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Significance of headwater streams and perennial springs in ecological monitoring in Shenandoah National Park

Shenandoah National Park has been monitoring water chemistry and benthic macroinvertebrates in stream ecosystems since 1979. These monitoring efforts were designed to assess the status and trends in stream condition associated with atmospheric deposition (acid rain) and changes in forest health due to gypsy moth infestations. The primary objective of the present research was to determine whether the current long-term macroinvertebrate and water-quality monitoring program in Shenandoah National Park was failing to capture important information on the status and trends in stream condition by not sufficiently representing smaller, headwater streams. The current benthic-macroinvertebrate and water-chemistry sampling designs do not include routine collection of data from streams with contributing watershed areas smaller than 100 hectares, even though these small streams represent the overwhelming proportion of total stream length in the park. In this study, we sampled headwater sites, including headwater stream reaches (contributing watershed area approximately 100 hectares (ha) and perennial springs, in the park for aquatic macroinvertebrates and water chemistry and compared the results with current and historical data collected at long-term ecological monitoring (LTEM) sites on larger streams routinely sampled as part of ongoing monitoring efforts. The larger purpose of the study was to inform ongoing efforts by park managers to evaluate the effectiveness and efficiency of the current aquatic monitoring program in light of other potential stressors (for example, climate change) and limited resources. Our results revealed several important findings that could influence management decisions regarding long-term monitoring of park streams. First, we found that biological indicators of stream condition at headwater sites and perennial springs generally were more indicative of lower habitat quality and were more spatially variable than those observed at sites on routinely monitored larger streams. We hypothesized that poorer stream condition observed in smaller streams was due to stream drying that occurs more frequently in headwater areas. We also found that biological and water-chemistry measures responded differently to landscape drivers. Variation in most biological endpoints was driven primarily by stream size and was only secondarily associated with bedrock geology. In contrast, water chemistry showed essentially the opposite pattern, with underlying geology explaining much of the variation and stream size being of secondary importance. Therefore, expanding the LTEM program to include headwater areas would yield substantially different biological information, whereas broad inferences regarding spatial patterns in water chemistry would probably not change. Although significant differences in community composition were observed among streams of different sizes, no taxa were unique to headwater sites. All taxa collected at the 45 headwater sites also had been collected at one or more LTEM sites during one or more years. This observation indicates that headwater sites in the park may be structured by biotic nestedness; consequently, focusing management efforts on preserving the species pool at the larger LTEM sites would likely result in the protection of most taxa parkwide. Finally, linkages (correlations) between water chemistry and biological measures of stream condition were signficantly stronger when assessed at the LTEM sites than when assessed at the springs or headwater sites, indicating that conditions at downstream sites may be better indicators of water-quality trends.

Virginia↗

Modelling dendritic ecological networks in space: anintegrated network perspective

Dendritic ecological networks (DENs) are a unique form of ecological networks that exhibit a dendritic network topology (e.g. stream and cave networks or plant architecture). DENs have a dual spatial representation; as points within the network and as points in geographical space. Consequently, some analytical methods used to quantify relationships in other types of ecological networks, or in 2-D space, may be inadequate for studying the influence of structure and connectivity on ecological processes within DENs. We propose a conceptual taxonomy of network analysis methods that account for DEN characteristics to varying degrees and provide a synthesis of the different approaches within the context of stream ecology. Within this context, we summarise the key innovations of a new family of spatial statistical models that describe spatial relationships in DENs. Finally, we discuss how different network analyses may be combined to address more complex and novel research questions. While our main focus is streams, the taxonomy of network analyses is also relevant anywhere spatial patterns in both network and 2-D space can be used to explore the influence of multi-scale processes on biota and their habitat (e.g. plant morphology and pest infestation, or preferential migration along stream or road corridors).

Ecology Letters↗

Strategic conservation planning for the Eastern North Carolina/Southeastern Virginia Strategic Habitat Conservation Team

The Eastern North Carolina/Southeastern Virginia Strategic Habitat Conservation Team (ENCSEVA) is a partnership among local federal agencies and programs with a mission to apply Strategic Habitat Conservation to accomplish priority landscape-level conservation within its geographic region. ENCSEVA seeks to further landscape-scale conservation through collaboration with local partners. To accomplish this mission, ENCSEVA is developing a comprehensive Strategic Habitat Conservation Plan (Plan) to provide guidance for its members, partners, and collaborators by establishing mutual conservation goals, objectives, strategies, and metrics to gauge the success of conservation efforts. Identifying common goals allows the ENCSEVA team to develop strategies that leverage joint resources and are more likely to achieve desired impacts across the landscape. The Plan will also provide an approach for ENCSEVA to meet applied research needs (identify knowledge gaps), foster adaptive management principles, identify conservation priorities, prioritize threats (including potential impacts of climate change), and identify the required capacity to implement strategies to create more resilient landscapes. ENCSEVA seeks to support the overarching goals of the South Atlantic Landscape Conservation Cooperative (SALCC) and to provide scientific and technical support for conservation at landscape scales as well as inform the management of natural resources in response to shifts in climate, habitat fragmentation and loss, and other landscape-level challenges (South Atlantic LCC 2012). The ENCSEVA ecoregion encompasses the northern third of the SALCC geography and offers a unique opportunity to apply landscape conservation at multiple scales through the guidance of local conservation and natural resource management efforts and by reporting metrics that reflect the effectiveness of those efforts (Figure 1). The Environmental Decision Analysis Team, housed within the North Carolina Cooperative Fish and Wildlife Research Unit at North Carolina State University, is assisting the ENCSEVA team in developing a scientifically sound basis for the Plan though the elicitation of expert knowledge and the organization of that knowledge using the Open Standards for the Practice of Conservation. The Open Standards for the Practice of Conservation is a framework that is well suited to incorporating decision-making tools such as Structured Decision Making and provides a multi-step process to conceptually organize conservation projects in a manner that enhances the rigor and transparency of expert and knowledge-based plans. It helps define explicit pathways from 2 planned conservation activities and ultimate impact, as well as indicators to measure success (Stem et al. 2005). Specifically, the framework identifies conservation targets, key ecological attributes, threats, and associated indicators to monitor responses given the implementation of a conservation action (Conservation Measures Partnership 2007). This report serves to provide a scientific foundation for the Plan by summarizing the expert opinion of wildlife biologists, ecologists, hydrologists, researchers, natural resource managers, and conservation practitioners regarding five environments (wetlands, riverine systems, estuaries, uplands, and barrier islands) within the ENCSEVA geography. Specifically, this report describes (1) the approach to elicit expert knowledge meant to support the strategic plan, (2) how this knowledge can inform collaborative conservation planning, and (3) a summary of opportunities available for the ENCSEVA team to address threats and impacts associated with climate change within the ecoregion.

Technical Bulletin↗

Integrating scientific knowledge into large-scale restoration programs: the CALFED Bay-Delta Program experience

Integrating science into resource management activities is a goal of the CALFED Bay-Delta Program, a multi-agency effort to address water supply reliability, ecological condition, drinking water quality, and levees in the Sacramento-San Joaquin Delta of northern California. Under CALFED, many different strategies were used to integrate science, including interaction between the research and management communities, public dialogues about scientific work, and peer review. This paper explores ways science was (and was not) integrated into CALFED's management actions and decision systems through three narratives describing different patterns of scientific integration and application in CALFED. Though a collaborative process and certain organizational conditions may be necessary for developing new understandings of the system of interest, we find that those factors are not sufficient for translating that knowledge into management actions and decision systems. We suggest that the application of knowledge may be facilitated or hindered by (1) differences in the objectives, approaches, and cultures of scientists operating in the research community and those operating in the management community and (2) other factors external to the collaborative process and organization.

Environmental Science and Policy↗

Mapping fire regime ecoregions in California

The fire regime is a central framing concept in wildfire science and ecology and describes how a range of wildfire characteristics vary geographically over time. Understanding and mapping fire regimes is important for guiding appropriate management and risk reduction strategies and for informing research on drivers of global change and altered fire patterns. Most efforts to spatially delineate fire regimes have been conducted by identifying natural groupings of fire parameters based on available historical fire data. This can result in classes with similar fire characteristics but wide differences in ecosystem types. We took a different approach and defined fire regime ecoregions for California to better align with ecosystem types, without using fire as part of the definition. We used an unsupervised classification algorithm to segregate the state into spatial clusters based on distinctive biophysical and anthropogenic attributes that drive fire regimes – and then used historical fire data to evaluate the ecoregions. The fire regime ecoregion map corresponded well with the major land cover types of the state and provided clear separation of historical patterns in fire frequency and size, with lower variability in fire severity. This methodology could be used for mapping fire regimes in other regions with limited historical fire data or forecasting future fire regimes based on expected changes in biophysical characteristics.

California↗

Evolutionary fire ecology: An historical account and future directions

The idea that fire acts as an evolutionary force contributing to shaping species traits started a century ago, but had not been widely recognized until very recently. Among the first to realize this force were E dward B . Poulton, R. D ale Guthrie, and E dwin V . Komarek in animals and W illis L . Jepson, W alter W . Hough, T om M . Harris, P hilip V . Wells, and R obert W . Mutch in plants. They were all ahead of their time in their evolutionary thinking. Since then, evolutionary fire ecology has percolated very slowly into the mainstream ecology and evolutionary biology; in fact, this topic is still seldom mentioned in textbooks of ecology or evolution. Currently, there is plenty of evidence suggesting that we cannot understand the biodiversity of our planet without considering the key evolutionary role of fire. But there is still research to be done in order to fully understand fire's contribution to species evolution and to predicting species responses to rapid global changes.

BioScience↗

Growth patterns of invasive Silver Carp in the Mississippi River basin

Silver Carp ( Hypophthalmichthys molitrix ) are an invasive fish in the Mississippi River Basin. Their rapid expansion over recent decades coupled with extraordinary growth rates have arguably caught many by surprise. Understanding the atypical growth rates that could be the driving force behind the Silver Carp's explosive expansion may be crucial for development of management strategies. Towards this goal, I synthetized existing data on the growth and maturity patterns of Silver Carp. I compiled 62 estimates of growth representing populations of native Silver Carp in East Asia and invasive Silver Carp in the Mississippi River Basin. A remarkably rapid increase in length at earlier ages, undocumented in their ancestral range, was a defining feature of Silver Carp in the Mississippi River Basin. Some of the fastest growth rates of Silver Carp were recorded in hypereutrophic floodplain lakes and at lower latitudes. Invasive Silver Carp frequently deviated from the growth patterns established by native species as evidenced by abnormally high growth coefficients ( K ) in relation to asymptotic length ( L ∞ ). There is evidence of genetic differentiation between native Silver Carp and those expanding in the Mississippi River Basin possibly resulting from genetic background of introductions, genetic drift, and ecological selection. There is also limited evidence of enemy release, allowing for reallocation of energy from defenses to the rapid growth; though speculative, this is a plausible hypothesis that merits further research. This overview of the growth patterns of invasive Silver Carp underscores the need for novel strategies to mitigate the rapid generation time induced by the atypical growth patterns.

Mississippi River basin↗

Development of high-throughput genomic resources to inform white-tailed deer population and disease management

White-tailed deer ( Odocoileus virginianus ) are the most abundant and widespread cervid in North America. Genetic data are used as a tool to monitor populations and make management decisions for this game species. However, the development and use of genomic tools that can generate a set of markers suitable for longitudinal genomic data collection, whether for management purposes or to study the demographic and evolutionary processes of widely distributed species, have been challenging. This is mainly due to the cost required to fully implement and interpret the data produced. Here, we generated whole genome resequencing data for 44 free-ranging deer from three regions in their central and eastern North American range and identified over 89 million single nucleotide polymorphisms (SNPs). We used a subset of these SNPs to develop two nested SNP tools, a high-density array (702,183 SNPs) and a medium-density array (72,723 SNPs) to support deer and chronic wasting disease (CWD) management and research. SNPs were selected to ensure an even distribution across scaffolds of the reference genome and include SNPs associated with CWD susceptibility. Using genotyping results for 469 deer from 15 states in the US and Mexico generated by the high-density array and 1335 deer from 18 states generated by the medium-density array, we assessed genotyping success across different populations and explored some insights into population structure. These genomic tools offer a standard set of markers that will enable researchers and managers to address important questions related to white-tailed deer and CWD management. Our SNP arrays also offer the opportunity to examine aspects of white-tailed deer ecology and evolutionary history that were previously difficult to address.

Molecular Ecology Resources↗

Managing climate change refugia for climate adaptation

Refugia have long been studied from paleontological and biogeographical perspectives to understand how populations persisted during past periods of unfavorable climate. Recently, researchers have applied the idea to contemporary landscapes to identify climate change refugia, here defined as areas relatively buffered from contemporary climate change over time that enable persistence of valued physical, ecological, and socio-cultural resources. We differentiate historical and contemporary views, and characterize physical and ecological processes that create and maintain climate change refugia. We then delineate how refugia can fit into existing decision support frameworks for climate adaptation and describe seven steps for managing them. Finally, we identify challenges and opportunities for operationalizing the concept of climate change refugia. Managing climate change refugia can be an important option for conservation in the face of ongoing climate change.

PLoS ONE↗

Columbia Environmental Research Center

The U.S. Geological Survey Columbia Environmental Research Center performs research to solve challenging environmental problems related to contaminants and habitat alterations in aquatic and terrestrial ecosystems. The research is interdisciplinary and pursued through partnerships within the U.S. Geological Survey and with national, international, state, and local agencies; nongovernmental organizations; and universities. Research is prioritized to provide science to the U.S. Department of the Interior and other natural resource management agencies to inform rehabilitation of degraded habitats and imperiled fish and wildlife populations. The Columbia Environmental Research Center was established in 1966 in Columbia, Missouri, as the U.S. Fish and Wildlife Service’s Fish Pesticide Research Laboratory; the Columbia Environmental Research Center was incorporated into the U.S. Geological Survey in 1996. The U.S. Geological Survey’s staff of 130 includes 90 scientists of which one-half have advanced degrees in ecology, toxicology, biology, biochemistry, chemistry, hydrology, geology, and information technology.

Missouri↗

Road impacts on the Baca National Wildlife Refuge, Colorado, with emphasis on effects to surface- and shallow ground-water hydrology - A literature review

A review of published research on unpaved road effects on surface-water and shallow ground-water hydrology was undertaken to assist the Baca National Wildlife Refuge, Colorado, in understanding factors potentially influencing refuge ecology. Few studies were found that addressed hydrological effects of roads on a comparable area of shallow slope in a semiarid region. No study dealt with road effects on surface- and ground-water supplies to ephemeral wetlands, which on the refuge are sustained by seasonal snowmelt in neighboring mountains. Road surfaces increase runoff, reduce infiltration, and serve as a sediment source. Roadbeds can interfere with normal surface- and ground-water flows and thereby influence the quantity, timing, and duration of water movement both across landscapes and through the soil. Hydrologic effects can be localized near the road as well as widespread and distant. The number, arrangement, and effectiveness of road-drainage structures (culverts and other devices) largely determine the level of hydrologic alteration produced by a road. Undesirable changes to natural hydrologic patterns can be minimized by considering potential impacts during road design, construction, and maintenance. Road removal as a means to restore desirable hydrologic conditions to landscapes adversely affected by roads has yet to be rigorously evaluated.

Colorado↗

Federal collaboration in science for invasive mammal management in U.S. National Parks and Wildlife Refuges of the Pacific Islands

Some of the most isolated islands in the Pacific Ocean are home to US National Parks and Wildlife Refuges. These islands are known for flora and fauna that occur nowhere else, but also for invasive species and other factors which have resulted in the disproportionate extinction of native species. The control of invasive mammals is the single most expensive natural resource management activity essential for restoring ecological integrity to parks in the Hawaiian Islands, American Samoa, and the islands of Guam and Saipan. Science-based applications supporting management efforts have been shaped by longstanding collaborative federal research programs over the past four decades. Consequently, feral goats (Capra hircus) have been removed from >690 km2 in National Parks, and feral pigs (Sus scrofa) have been removed from >367 km2 of federal lands of Hawai‘i, bringing about the gradual recovery of forest ecosystems. The exclusion of other non-native ungulates and invasive mammals is now being undertaken with more sophisticated control techniques and fences. New fence designs are now capable of excluding feral cats (Felis catus) from large areas to protect endangered native waterfowl and nesting seabirds. Rodenticides which have been tested and registered for hand and aerial broadcast in Hawai‘i have been used to eradicate rats from small offshore islands to protect nesting seabirds and are now being applied to montane environments of larger islands to protect forest birds. Forward-looking infrared radar (FLIR) is also being applied to locate wild ungulates which were more recently introduced to some islands. All invasive mammals have been eradicated from some remote small islands, and it may soon be possible to manage areas on larger islands to be free of invasive mammals at least during seasonally important periods for native species.

Conference Paper↗

Ciscoes ( Coregonus , subgenus Leucichthys ) of the Laurentian Great Lakes and Lake Nipigon

This study of the ciscoes ( Coregonus , subgenus Leucichthys ) of the Great Lakes and Lake Nipigon represents a furtherance through 2015 of field research initiated by Walter Koelz in 1917 and continued by Stanford Smith in the mid-1900s—a period spanning nearly a century. Like Koelz’s study, this work contains information on taxonomy, geographical distribution, ecology, and status of species (here considered forms). Of the seven currently recognized forms ( C. artedi , C. hoyi , C. johannae , C. kiyi , C. nigripinnis , C. reighardi , and C. zenithicus ) described by Koelz as major in his 1929 monograph, two ( C. johannae and C. reighardi ) are extinct. In addition, C. alpenae , described by Koelz but subsequently synonymized with C. zenithicus , although extinct, is recognized as valid making a total of eight major forms. Six of these forms, all but C. artedi and C. hoyi , have been lost from Lake Michigan, and seven have been lost from Lake Huron, leaving in Lake Huron only C. artedi and an introgressed deepwater form that we term a hybrid swarm. C. artedi appears, like its sister form C. alpenae , to have been lost from Lake Erie. Only C. artedi remains extant in Lake Ontario, its three sister forms ( C. hoyi , C. kiyi , and C. reighardi ) having disappeared long ago. Lakes Superior and Nipigon have retained their original species flocks consisting of four forms each: C. artedi , C. hoyi , and C. zenithicus in both lakes; C. kiyi in Lake Superior; and C. nigripinnis in Lake Nipigon. Morphological deviations from the morphotypes described by Koelz have been modest in contemporary samples. Overall, C. kiyi and C. artedi were the most morphologically stable forms while C. hoyi , C. nigripinnis , and C. zenithicus were the least stable. Although contemporary populations of C. artedi from Lakes Michigan and Huron are highly diverged from the morphotypes described by Koelz, the contemporary samples were of undescribed deep-bodied forms unlikely to have been sampled by Koelz because of their association with bays. Of the two intact species flocks, Lake Nipigon’s was much less stable morphologically than Lake Superior’s even though Lake Nipigon is far less disturbed. Two priorities for research are determining the role of developmental plasticity in morphological divergence, especially within C. zenithicus of Lake Superior, and the basis for morphological divergence in C. artedi .

Lake Nipigon, Laurentian Great Lakes↗

Minimizing effects of methodological decisions on interpretation and prediction in species distribution studies: An example with background selection

Evaluating the conditions where a species can persist is an important question in ecology both to understand tolerances of organisms and to predict distributions across landscapes. Presence data combined with background or pseudo-absence locations are commonly used with species distribution modeling to develop these relationships. However, there is not a standard method to generate background or pseudo-absence locations, and method choice affects model outcomes. We evaluated combinations of both model algorithms (simple and complex generalized linear models, multivariate adaptive regression splines, Maxent, boosted regression trees, and random forest) and background methods (random, minimum convex polygon, and continuous and binary kernel density estimator (KDE)) to assess the sensitivity of model outcomes to choices made. We evaluated six questions related to model results, including five beyond the common comparison of model accuracy assessment metrics (biological interpretability of response curves, cross-validation robustness, independent data accuracy and robustness, and prediction consistency). For our case study with cheatgrass in the western US, random forest was least sensitive to background choice and the binary KDE method was least sensitive to model algorithm choice. While this outcome may not hold for other locations or species, the methods we used can be implemented to help determine appropriate methodologies for particular research questions.

Ecological Modelling↗

Optimal sampling design for spatial capture‐recapture

Spatial capture‐recapture (SCR) has emerged as the industry standard for estimating population density by leveraging information from spatial locations of repeat encounters of individuals. The precision of density estimates depends fundamentally on the number and spatial configuration of traps. Despite this knowledge, existing sampling design recommendations are heuristic and their performance remains untested for most practical applications. To address this issue, we propose a genetic algorithm that minimizes any sensible, criteria‐based objective function to produce near‐optimal sampling designs. To motivate the idea of optimality, we compare the performance of designs optimized using three model‐based criteria related to the probability of capture. We use simulation to show that these designs out‐perform those based on existing recommendations in terms of bias, precision, and accuracy in the estimation of population size. Our approach, available as a function in the R package oSCR, allows conservation practitioners and researchers to generate customized and improved sampling designs for wildlife monitoring.

Ecology↗

Use of dynamic occupancy models to assess the response of Darters (Teleostei: Percidae) to varying hydrothermal conditions in a southeastern United States tailwater

During the past 100 years, most large rivers in North America have been altered for flood control, hydropower, navigation or water supply development. Although these activities clearly provide important human services, their associated environmental disturbances can profoundly affect stream-dwelling organisms. We used dynamic multi-species occupancy models combined with a trait-based approach to estimate the influence of site-level and species-level characteristics on patch dynamic rates for 15 darter species native to the Elk River, a large, flow-regulated Tennessee River tributary in Tennessee and Alabama. Dynamic occupancy modelling results indicated that for every 2.5 °C increase in stream temperature, darters were 3.94 times more likely to colonize previously unoccupied stream reaches. Additionally, large-bodied darter species were 3.72 times more likely to colonize stream reaches compared with small-bodied species, but crevice-spawning darter species were 5.24 times less likely to colonize previously unoccupied stream reaches. In contrast, darters were 2.21 times less likely to become locally extinct for every 2.5 °C increase in stream temperature, but high stream discharge conditions elevated the risk of local extinction. Lastly, the presence of populations in neighbouring upstream study reaches contributed to a lower risk of extinction, whereas the presence of populations in neighbouring downstream study reaches contributed to higher rates of colonization. Our study demonstrates the application of a trait-based approach combined with a metapopulation framework to assess the patch dynamics of darters in a regulated river. Results from our study will provide a baseline for evaluating the ecological consequences of alternative dam operations.

River Research and Applications↗

Thermal characteristics of amphibian microhabitats in a fire-disturbed landscape

Disturbance has long been a central issue in amphibian conservation, often regarding negative effects of logging or other forest management activities, but some amphibians seem to prefer disturbed habitats. After documenting increased use of recently burned forests by boreal toads (Bufo boreas), we hypothesized that burned habitats provided improved thermal opportunities in terrestrial habitats. We tested this hypothesis by conducting a radio telemetry study of habitat use (reported previously) and by using physical models that simulated the temperature of adult toads. We deployed 108 physical models in and adjacent to a 1-year old burn using a fully-replicated design with three burn severities (unburned, partial, high severity) and four microhabitats (open surface, under vegetation, under log, in burrow). Model temperatures were compared to a range of preferred temperatures in published studies. We found 70% more observations within the preferred temperature range of B. boreas in forests burned with high severity than in unburned areas. Burned forest was warmer than unburned forest across all microhabitats, but the largest relative difference was in burrows, which averaged 3 ??C warmer in high-severity burn areas and remained warmer though the night. More than twice as many observations were within the preferred temperature range in high-severity burrows than in unburned burrows. Areas burned with high severity were still warmer than unburned forest 3 years after the fire. Habitat use of toads during the concurrent radio telemetry study matched that predicted by the physical models. These results suggest there are fitness-linked benefits to toads using burned habitats, such as increased growth, fertility, and possibly disease resistance. However, increased soil temperatures that result from wildfire may be detrimental to other amphibian species that prefer cooler temperatures and stable environments. More broadly, our data illustrate the use of physical models to measure and interpret changes that amphibians may experience from disturbance, and highlight the need for research linking vital rates such as growth and survival to disturbance.

Forest Ecology and Management↗

Reconstructing the response of C3 and C4 plants to decadal-scale climate change during the late Pleistocene in southern Illinois using isotopic analyses of calcified rootlets

The ??13C and ??18O values of well-preserved carbonate rhizoliths (CRs) provide detailed insights into changes in the abundance of C3 and C4 plants in response to approximately decadal-scale changes in growing-season climate. We performed stable isotope analyses on 35-40 CRs sampled at 1-cm intervals from an 18-cm-thick paleosol formed in southern Illinois during Wisconsin interstadial 2. Minimum ??13C values show little variation with depth, whereas maximum values vary dramatically, and average values show noticeable variability; maximum ??18O values vary less than the minimum ??18O values. These findings indicate that a diverse and stable C3 flora with a limited number of C4 grass species prevailed during this interval, and suggest that the maximum growing-season temperatures were relatively stable, but minimum growing-season temperatures varied considerably. Two general patterns characterize the relationships between the ??13C and ??18O values obtained from the 1-cm samples. In some cases, low ??13C values correspond to low ??18O values and high ??13C values correspond to high ??18O values, suggesting that cooler growing-season temperatures favored C3 and warmer growing-season temperatures favored C4 plants. In other cases, low ??13C values correspond to high ??18O values, likely suggesting that wetter growing-season conditions were favorable to C3 plants. The high density of well-preserved CRs in this paleosol provides a unique opportunity to study detailed ecological responses to high-resolution variability in growing-season climate. ?? 2006 University of Washington.

Quaternary Research↗