structure and environmental consequences of these fires are poorly understood. High-resolution charcoal and pollen records from 16 lakes were analyzed to reconstruct the fire and vegetation history of the last 1,000 y. Diatom, chironomid, and element concentration data were examined to identify disturbance-related limnobiotic and biogeochemical changes within burned watersheds. At most sites, several high-severity fire events occurred within the first two centuries of Māori arrival and were often accompanied by a transformation in vegetation, slope stability, and lake chemistry. Proxies of past climate suggest that human activity alone, rather than unusually dry or warm conditions, was responsible for this increased fire activity. The transformation of scrub to grassland by Europeans in the mid-19th century triggered further, sometimes severe, watershed change, through additional fires, erosion, and the introduction of nonnative plant species. Alteration of natural disturbance regimes had lasting impacts, primarily because native forests had little or no previous history of fire and little resilience to the severity of burning. Anthropogenic burning in New Zealand highlights the vulnerability of closed-canopy forests to novel disturbance regimes and suggests that similar settings may be less resilient to climate-induced changes in the future.     

Late Carboniferous paleoichnology reveals the oldest full-body impression of a flying insect

Insects were the first animals to evolve powered flight and did so perhaps 90 million years before the first flight among vertebrates. However, the earliest fossil record of flying insect lineages (Pterygota) is poor, with scant indirect evidence from the Devonian and a nearly complete dearth of material from the Early Carboniferous. By the Late Carboniferous a diversity of flying lineages is known, mostly from isolated wings but without true insights into the paleoethology of these taxa. Here, we report evidence of a full-body impression of a flying insect from the Late Carboniferous Wamsutta Formation of Massachusetts, representing the oldest trace fossil of Pterygota. Through ethological and morphological analysis, the trace fossil provides evidence that its maker was a flying insect and probably was representative of a stem-group lineage of mayflies. The nature of this current full-body impression somewhat blurs distinctions between the systematics of traces and trace makers, thus adding to the debate surrounding ichnotaxonomy for traces with well-associated trace makers.     

The magnetic polarity stratigraphy of the Mauch Chunk Formation, Pennsylvania

Three sections of Chesterian Mauch Chunk Formation in Pennsylvania have been studied paleomagnetically to determine a Late Mississippian magnetic polarity stratigraphy. The upper section at Lavelle includes a conglomerate with abundant red siltstone rip-up clasts that yielded a positive conglomerate test. All samples were subjected to progressive thermal demagnetization to temperatures as high as 700 degrees C. Two components of magnetization were isolated: a synfolding "B" component and the prefolding "C" component. The conglomerate test is positive, indicating that the C component was acquired very early in the history of the sediment. A coherent pattern of magnetic polarity reversals was identified. Five magnetozones were identified in the upper Lavelle section, which yields a pattern that is an excellent match with the pattern of reversals obtained from the upper Mauch Chunk at the original type section of the Mississippian/Pennsylvanian boundary at Pottsville, PA. The frequency of reversals in the upper Mississippian, as identified in the Mauch Chunk Formation, is approximately one to two per million years, which is an average for field reversal through time.