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Fire and the Miocene expansion of C4 grasslands

C 4 photosynthesis had a mid-Tertiary origin that was tied to declining atmospheric CO 2 , but C 4 -dominated grasslands did not appear until late Tertiary. According to the ‘CO 2 -threshold’ model, these C 4 grasslands owe their origin to a further late Miocene decline in CO 2 that gave C 4 grasses a photosynthetic advantage. This model is most appropriate for explaining replacement of C 3 grasslands by C 4 grasslands, however, fossil evidence shows C 4 grasslands replaced woodlands. An additional weakness in the threshold model is that recent estimates do not support a late Miocene drop in p CO 2 . We hypothesize that late Miocene climate changes created a fire climate capable of replacing woodlands with C 4 grasslands. Critical elements were seasonality that sustained high biomass production part of year, followed by a dry season that greatly reduced fuel moisture, coupled with a monsoon climate that generated abundant lightning-igniting fires. As woodlands became more open from burning, the high light conditions favoured C 4 grasses over C 3 grasses, and in a feedback process, the elevated productivity of C 4 grasses increased highly combustible fuel loads that further increased fire activity. This hypothesis is supported by paleosol data that indicate the late Miocene expansion of C 4 grasslands was the result of grassland expansion into more mesic environments and by charcoal sediment profiles that parallel the late Miocene expansion of C 4 grasslands. Many contemporary C 4 grasslands are fire dependent and are invaded by woodlands upon cessation of burning. Thus, we maintain that the factors driving the late Miocene expansion of C 4 were the same as those responsible for maintenance of C 4 grasslands today.

Ecology Letters

The California Valley grassland

Grasslands are distributed throughout California from Oregon to Baja California Norte and from the coast to the desert (Brown 1982) (Figure 1). This review will focus on the dominant formation in cismontane California, a community referred to as Valley Grassland (Munz 1959). Today, Valley Grassland is dominated by non-native annual grasses in genera such as Avena (wild oat), Bromus (brome grass), and Hordeum (barley), and is often referred to as the California annual grassland. On localized sites, native perennial bunchgrasses such as Stipa pultra (purple needle grass) may dominate and such sites are interpreted to be remnants of the pristine valley grassland. In northwestern California a floristically distinct formation of the Valley Grassland, known as Coast Prairie (Munz 1959) or Northern Coastal Grassland (Holland and Keil 1989) is recognized. The dominant grasses include many native perennial bunchgrasses in genera such as Agrostis , Calamagrostis , Danthonia , Deschampsia , Festuca , Koeleria and Poa (Heady et al. 1977). Non-native annuals do not dominate, but on some sites non-native perennials like Anthoxanthum odoratum may colonize the native grassland (Foin and Hektner 1986). Elevationally, California's grasslands extend from sea level to at leas 1500 m. The upper boundary is vague because montane grassland formations are commonly referred to as meadows; a community which Munz (1959) does not recognize. Holland and Keil (1989) describe the montane meadow as an azonal community; that is, a community restricted not so much to a particular climatic zone but rather controlled by substrate characteristics. They consider poor soil-drainage an over-riding factor in the development of montane meadows and, in contrast to grasslands, meadows often remain green through the summer drought. Floristically, meadows are composed of graminoids; Cyperaceae, Juncaceae, and rhizomatous grasses such as Agropyron (wheat grass). Some bunchgrasses, such as Muhlenbergia rigens , are found in both montane meadows and moister grasslands. Forbs when present, are typically perennials. East of the interior ranges, grasslands are uncommon although native perennial bunchgrasses in genera such as Stipa , Hilaria and Aristida are common in steppe and desert scrub. Today, Valley Grassland covers nearly 7 million ha or 17% or the state (Huenneke 1989), although other sources list less than half this amount (Jones and Stokes 1987). There is some evidence that extent of the grassland region has not changed since pre-European conditions, although the spatial distribution of grasslands has likely changed substantially (Huenneke 1989). That is, many current grasslands previously may have been dominated by other vegetation types and vice versa. Without question, many former grasslands have been converted to agricultural and urban use (Barry 1972). The Valley Grassland community occurs in regions characterized by a broad range of climatic conditions. Average January temperatures may range from 5°C to 15°C and July temperatures from 15°C to 30°C (NOAA 1988). Annual precipitation ranges from approximately 12 cm to over 200 cm, although all sites are characterized by a summer drought of 4-8 months (Heady 1977). Grasslands are well developed on deep, fine-textured soils although they are not restricted to such conditions (Wells 1962, Adams 1964, Heady 1977).

California

Dormant-season prescribed fires can enhance forage quality for two growing seasons in the northwestern Great Plains

Fire is an ecological disturbance that can accelerate nutrient cycling and alter herbivore distribution in grasslands and shrublands. Historically and today, humans have used prescribed fire to enhance forage quality for wild and domestic herbivores, and to achieve other management objectives. Contextual factors such as ecosystem type, seasonality, and fuel characteristics can determine how prescribed fire affects forage quality. For example, forage quality enhancements are relatively short lived in shortgrass steppe and subhumid grasslands but may persist for multiple years in northern mixed-grass prairie and sagebrush steppe. Outcomes near ecosystem boundaries are uncertain. Using 152 independent, small-plot, prescribed burns completed over multiple years at 36 sites in Wyoming and South Dakota, USA, we tested the effects of ecosystem type (northern mixed-grass prairie vs. sagebrush grassland ecotone), burn seasonality (fall vs. spring), and fuel characteristics (ambient vs. added fuel) on forage quality. Across ecosystems, fall fires had strong positive effects on forage quality parameters during the first growing season after fire, and energy benefits that persisted into the second growing season after fire. In northern mixed-grass prairie, spring and fall prescribed fires implemented before the same growing season yielded similar forage quality enhancements for one season. Finally, artificially increased fuel loads had few effects on forage quality. Although forage quality rarely dropped below critical nutrition thresholds for ruminants regardless of burning, our findings suggest relatively consistent, positive forage quality responses to prescribed fire across ecosystem types, seasonality of dormant-season burns, and fuel characteristics. Many improvements lasted through one growing season, and some persisted into a second growing season. Managers could take advantage of short-term nutritional benefits via adaptive management strategies that allow animals access to burned and unburned areas.

South Dakota, Wyoming

Science needs of southeastern grassland species of conservation concern: A framework for species status assessments

The unglaciated southeastern United States is a biodiversity hotspot, with a disproportionate amount of this biodiversity concentrated in grasslands. Like most hotspots, the Southeast is also threatened by human activities, with the total reduction of southeastern grasslands estimated as 90 percent (upwards to 100 percent for some types) and with many threats escalating today. This report summarizes the results of a multistakeholder workshop organized by the Southeastern Grasslands Initiative and the U.S. Geological Survey, held in January 2020 to provide a scientific needs assessment to help inform the Species Status Assessment (SSA) process under the U.S. Endangered Species Act, with a focus on grassland species and communities of conservation concern in the southeastern United States. This report reviews the ecology of southeastern grasslands, including influences on their origin, maintenance, and high species richness and endemism; presents findings from the workshop; and discusses science questions, hypotheses, and possibilities for future research projects to help fill key knowledge gaps. Participants in the January 2020 workshop, representing diverse expertise in various topics in southeastern grassland ecology, were tasked with identifying major threats to grassland species in the Southeast as well as potential ways to make the SSA process more efficient and effective. An underlying assumption and starting place for workshop discussion was that an ecosystem-based approach to the SSA process is more cost-efficient than a species-by-species approach, in large part because many species with similar biological requirements can be addressed by the same actions. Nevertheless, one partner in this effort, the U.S. Fish and Wildlife Service, does require specific attention be given to taxa that have been petitioned for Federal listing, though as often as possible these taxa are considered alongside a larger group of priority taxa with an ecosystem approach. For group discussions, workshop participants followed a modified “World Café” method, a structured conversational approach for knowledge sharing. Group discussions focused on five categories of threats to grassland communities and species: (1) habitat loss, fragmentation, and disruption of functional population connectivity; (2) climate change, especially changes in temperature and precipitation, including intensity and seasonality, and impacts on soil moisture, groundwater levels, and other ecosystem parameters; (3) changes to disturbance regimes, as influenced by climate and land-use change, extinctions, and human attitudes and behaviors; (4) invasive species (not limited to nonnative species); and (5) localized or subregional impacts such as sea-level rise. In addition to group discussions, workshop participants—as well as other grassland experts who were unable to attend the workshop—completed a preworkshop survey concerning challenges and opportunities for grassland conservation. Findings reported here under each of these topics represent ideas, problems, hypotheses, and questions identified by a diverse community of grassland managers and researchers which may be addressed by future research and monitoring in southeastern grassland ecosystems to help guide science-based conservation of grassland-dependent species.

Alabama, Arkansas, Florida, Georgia, Kentucky, Lou

Using conservation value to assess land restoration and management alternatives across a degraded oak savanna landscape

1. Managers considering restoration of landscapes often face a fundamental challenge - what should be the habitat composition of the restored landscape? We present a method for evaluating an important conservation trade-off inherent in making that decision. 2. Oak savannas and grasslands were historically widespread across central North America but are now rare. Today, in north-west Indiana, USA, habitats spanning a range of woody vegetation density, from nearly treeless open habitats to forests, occur across the conserved landscape where savannas probably once dominated. To understand the benefits of different potential landscape compositions, we evaluated how different proportions of five habitats - open, savanna, woodland, scrub and forest - might affect the conservation value of the north-west Indiana landscape for birds. Two variables of potential conservation importance were examined: species diversity, a measure of avian community richness, and conservation index, the percentage of a bird species' global population occurring on a hectare of landscape, summed across all bird species present. Higher values of conservation index were associated with higher local densities of globally more rare and more threatened species. 3. Conservation index and species diversity were correlated negatively across hypothetical landscapes composed of different proportions of the five habitats. Therefore, a management trade-off existed between conservation index and species diversity because landscapes that maximized species diversity differed from landscapes that maximized conservation index. 4. A landscape of 50% open, 22% savanna, 15% scrub and 13% forest was predicted to represent a compromise at which conservation index and species diversity reached the same percentage of their maxima. In contrast, the current landscape is dominated by forest. 5. Synthesis and applications. We quantified the trade-off between two potential aspects of a landscape's conservation value for birds - the landscape's ability to promote avian species diversity and the landscape's use by threatened avian species. This quantification allowed us to evaluate the ability of different landscape compositions to achieve preferable trade-off compromises, such as maximizing diversity for a given level of landscape use by threatened species. Managers can use these trade-off results to determine which landscape compositions are associated with particular conservation and management priorities.

Journal of Applied Ecology

Environmental factors that influence the location of crop agriculture in the conterminous United States

This report presents and describes high-resolution geospatial data identifying the range of environmental conditions that influence the location of cropped agricultural lands in the conterminous United States. Also presented are estimates of the extent of land where environmental constraints limit agricultural production (marginal land) and the extents of land where modifications overcome environmental constraints. The report is the result of the compilation and manipulation of datasets from numerous sources; it consists of an explanatory text and a series of appendixes and associated tables that document the data sources and data-manipulation methods in detail. Environmental factors that influence the extent of crop agriculture are terrain, climate, soil properties, and soil water. It is the combination of these four factors that allow specific crops to be grown in certain areas. Today, in order to maximize production, most of the cultivated croplands and grasslands for commercial agriculture are in areas where crops and livestock are well suited to local conditions. In the United States, cropland (row crops, closely sown crops (except hay), fruits, nuts, vegetables) occupies about 13 percent of the total land area. Grassland and rangeland occupy another 41 percent of the land area. Most crops are grown on land with shallow slope where the temperature, precipitation, and soils are favorable. In areas that are too steep, wet, or dry, landscapes have been modified to allow cultivation. Some of the limitations of the environmental factors that determine the location of agriculture can be overcome through modifications, but others cannot. On a larger-than-field scale, agricultural modifications commonly influence water availability through irrigation and (or) drainage and soil fertility and (or) organic-matter content through amendments such as manure, commercial fertilizer and lime. In general, it is not feasible to modify the other environmental factors, soil texture, soil depth, soil mineralogy, temperature, and terrain at large scales.

Scientific Investigations Report

Evolution of CAM and C 4 carbon-concentrating mechanisms

Mechanisms for concentrating carbon around the Rubisco enzyme, which drives the carbon-reducing steps in photosynthesis, are widespread in plants; in vascular plants they are known as crassulacean acid metabolism (CAM) and C 4 photosynthesis. CAM is common in desert succulents, tropical epiphytes, and aquatic plants and is characterized by nighttime fixation of CO2. The proximal selective factor driving the evolution of this CO2-concentrating pathway is low daytime CO2, which results from the unusual reverse stomatal behavior of terrestrial CAM species or from patterns of ambient CO2 availability for aquatic CAM species. In terrestrials the ultimate selective factor is water stress that has selected for increased water use efficiency. In aquatics the ultimate selective factor is diel fluctuations in CO2 availability for palustrine species and extreme oligotrophic conditions for lacustrine species. C4 photosynthesis is based on similar biochemistry but carboxylation steps are spatially separated in the leaf rather than temporally as in CAM. This biochemical pathway is most commonly associated with a specialized leaf anatomy known as Kranz anatomy; however, there are exceptions. The ultimate selective factor driving the evolution of this pathway is excessively high photorespiration that inhibits normal C3 photosynthesis under high light and high temperature in both terrestrial and aquatic habitats. CAM is an ancient pathway that likely has been present since the Paleozoic era in aquatic species from shallow-water palustrine habitats. While atmospheric CO2 levels have undoubtedly affected the evolution of terrestrial plant carbon-concentrating mechanisms, there is reason to believe that past atmospheric changes have not played as important a selective role in the aquatic milieu since palustrine habitats today are not generally carbon sinks, and the selective factors driving aquatic CAM are autogenic. Terrestrial CAM, in contrast, is of increasing selective value under extreme water deficits, and undoubtedly, high Mesozoic CO2 levels reduced the amount of landscape perceived by plants as water limited. Late Tertiary and Quaternary reductions in atmospheric CO2, coupled with increasing seasonality, were probably times of substantial species radiation and ecological expansion for CAM plants. C4 photosynthesis occurs in only about half as many families as CAM, and three-fourths of C4 species are either grasses or sedges. Molecular phylogenies indicate C4 is a more recent innovation than CAM and that it originated in the mid-Tertiary, 20–30 Ma, although some data support an earlier origin. While the timing of the origin of C4 remains controversial, the nearly explosive increase in C4 species is clearly documented in the late Miocene, 4–7 Ma. Increasing seasonality has been widely suggested as an important climatic stimulus for this C4 expansion. Alternatively, based on models of photosynthetic quantum yield at different temperatures and CO2 concentration, it has been hypothesized that the late Miocene C4 expansion resulted from declining atmospheric CO2 levels. This model is most appropriate for explaining the transition from C3 grasslands to C4 grasslands but by itself may not be sufficient to explain the more likely scenario of a late Miocene transition from C3 woodland/ savanna to C4 grasslands. A largely unexplored hypothesis is that climatic changes in late Miocene altered disturbance regimes, in particular the incidence of fires, which today are often associated with maintenance of C4 grasslands. Oceanic charcoal sediments that appear to represent Aeolian deposits from continental wildfires follow a strikingly similar pattern of explosive increase in late Miocene. Climate, CO2, and disturbance are not mutually exclusive explanations and probably all acted in concert to promote the expansion of C4 grasslands. More recently, late Quaternary changes in CO2 may have been responsible for driving major changes in the landscape distribution of C4 species. The theory is sound; however, many of the studies cited in support of this model are open to alternative interpretations, and none has eliminated climatic factors as important selective agents. CAM and C4 evolution required coupling of biochemical pathways with structural changes in photosynthetic tissues, succulence in CAM and Kranz in C4. This was apparently accomplished by piecemeal evolution beginning with mechanisms for recapturing respiratory CO2, although this need not have been so in aquatic CAM species. It has been proposed that the extreme rarity of both pathways in the same plant results from biochemical and structural incompatibilities (Sage 2002). Equally important is the fact that the selective environments are quite different, with CAM evolution thriving on stressful sites inhospitable to C3 species whereas C4 evolution has selected for rapid growth capable of outcompeting associated C3 plants.

International Journal of Plant Sciences

Changes in conservation value from grasslands to savannas to forests: How a temperate canopy cover gradient affects butterfly community composition

Temperate savannas and grasslands are globally threatened. In the Midwest United States of America (USA), for example, oak savannas persist today at a small percentage of recent historic coverage. Therefore, restoration of habitats of low and intermediate canopy cover is a landscape conservation priority that often emphasizes returning tree density to a savanna-like target value. Understanding how animal species react to such changes in vegetation structure is important for assessing the value of these restoration plans. We examined how butterfly community attributes in northwest Indiana USA, including community composition, richness, and abundance responded to a grassland-to-forest gradient of canopy cover. Butterfly community composition under intermediate canopy cover differed significantly from community composition in the most open or closed-canopy habitats. Composition of the plant community in flower was a significant predictor of three assessed attributes of the butterfly community—composition, richness, and abundance. Phenology, expressed as day-of-the-year, was also a strong predictor of these butterfly community attributes. Few butterfly species were habitat specialists as adults although canopy cover was a more important predictor of adult community composition than of richness or abundance of butterflies. Therefore, adult butterfly community differences along the canopy cover gradient were less about butterfly communities filled with habitat specialists for different canopy-defined habitats and more about gradual changes in community composition along this gradient. Overall, butterfly community richness was predicted to peak at about 34% canopy cover, butterfly abundance at about 53% canopy cover, community conservation value at about 59% canopy cover, and a combination of desirable conservation attributes–high diversity, high abundance, and high conservation value–was predicted to reach a peak of co-occurrence at about 67% canopy cover suggesting that habitats of intermediate canopy cover might be particularly effective for butterfly conservation in this region.

Indiana

Mapping grasslands suitable for cellulosic biofuels in the Greater Platte River Basin, United States

Biofuels are an important component in the development of alternative energy supplies, which is needed to achieve national energy independence and security in the United States. The most common biofuel product today in the United States is corn-based ethanol; however, its development is limited because of concerns about global food shortages, livestock and food price increases, and water demand increases for irrigation and ethanol production. Corn-based ethanol also potentially contributes to soil erosion, and pesticides and fertilizers affect water quality. Studies indicate that future potential production of cellulosic ethanol is likely to be much greater than grain- or starch-based ethanol. As a result, economics and policy incentives could, in the near future, encourage expansion of cellulosic biofuels production from grasses, forest woody biomass, and agricultural and municipal wastes. If production expands, cultivation of cellulosic feedstock crops, such as switchgrass ( Panicum virgatum L. ) and miscanthus ( Miscanthus species ), is expected to increase dramatically. The main objective of this study is to identify grasslands in the Great Plains that are potentially suitable for cellulosic feedstock (such as switchgrass) production. Producing ethanol from noncropland holdings (such as grassland) will minimize the effects of biofuel developments on global food supplies. Our pilot study area is the Greater Platte River Basin, which includes a broad range of plant productivity from semiarid grasslands in the west to the fertile corn belt in the east. The Greater Platte River Basin was the subject of related U.S. Geological Survey (USGS) integrated research projects.

Colorado;Nebraska;South Dakota;Wyoming

The Pothole Hydrology-Linked Systems Simulator (PHyLiSS)—Development and application of a systems model for prairie-pothole wetlands

The North American Prairie Pothole Region covers about 770,000 square kilometers of the United States and Canada (including parts of 5 States and 3 provinces: North Dakota, South Dakota, Montana, Minnesota, Iowa, Saskatchewan, Manitoba, and Alberta). The Laurentide Ice Sheet shaped the landscape of the region about 12,000 to 14,000 years ago. The retreat of the ice sheet left behind low-permeability glacial till and a landscape dotted with millions of depressions known today as prairie potholes. The wetlands that subsequently formed in these depressions, prairie-pothole wetlands, provide critical migratory-bird habitat and support dynamic aquatic communities. Extensive grasslands and productive agricultural systems surround these wetland ecosystems. In prairie-pothole wetlands, the compositions of plant, invertebrate, and vertebrate communities are highly dependent on hydrogeochemical conditions. Regional climate shifts between wet and dry periods affect the length of time that wetlands contain ponded surface water and the chemistry of that ponded water. Land-use change can exacerbate or reduce the effects of climate on wetland hydrology and water chemistry. A mechanistic understanding of the relation among climate, land use, hydrology, chemistry, and biota in prairie-pothole wetlands is needed to better understand the complex, and often interacting, effects of climate and land use on prairie-pothole wetland systems and to facilitate climate and land-use change adaptation efforts. The Pothole Hydrology-Linked Systems Simulator (PHyLiSS) model was developed to address this need. The model simulates water-surface elevation dynamics in prairie-pothole wetlands and quantifies changes in salinity. The PHyLiSS model is unique among other wetland models because it accommodates differing sizes and morphometries of wetland basins, is not dependent on a priori designations of wetland class, and allows for functional changes associated with dynamic shifts in ecohydrological states. The PHyLiSS model also has the capability to simulate wetland salinity, and potential future iterations will also simulate the effects of changing hydrology and geochemical conditions on biota. This report documents the development of the hydrological and geochemical components of the PHyLiSS model and provides example applications.

Open-File Report

Herbicide treatment of invasive Vinca major growing with endangered Galium buxifolium, an island endemic

Galium buxifolium E. Greene [Rubiaceae] (sea-cliff bedstraw) is a small shrub restricted to San Miguel and Santa Cruz Islands, in the California Channel Islands. Almost all of the 26 known populations grow on vertical north-facing sea cliffs in native scrub, sandwiched between the sea below and non-native annual grasslands on the terraces above. A notable exception is a popula?tion at Pelican Bay on Santa Cruz Island, growing on the cliff and on thin terrace soils above the cliff in a stand of coastal bluff scrub that is recovering from more than a century of sheep grazing (Figure 1). Ironically, this stand is near the location of the historic Eaton Resort, a charismatic inn frequented by Hollywood glitterati in the early 1900s. Several landscape ornamentals planted there persist today in the area of the Galium population. Italian stone pine (Pinus pinaea) and eucalyptus (Eucalyptus spp.) tower over the site while the invasive groundcover Vinca major L. [Apocynaceae] (greater periwinkle) forms dense mats at the cliff edge that are spreading into the developing native plant stand. Wherever the Galium is found it is intermixed with other native scrub plants in dense communities on moist ocean bluffs. Historic notes and herbarium collections indicate that it may have been more widespread on nearshore terraces adjacent to sea-cliff populations before conversion to grassland. Since Pelican Bay is the only site that we know of where Galium is spreading onto the nearby terrace, we wanted to know whether the Vinca posed a roadblock to upslope population expansion. In 2005 we mapped Vinca and Galium at the site. We also measured Galium individuals to see where the smaller, younger plants were to better understand where the population is expanding. We observed that 1) both the Galium and the Vinca appear to be spreading from the cliff face upslope onto a series of rock outcrops, stone walls and benches, 2) the native scrub community is recovering at the site and 3) the Vinca appears to be moving into the native scrub where it overtops small plants, including those Galium in the smallest size classes. This observed pattern of Vinca displacement of native vegetation has been noted in other places, where it is treated as an invasive weed. We concluded that Vinca may pose a threat to the expansion of both the native scrub and the Galium population that it supports. Therefore, we worked with the US Fish and Wildlife Service and others to develop a research project investigating 1) best techniques for killing Vinca within the boundaries of an endangered plant population and 2) demographic response of Galium to the treatment. Our intent is to push Vinca back to the vertical cliff face to give the natives a chance to establish a vigorous stand. Our conservation goal is to encourage natural establishment of new Galium plants on the terrace along with expansion of the native coastal bluff scrub and Galium population. Our immediate treatment objective is to reduce live Vinca cover by 90 % on the accessible upslope portions of the habitat. Our recovery objective is no net loss of Galium plants 2006-2016.

California

Late Quaternary environmental change inferred from phytoliths and other soil-related proxies: Case studies from the central and southern Great Plains, USA

This study investigates stable carbon isotopes ( δ 13 C), opal phytolith assemblages, burnt phytoliths, microscopic charcoal and Sporormiella spores from modern soils and paleosols in Kansas and Oklahoma. Grass and dicot phytoliths in combination with δ 13 C are used as proxies for reconstructing the structure of grasslands and woodlands. Burnt grass phytoliths and microscopic charcoal are evaluated as proxies for reconstructing paleofire incidence. Concentrations of the fungal spore Sporormiella are used as a proxy for assessing large herbivore activity. These proxies were tested on various modern grassland communities of the central and southern Great Plains, including areas with bison, cattle, and small herbivores, and areas under different fire frequencies. Opal phytolith assemblages and δ 13 C values show that before cal 11 ka, C 3 grasses and woody plants predominated in areas that today are dominated by C 4 grasses. The origin of the shortgrass prairie dates back to about cal 10 ka. The origin of the tallgrass prairie, however, is not clear as phytolith data show variable assemblages throughout the Holocene (mixed-grass, tallgrass, and tallgrass–woodland mosaic). Different proxies (burnt phytoliths vs. charcoal) reveal different fire frequencies, but it is apparent that microfossil evidence for fire incidence is closely related to the abundance of woody plants in the landscape. Before cal 12 ka, soils show somewhat elevated concentration of Sporormiella , but lower concentrations than the modern high-density bison and cattle grazing areas. Throughout the Holocene, Sporormiella frequencies are low, which suggests lower large ungulate densities and perhaps high mobility.

Colorado, Kansas, New Mexico, Oklahoma, Texas

Middle Pliocene vegetation: Reconstructions, paleoclimatic inferences, and boundary conditions for climate modeling

The general characteristics of global vegetation during the middle Pliocene warm period can be reconstructed from fossil pollen and plant megafossil data. The largest differences between Pliocene vegetation and that of today occurred at high latitudes in both hemispheres, where warming was pronounced relative to today. In the Northern Hemisphere coniferous forests lived in the modern tundra and polar desert regions, whereas in the Southern Hemisphere southern beech apparently grew in coastal areas of Antarctica. Pliocene middle latitude vegetation differed less, although moister-than-modern conditions supported forest and woodland growth in some regions now covered by steppe or grassland. Pliocene tropical vegetation reflects essentially modem conditions in some regions and slightly cooler-than-or warmer-than- modern climates in other areas. Changes in topography induced by tectonics may be responsible for many of the climatic changes since the Pliocene in both middle and lower latitudes. However, the overall latitudinal progression of climatic conditions on land parallels that seen in the reconstruction of middle Pliocene sea-surface temperatures. Pliocene paleovegetational data was employed to construct a 2 ° × 2 ° global grid of estimated mid-Pliocene vegetational cover for use as boundary conditions for numerical General Circulation Model simulations of middle Pliocene climates. Continental outlines and topography were first modified to represent the Pliocene landscape on the 2 ° × 2 ° grid. A modern 1 ° × 1 ° vegetation grid was simplified and mapped on this Pliocene grid, and then modified following general geographic trends evident in the Pliocene paleovegetation data set.

Marine Micropaleontology

Second chance for the plains bison

Before European settlement the plains bison ( Bison bison bison ) numbered in the tens of millions across most of the temperate region of North America. Within the span of a few decades during the mid- to late-1800s its numbers were reduced by hunting and other factors to a few hundred. The plight of the plains bison led to one of the first major movements in North America to save an endangered species. A few individuals and the American Bison Society rescued the remaining animals. Attempts to hybridize cattle and bison when bison numbers were low resulted in extensive cattle gene introgression in bison. Today, though approximately 500,000 plains bison exist in North America, few are free of cattle gene introgression, 96% are subject to anthropogenic selection for commodity production, and only 4% are in herds managed primarily for conservation purposes. Small herd size, artificial selection, cattle-gene introgression, and other factors threaten the diversity and integrity of the bison genome. In addition, the bison is for all practical purposes ecologically extinct across its former range, with multiple consequences for grassland biodiversity. Urgent measures are needed to conserve the wild bison genome and to restore the ecological role of bison in grassland ecosystems. Socioeconomic trends in the Great Plains, combined with new information about bison conservation needs and new conservation initiatives by both the public and public sectors, have set the stage for significant progress in bison conservation over the next few years.

Biological Conservation

A 16,000 14C yr B.P. packrat midden series from the USA-Mexico Borderlands

A new packrat midden chronology from Playas Valley, southwestern New Mexico, is the first installment of an ongoing effort to reconstruct paleovegetation and paleoclimate in the U.S.A.-Mexico Borderlands. Playas Valley and neighboring basins supported pluvial lakes during full and/or late glacial times. Plant macrofossil and pollen assemblages from nine middens in the Playas Valley allow comparisons of two time intervals: 16,000-10,000 and 4000-0 14C yr B.P. Vegetation along pluvial lake margins consisted of open pinyon-juniper communities dominated by Pinus edulis, Juniperus scopulorum, Juniperus cf. coahuilensis, and a rich understory of C4 annuals and grasses. This summer-flowering understory is also characteristic of modern desert grassland in the Borderlands and indicates at least moderate summer precipitation. P. edulis and J. scopulorum disappeared or were rare in the midden record by 10,670 14C yr B.P. The late Holocene is marked by the arrival of Chihuahuan desert scrub elements and few departures as the vegetation gradually became modern in character. Larrea tridentata appears as late as 2190 14C yr B.P. based on macrofossils, but may have been present as early as 4095 14C yr B.P. based on pollen. Fouquieria splendens, one of the dominant desert species present at the site today, makes its first appearance only in the last millennium. The midden pollen assemblages are difficult to interpret; they lack modern analogs in surface pollen assemblages from stock tanks at different elevations in the Borderlands. ?? 2003 University of Washington. Published by Elsevier Inc. All rights reserved.

Quaternary Research

Wyoming and Landsat

Wyoming has the smallest population of any State—fewer than 600,000 people—but an abun­dance of wildlife. The largest number of pronghorn ( Antilocapra americana ), often called antelope, and the biggest public bison ( Bison bison ) herd in the United States live in Wyoming, which also hosts elk ( Cervus elaphus ), moose ( Alces americanus ), bighorn sheep ( Ovis canadensis ), black bears ( Ursus americanus ), and grizzly bears ( U. arctos ). The terrain of the Nation’s 10th largest State varies from the Black Hills to the Rocky Mountains, and from Great Plains grassland to Wyoming Basin desert sagebrush. Natural resources attract millions of visitors annually, especially to the country’s first national park, Yellowstone, and the ski slopes in Grand Teton National Park and elsewhere. Natural resources account for much of the employment in the Cowboy State, from tourism and ranching to coal, natural gas, and oil mining. To help monitor these natural resources, researchers and land managers have relied on USGS Landsat imagery and data. The scale of images from the first Landsat satellites helped geologists detect previously unknown uranium deposits in Wyoming, which is the leading State for uranium production. Today, land managers and researchers take advantage of Landsat and its 50-year archive to explore landscape change over time—and continue to make new discoveries.

Wyoming

Late Quaternary environmental change in eastern Beringia

Eastern Beringia (Alaska and western Yukon) is an extensive, high-latitude region of North America that remained largely unglaciated throughout the Quaternary. Consequently, its sedimentary deposits preserve long-term environmental records that have intrigued scientists for nearly a century. Recent advances in palaeoecological proxies and dating methods have proved critical in addressing long-standing questions about regional late Quaternary environmental change. At the same time, they have led to new and sometimes controversial hypotheses. This review covers recent discoveries and unresolved questions focused on the period 57,000–10,000 calendar years before C.E. 1950 (cal yr BP). The middle Wisconsin interstadial (57,000–30,000 cal yr BP) was a period of relative warmth in eastern Beringia, compared with the late Wisconsin (30,000–14,000 cal yr BP). Early in the interstadial occasional Picea woodland was present amongst widespread shrub tundra. Palaeoecological, sedimentary and isotopic data indicate that climate was cooler and drier than the Holocene, with high rates of aeolian activity. Megafauna typically associated with the ‘mammoth steppe’ ecosystem (woolly mammoth [ Mammuthus primigenius ], horse [ Equus ] and steppe-bison [ Bison priscus ]) were present in some abundance. The transition towards late Wisconsin cold-stage conditions (35,000–30,000 cal yr BP) coincided with the establishment of the Bering Land Bridge and featured expansion of spatially varied, herbaceous vegetation, sometimes associated with deep active layers. Sedimentary DNA ( seda DNA) and macrofossil evidence show vegetation was not a prairie-like grassland, and the term “steppe-tundra” is a better descriptor. Permafrost pore-ice isotopic (δ 18 O) records suggest a step change in one or more climate drivers ca. 30,000 cal yr BP, by which time steppe-tundra was established across eastern Beringia. It remains uncertain whether Picea survived cold-stage conditions within isolated refugia, or whether it recolonized from south of the Laurentide-Cordilleran ice sheets. Genetic data suggest that Picea probably survived in situ ; however, there is no definitive fossil evidence to support this. The end-Pleistocene transition from steppe-tundra to shrub tundra began ca. 15,000 cal yr BP and took place within decades at local scales. The expansion of woody taxa coincided with rising sea levels, reduced sea-ice extent and an abrupt shift in atmospheric circulation that enhanced precipitation. During this time, Earth's orbital configuration caused high early-summer temperatures and strong seasonality, creating growing conditions very different from today. The vegetation consisted of Salix and Betula shrub tundra with open areas of herbs and graminoids. During the deglacial warming trend, the Younger Dryas oscillation (12,800–11,700 cal yr BP) was variably expressed. It is generally evident in records affected by adjacent oceans but can be absent at sites in continental areas. These past conditions and paleoenvironmental changes have implications for contemporary issues: hypotheses about Pleistocene mammalian extinction; sensitivity of eastern Beringia to major oceanic reorganizations and high-frequency climate variability; the nature of woody plant expansion with climate warming; grazing, hydroclimate and fire as controls over ecosystems; the efficacy of “Pleistocene rewilding” for carbon capture.

Beringia