Freeze tolerance influenced forest cover and hydrology during the Pennsylvanian.

The distribution of forest cover alters Earth surface mass and energy exchange and is controlled by physiology, which determines plant environmental limits. Ancient plant physiology, therefore, likely affected vegetation-climate feedbacks. We combine climate modeling and ecosystem-process modeling to simulate arboreal vegetation in the late Paleozoic ice age. Using GENESIS V3 global climate model simulations, varying pCO2, pO2, and ice extent for the Pennsylvanian, and fossil-derived leaf C:N, maximum stomatal conductance, and specific conductivity for several major Carboniferous plant groups, we simulated global ecosystem processes at a 2° resolution with Paleo-BGC. Based on leaf water constraints, Pangaea could have supported widespread arboreal plant growth and forest cover. However, these models do not account for the impacts of freezing on plants. According to our interpretation, freezing would have affected plants in 59% of unglaciated land during peak glacial periods and 73% during interglacials, when more high-latitude land was unglaciated. Comparing forest cover, minimum temperatures, and paleo-locations of Pennsylvanian-aged plant fossils from the Paleobiology Database supports restriction of forest extent due to freezing. Many genera were limited to unglaciated land where temperatures remained above -4 °C. Freeze-intolerance of Pennsylvanian arboreal vegetation had the potential to alter surface runoff, silicate weathering, CO2 levels, and climate forcing. As a bounding case, we assume total plant mortality at -4 °C and estimate that contracting forest cover increased net global surface runoff by up to 6.1%. Repeated freezing likely influenced freeze- and drought-tolerance evolution in lineages like the coniferophytes, which became increasingly dominant in the Permian and early Mesozoic.

Carboniferous plant physiology breaks the mold.

How plants have shaped Earth surface feedbacks over geologic time is a key question in botanical and geological inquiry. Recent work has suggested that biomes during the Carboniferous Period contained plants with extraordinary physiological capacity to shape their environment, contradicting the previously dominant view that plants only began to actively moderate the Earth's surface with the rise of angiosperms during the Mesozoic Era. A recently published Viewpoint disputes this recent work, thus here, we document in detail, the mechanistic underpinnings of our modeling and illustrate the extraordinary ecophysiological nature of Carboniferous plants.

Paleoecological and paleoenvironmental interpretation of three successive macrofloras and palynofloras from the Kola Switch locality, lower Permian (Archer City Formation, Bowie Group) of Clay County, Texas, USA.

Fossil floras have been recovered from a unique deposit of early Permian age in North-Central Texas. The site, Kola Switch, preserves three distinct floras in different lithofacies, in a succession from a single outcrop. The sedimentary environment appears to be a floodplain channel fill of primarily siltstones and claystones. The lowermost flora, preserved in a kaolinitic siltstone, indicates active water flow. It is dominated by plants typical of well-drained substrates, dominated by Sphenopteris germanica, and contains no wetland elements. The middle flora is from a finely laminated carbonaceous claystone and is dominated by marattialean tree ferns, with no elements from habitats typical of seasonal moisture availability. It contains no roots and appears to have formed as a floating peat mat. The upper flora is a mixed assemblage of wetland taxa and those typical of well-drained soil environments or a seasonal rainfall regime. Unlike the two lower floras, it has a relatively even distribution of dominance and is the most diverse of the three assemblages. Palynofloras also were recovered from each of these beds. The palynofloras, although varying between and even within the beds, indicate a common background species pool during the time interval sampled, suggesting that these distinct floras reflect local changes in microhabitat conditions under a constant climatic background. The palynoflora from each bed has characteristics in common with the macroflora of that bed, but also distinct differences. Together, the macroflora and microflora provide an unusually broad picture of this site through time. Kola Switch compares favorably with the recently described flora from the nearby Sanzenbacher Ranch site of approximately the same age and also with floras of Rotliegend age from Central Europe.

A hidden cradle of plant evolution in Permian tropical lowlands.

The latitudinal biodiversity gradient today has deep roots in the evolutionary history of Earth's biota over geologic time. In the marine realm, earliest fossil occurrences at low latitudes reveal a tropical cradle for many animal groups. However, the terrestrial fossil record-especially from drier environments that are thought to drive evolutionary innovation-is sparse. We present mixed plant-fossil assemblages from Permian equatorial lowlands in present-day Jordan that harbor precocious records of three major seed-plant lineages that all became dominant during the Mesozoic, including the oldest representative of any living conifer family. These finds offer a glimpse of the early evolutionary origins of modern plant groups in disturbance-prone tropical habitats that are usually hidden from observation.

Dynamic Carboniferous tropical forests: new views of plant function and potential for physiological forcing of climate.

Contents 1333 I. 1334 II. 1335 III. 1339 IV. 1344 V. 1347 VI. 1347 1348 1348 References 1348 SUMMARY: The Carboniferous, the time of Earth's penultimate icehouse and widespread coal formation, was dominated by extinct lineages of early-diverging vascular plants. Studies of nearest living relatives of key Carboniferous plants suggest that their physiologies and growth forms differed substantially from most types of modern vegetation, particularly forests. It remains a matter of debate precisely how differently and to what degree these long-extinct plants influenced the environment. Integrating biophysical analysis of stomatal and vascular conductivity with geochemical analysis of fossilized tissues and process-based ecosystem-scale modeling yields a dynamic and unique perspective on these paleoforests. This integrated approach indicates that key Carboniferous plants were capable of growth and transpiration rates that approach values found in extant crown-group angiosperms, differing greatly from comparatively modest rates found in their closest living relatives. Ecosystem modeling suggests that divergent stomatal conductance, leaf sizes and stem life span between dominant clades would have shifted the balance of soil-atmosphere water fluxes, and thus surface runoff flux, during repeated, climate-driven, vegetation turnovers. This synthesis highlights the importance of 'whole plant' physiological reconstruction of extinct plants and the potential of vascular plants to have influenced the Earth system hundreds of millions of years ago through vegetation-climate feedbacks.

Delayed fungal evolution did not cause the Paleozoic peak in coal production.

Organic carbon burial plays a critical role in Earth systems, influencing atmospheric O2 and CO2 concentrations and, thereby, climate. The Carboniferous Period of the Paleozoic is so named for massive, widespread coal deposits. A widely accepted explanation for this peak in coal production is a temporal lag between the evolution of abundant lignin production in woody plants and the subsequent evolution of lignin-degrading Agaricomycetes fungi, resulting in a period when vast amounts of lignin-rich plant material accumulated. Here, we reject this evolutionary lag hypothesis, based on assessment of phylogenomic, geochemical, paleontological, and stratigraphic evidence. Lignin-degrading Agaricomycetes may have been present before the Carboniferous, and lignin degradation was likely never restricted to them and their class II peroxidases, because lignin modification is known to occur via other enzymatic mechanisms in other fungal and bacterial lineages. Furthermore, a large proportion of Carboniferous coal horizons are dominated by unlignified lycopsid periderm with equivalent coal accumulation rates continuing through several transitions between floral dominance by lignin-poor lycopsids and lignin-rich tree ferns and seed plants. Thus, biochemical composition had little relevance to coal accumulation. Throughout the fossil record, evidence of decay is pervasive in all organic matter exposed subaerially during deposition, and high coal accumulation rates have continued to the present wherever environmental conditions permit. Rather than a consequence of a temporal decoupling of evolutionary innovations between fungi and plants, Paleozoic coal abundance was likely the result of a unique combination of everwet tropical conditions and extensive depositional systems during the assembly of Pangea.

Holocene shifts in the assembly of plant and animal communities implicate human impacts.

Understanding how ecological communities are organized and how they change through time is critical to predicting the effects of climate change. Recent work documenting the co-occurrence structure of modern communities found that most significant species pairs co-occur less frequently than would be expected by chance. However, little is known about how co-occurrence structure changes through time. Here we evaluate changes in plant and animal community organization over geological time by quantifying the co-occurrence structure of 359,896 unique taxon pairs in 80 assemblages spanning the past 300 million years. Co-occurrences of most taxon pairs were statistically random, but a significant fraction were spatially aggregated or segregated. Aggregated pairs dominated from the Carboniferous period (307 million years ago) to the early Holocene epoch (11,700 years before present), when there was a pronounced shift to more segregated pairs, a trend that continues in modern assemblages. The shift began during the Holocene and coincided with increasing human population size and the spread of agriculture in North America. Before the shift, an average of 64% of significant pairs were aggregated; after the shift, the average dropped to 37%. The organization of modern and late Holocene plant and animal assemblages differs fundamentally from that of assemblages over the past 300 million years that predate the large-scale impacts of humans. Our results suggest that the rules governing the assembly of communities have recently been changed by human activity.

Growth habit of the late Paleozoic rhizomorphic tree-lycopsid family Diaphorodendraceae: phylogenetic, evolutionary, and paleoecological significance.

PREMISE OF THE STUDY: Rhizomorphic lycopsids evolved the tree habit independently of all other land plants. Newly discovered specimens allow radical revision of our understanding of the growth architectures of the extinct Paleozoic sister-genera Synchysidendron and Diaphorodendron. METHODS: Detailed descriptions of six remarkable adpression specimens from the Pennsylvanian of the USA and three casts from the late Mississippian of Scotland are used to revise and reanalyze a previously published morphological cladistic matrix and to reinterpret their remarkable growth forms. KEY RESULTS: Contrary to previous assertions, Synchysidendron resembled Diaphorodendron in having a distinct and relatively complex growth habit that emphasized serially homologous, closely spaced, deciduous lateral branches at the expense of reduced monocarpic crown branches. Lateral branches originated through several strongly anisotomous dichotomies before producing during extended periods large numbers of Achlamydocarpon strobili. The comparatively large diameter of abscission scars remaining on the main trunk and the emergence of branches above the horizontal plane suggest that the lateral branch systems were robust. Lateral branches were borne in two opposite rows on the main trunk and continued upward into an isotomously branched, determinate crown; their striking distichous arrangement caused preferred orientation of fallen trunks on bedding planes. CONCLUSIONS: This discovery identifies the plagiotropic growth habit, dominated by serial lateral branches, as ubiquitous in the Diaphorodendraceae and also as unequivocally primitive within Isoetales s.l., a conclusion supported by both the revised morphological cladistic analysis and relative first appearances of taxa in the fossil record. Previously assumed complete homology between crown branching in Lepidodendraceae and that of all earlier-divergent genera requires reassessment. Saltational phenotypic transitions via modification of key developmental switches remains the most credible explanation for architectural evolution in the group. The resulting architecture allowed Diaphorodendraceae to co-dominate disturbed, clastic, equatorial wetlands from the Asbian to the Early Permian.

Conservatism of Late Pennsylvanian vegetational patterns during short-term cyclic and long-term directional environmental change, western equatorial Pangea.

Patterns of plant distribution by palaeoenvironment were examined across the Pennsylvanian-Permian transition in North-Central Texas. Stratigraphically recurrent packages of distinct lithofacies, representing different habitats, contain qualitatively and quantitatively different macrofloras and microfloras. The species pools demonstrate niche conservatism, remaining closely tied to specific habitats, during both short-term cyclic environmental change and a long-term trend of increasing aridity. The deposits examined principally comprise the terrestrial Markley and its approximate marine equivalent, the Harpersville Formation and parts of lower Archer City Formation. Fossiliferous deposits are lens-like, likely representing fill sequences of channels formed during abandonment phases. Palaeosols, represented by blocky mudstones, comprise a large fraction of the deposits. They suggest progressive climate change from minimally seasonal humid to seasonal subhumid to seasonal dry subhumid. Five lithofacies yielded plants: kaolinite-dominated siltstone, organic shale, mudstone beds within organic shale, coarsening upward mudstone-sandstone interbeds and channel sandstone. Both macro- and microflora were examined. Lithofacies proved compositionally distinct, with different patterns of dominance diversity. Organic shales (swamp deposits), mudstone partings (swamp drainages) and coarsening upward mudstone-sandstone interbeds (floodplains) typically contain Pennsylvanian wetland vegetation. Kaolinite-dominated siltstones and (to the extent known) sandstones contain taxa indicative of seasonally dry substrates. Some kaolinite-dominated siltstones and organic shales/coals yielded palynomorphs. Microfloras are more diverse, with greater wetland-dryland overlap than macrofloras. It appears that these two floras were coexistent at times on the regional landscape.

CO2-forced climate and vegetation instability during Late Paleozoic deglaciation.

The late Paleozoic deglaciation is the vegetated Earth's only recorded icehouse-to-greenhouse transition, yet the climate dynamics remain enigmatic. By using the stable isotopic compositions of soil-formed minerals, fossil-plant matter, and shallow-water brachiopods, we estimated atmospheric partial pressure of carbon dioxide (pCO2) and tropical marine surface temperatures during this climate transition. Comparison to southern Gondwanan glacial records documents covariance between inferred shifts in pCO2, temperature, and ice volume consistent with greenhouse gas forcing of climate. Major restructuring of paleotropical flora in western Euramerica occurred in step with climate and pCO2 shifts, illustrating the biotic impact associated with past CO2-forced turnover to a permanent ice-free world.

THE PENNSYLVANIAN-PERMIAN VEGETATIONAL TRANSITION: A TERRESTRIAL ANALOGUE TO THE ONSHORE-OFFSHORE HYPOTHESIS.

An analysis of 68 floras from the Pennsylvanian and Early Permian of Euramerica reveals distinct patterns of environmental distribution. Wetland assemblages are the most commonly encountered floras from the Early and Middle Pennsylvanian. Floras from drier habitats characterize the Permian. Both wetland and dry-site floras occur in the Late Pennsylvanian, but floristic overlap is minimal, which implies strong environmental controls on the distributions of the component species. Drier habitats appear to be the sites of first appearance of orders that become prominent during the Late Permian and Mesozoic. Higher taxa originated in physically heterogeneous, drier habitats, which were geographically marginal throughout most of the Pennsylvanian. They then moved into the lowlands during periods of climatic drying in the Permian, replacing older wetland vegetation. This pattern is analogous to the marine onshore-offshore pattern of origination and migration. The derivation of Mesozoic wetland clades from the Permian dry-lowland vegetation completes the parallel. The similarities of the marine and terrestrial patterns suggest that the combination of evolutionary opportunity, created by physical heterogeneity of the environment, and migrational opportunity, created by changing extrinsic conditions, may be underlying factors that transcend the specifics of organism and environment.