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Polar and brown bear genomes reveal ancient admixture and demographic footprints of past climate change

Polar bears (PBs) are superbly adapted to the extreme Arctic environment and have become emblematic of the threat to biodiversity from global climate change. Their divergence from the lower-latitude brown bear provides a textbook example of rapid evolution of distinct phenotypes. However, limited mitochondrial and nuclear DNA evidence conflicts in the timing of PB origin as well as placement of the species within versus sister to the brown bear lineage. We gathered extensive genomic sequence data from contemporary polar, brown, and American black bear samples, in addition to a 130,000- to 110,000-y old PB, to examine this problem from a genome-wide perspective. Nuclear DNA markers reflect a species tree consistent with expectation, showing polar and brown bears to be sister species. However, for the enigmatic brown bears native to Alaska's Alexander Archipelago, we estimate that not only their mitochondrial genome, but also 5–10% of their nuclear genome, is most closely related to PBs, indicating ancient admixture between the two species. Explicit admixture analyses are consistent with ancient splits among PBs, brown bears and black bears that were later followed by occasional admixture. We also provide paleodemographic estimates that suggest bear evolution has tracked key climate events, and that PB in particular experienced a prolonged and dramatic decline in its effective population size during the last ca. 500,000 years. We demonstrate that brown bears and PBs have had sufficiently independent evolutionary histories over the last 4–5 million years to leave imprints in the PB nuclear genome that likely are associated with ecological adaptation to the Arctic environment.

Proceedings of the National Academy of Sciences

Prevention, early detection and containment of invasive, nonnative plants in the Hawaiian Islands: current efforts and needs

Introduction: Invasive, non-native plants (or environmental weeds) have long been recognized as a major threat to the native biodiversity of oceanic islands (Cronk & Fuller, 1995; Denslow, 2003). Globally, several hundred non-native plant species have been reported to have major impacts on natural areas on oceanic islands (Kueffer et al ., 2009). In Hawaii, at least some 50 non-native plant species reach dominance in natural areas (Kueffer et al ., 2009) and many of them are known to impact ecosystem processes or biodiversity. One example is the invasive Australian tree fern ( Cyathea cooperi ), which has been shown to be very efficient at utilizing soil nitrogen and can grow six times as rapidly in height, maintain four times more fronds, and produce significantly more fertile fronds per month than the native Hawaiian endemic tree ferns, Cibotium spp. (Durand & Goldstein, 2001a, b). Additionally, while native tree ferns provide an ideal substrate for epiphytic growth of many understory ferns and flowering plants, the Australian tree fern has the effect of impoverishing the understory and failing to support an abundance of native epiphytes (Medeiros & Loope, 1993). Other notorious examples of invasive plant species problematic for biodiversity and ecosystem processes in Hawaii include miconia ( Miconia calvescens ), strawberry guava ( Psidium cattleianum ), albizia ( Falcataria moluccana ), firetree ( Morella faya ), clidemia ( Clidemia hirta ), kahili ginger ( Hedychium gardnerianum ), and fountain grass ( Pennisetum setaceum ), to name just a few. Fireweed ( Senecio madagascariensis ) is a recent example of a seriously problematic invasive species for Hawaii’s agriculture and is damaging certain high-elevations native ecosystems as well. The threat of invasive plants has long been recognized in Hawaii and is well documented (e.g. Cox, 1999; Loope & Kraus, 2009 in press; Loope et al ., 2004; Mooney & Drake, 1986; Stone & Scott, 1985; Stone et al. , 1992). In many respects, Hawaii may be near the forefront among national and international efforts to address the burgeoning threat of invasive plants, perhaps especially in the field of outreach and education (Holt, 1996; Van Driesche & Van Driesche, 2000). However, given the scale of the problem many challenges still need to be addressed and gaps in the existing management system need to be identified. In particular, it appears that new non-native plant species are still introduced to the Hawaiian Islands at a high rate with little or no regard for their potential invasiveness. In fact, a Pacific-wide and a global survey of non-native plants on oceanic islands have both shown that on Hawaii among all archipelagos by far the highest number of problematic invasive species known from other areas in the world is already present (Denslow et al . 2009, Kueffer et al . 2009). Hawaii lacks an effective mechanism for tracking what species are present or incoming. For instance, early detection nursery surveys conducted on Maui in 2008 found over 300 species of cultivated vascular plants that have not previously been recorded in Hawaii (Starr et al. , in prep.). In spite of an innovative Hawaii Biological Survey (e.g. Eldredge & Evenhuis, 2003), there is no mechanism for recording presence of a species until it becomes naturalized. Some of these new introductions may quickly become serious pests. Fireweed, first recorded in Hawaii on the Big Island in the early 1980s, is now considered one of the Kueffer & Loope 2009 5/48 worst weeds of pastures and is also invading natural areas from near sea level to above 10,000 feet. Although the cultivated and as yet non-invasive Cortaderia selloana has been present in Hawaii for 50 years or more, the morphologically similar Cortaderia jubata was simultaneously found to be present on Maui and invading on a large scale in 1989. It played an important role in inspiring the establishment of the Maui Invasive Species Committee (MISC) in 1997, and MISC now spends roughly $200,000 per year removing and containing C. jubata to keep it from becoming widespread in high elevation conservation lands of East and West Maui. The existence of many similar examples shows that to date regulatory action to prevent new invasive plant species from establishing and spreading in Hawaii has not yet been as successful as it needs to be. In particular, because some problematic invasive species known from other areas in the world (Kueffer et al ., 2009; Weber, 2003) have not yet been recorded from Hawaii, preventive measures against the introduction and spread of such likely invasive species is therefore an urgent need for Hawaii. Indeed, regulation of importation and early detection and eradication of introduced species before they become abundant and widespread are widely considered the most cost-efficient and often only effective measures against the threat of new invasive species (Kueffer & Hirsch Hadorn, 2008; Wittenberg & Cock, 2001). Timing seems favorable for Hawaii to achieve effective protection against the threat of new invasive species through prevention, early detection, and eradication/containment. Through the establishment and evolution of Invasive Species Committees (ISCs) on each major Hawaiian island, the institutional capacity has been built up for prevention, early detection, containment, and outreach at an island scale. Weed risk assessment (Daehler et al ., 2004) and early detection methodologies (Starr et al. , in review-a, b) have been developed and tested specifically for Hawaii. Containment strategies have been successful (e.g., Special Ecological Areas in Hawaii Volcanoes National Park), and so have eradications of particular species on an island scale (e.g. mullein ( Verbascum thapsus ) and other species on Maui, fireweed ( Senecio madagascariensis ) on Kauai). These successful management strategies may be further strengthened through recently developed novel approaches in research (e.g. remote sensing, species distribution modelling, and molecular genetics tools). Another major recent achievement is the gained support of the plant industry for preventive measures against invasive species (see p. 13ff). Last but not least, regulatory action is also moving forward. Passage of House Bill 2517 by the 2008 Hawaii House and Senate and prompt signing of the bill into law by the Governor provides hope that action to ban the sale of a meaningful suite of restricted weeds can quickly proceed through the rulemaking phase into the implementation phase. This report documents these achievements and experiences and provides a range of perspectives on how to further develop prevention, early detection and containment of invasive species in Hawaii. The report is based on a symposium and workshop held at the 2008 Hawaii Conservation Conference in Honolulu on 31 July 2008.

Hawaii

Threats posed by the Fungal Kingdom to humans, wildlife, and agriculture

The Fungal Kingdom includes at least six million eukaryotic species and is remarkable with respect to its profound impact on global health, biodiversity, ecology, agriculture, manufacturing, and biomedical research. Approximately 625 fungal species have been reported to infect vertebrates, 200 of which can be human-associated, either as commensals and members of our microbiome or as pathogens that cause infectious diseases. These organisms pose a growing threat to human health with the global increase in the incidence of invasive fungal infections, prevalence of fungal allergy, and the evolution of fungal pathogens resistant to some or all current classes of antifungals. More broadly, there has been an unprecedented and worldwide emergence of fungal pathogens impacting animal and plant biodiversity. Approximately 8,000 species of fungi and Oomycetes are associated with plant disease. Indeed, across agriculture, such fungal diseases of plants include new devastating epidemics of trees and jeopardize food security worldwide by causing epidemics in staple and commodity crops that feed billions. Further, ingestion of mycotoxins contributes to ill health and causes cancer. Coordinated international research efforts, enhanced technology translation, and greater policy outreach by scientists are needed to more fully understand the biology and drivers that underlie the emergence of fungal diseases and to mitigate against their impacts. Here, we focus on poignant examples of emerging fungal threats in each of three areas: human health, wildlife biodiversity, and food security.

mBio

Upper Rio Grande Basin water-resource status and trends: Focus area study review and synthesis

The Upper Rio Grande Basin (URGB) is a critical international water resource under pressure from a myriad of climatic, ecological, infrastructural, water-use, and legal constraints. The objective of this study is to provide a comprehensive assessment of the spatial distribution and temporal trends of selected water-budget components (snow processes, evapotranspiration (ET), streamflow processes, and groundwater storage) using integrated analyses, such as watershed modeling and water availability and use data in the URGB over the past three decades. A spatially distributed snow evolution modeling system simulated snowpack processes over 34 years (1984–2017). It highlighted snow water equivalent declines from -35 to -77 mm/decade with widespread variability across elevation zones and land cover types. Gridded actual ET data from the SSEBop model were developed and tested for the URGB and demonstrated that all land-cover types had significant decreasing trends (1986-2015) ranging from -14 to -80 mm/decade. Conductivity-mass-balance (CMB) hydrograph separation results found that baseflow forms a large component of total streamflow, ranging from 29 to 69% (49% average) of total streamflow at 17 URGB sites upstream of Albuquerque, NM. Three of 4 graphical hydrograph separation methods in the U.S. Geological Survey Groundwater Toolbox were found to be inappropriate for estimating baseflow in the URGB; the most promising method, baseflow index (BFI) Standard, was optimized using CMB data and tested at three URGB sites, with resulting overestimation of 0 to 47%. Simulated changes in groundwater storage were extracted from historical and recent groundwater-flow models of select alluvial basins (San Luis, Española, Middle Rio Grande, and Tularosa-Hueco). In general, decreases in groundwater storage were observed from 1903 to 2013 except for the San Luis alluvial basin (Colorado), where periods of recovery are observed. The PRMS hydrologic model was successfully calibrated for 9 near-native subbasins (Nash-Sutcliffe efficiency 0.47 to 0.85) and parameters translated to the remaining subbasins; compared to simulated near-native flows (with minimal influence of reservoirs or diversions), observed Rio Grande streamgage flows demonstrated reductions of 40% or more for New Mexico and Texas areas of the basin. Significant decreasing trends (1980-2015) in precipitation, snowmelt rate, streamflow, and baseflow were observed at many of the 12 streamgage basins studied, which suggests that the decreasing trends for actual ET may be related to overall decreasing water availability in the basin, with negative implications for agricultural production and groundwater abstraction. Water security concerns arise from our findings of higher fraction precipitation as rain, slower snowmelt rates leading to decreasing streamflow production, and an increasing fraction of baseflow, all of which will affect the timing and magnitude of water available for human needs in the basin.

Upper Rio Grande Basin

Computationally efficient statistical differential equation modeling using homogenization

Statistical models using partial differential equations (PDEs) to describe dynamically evolving natural systems are appearing in the scientific literature with some regularity in recent years. Often such studies seek to characterize the dynamics of temporal or spatio-temporal phenomena such as invasive species, consumer-resource interactions, community evolution, and resource selection. Specifically, in the spatial setting, data are often available at varying spatial and temporal scales. Additionally, the necessary numerical integration of a PDE may be computationally infeasible over the spatial support of interest. We present an approach to impose computationally advantageous changes of support in statistical implementations of PDE models and demonstrate its utility through simulation using a form of PDE known as “ecological diffusion.” We also apply a statistical ecological diffusion model to a data set involving the spread of mountain pine beetle (Dendroctonus ponderosae) in Idaho, USA.

Journal of Agricultural, Biological, and Environme

Maintenance and dissemination of avian-origin influenza A virus within the northern Atlantic Flyway of North America

Wild waterbirds, the natural reservoirs for avian influenza viruses, undergo migratory movements each year, connecting breeding and wintering grounds within broad corridors known as flyways. In a continental or global view, the study of virus movements within and across flyways is important to understanding virus diversity, evolution, and movement. From 2015 to 2017, we sampled waterfowl from breeding (Maine) and wintering (Maryland) areas within the Atlantic Flyway (AF) along the east coast of North America to investigate the spatio-temporal trends in persistence and spread of influenza A viruses (IAV). We isolated 109 IAVs from 1,821 cloacal / oropharyngeal samples targeting mallards (Anas platyrhynchos) and American black ducks (Anas rubripes) , two species having ecological and conservation importance in the flyway that are also host reservoirs of IAV. Isolates with >99% nucleotide similarity at all gene segments were found between eight pairs of birds in the northern site across years, indicating some degree of stability among genome constellations and the possibility of environmental persistence. No movement of whole genome constellations were identified between the two parts of the flyway, however, virus gene flow between the northern and southern study locations was evident. Examination of banding records indicate direct migratory waterfowl movements between the two locations within an annual season, providing a mechanism for the inferred viral gene flow. Bayesian phylogenetic analyses provided evidence for virus dissemination from other North American wild birds to AF dabbling ducks (Anatinae), shorebirds (Charidriformes), and poultry (Galliformes). Evidence was found for virus dissemination from shorebirds to gulls (Laridae), and dabbling ducks to shorebirds and poultry. The findings from this study contribute to the understanding of IAV ecology in waterfowl within the AF.

Chesapeake Bay

Cenozoic planktonic marine diatom diversity and correlation to climate change

Marine planktonic diatoms export carbon to the deep ocean, playing a key role in the global carbon cycle. Although commonly thought to have diversified over the Cenozoic as global oceans cooled, only two conflicting quantitative reconstructions exist, both from the Neptune deep-sea microfossil occurrences database. Total diversity shows Cenozoic increase but is sample size biased; conventional subsampling shows little net change. We calculate diversity from a separately compiled new diatom species range catalog, and recalculate Neptune subsampled-in-bin diversity using new methods to correct for increasing Cenozoic geographic endemism and decreasing Cenozoic evenness. We find coherent, substantial Cenozoic diversification in both datasets. Many living cold water species, including species important for export productivity, originate only in the latest Miocene or younger. We make a first quantitative comparison of diatom diversity to the global Cenozoic benthic ∂ 18 O (climate) and carbon cycle records (∂ 13 C, and 20-0 Ma pCO 2 ). Warmer climates are strongly correlated with lower diatom diversity (raw: rho = .92, p<.001; detrended, r = .6, p = .01). Diatoms were 20% less diverse in the early late Miocene, when temperatures and pCO 2 were only moderately higher than today. Diversity is strongly correlated to both ∂ 13 C and pCO 2 over the last 15 my (for both: r>.9, detrended r>.6, all p<.001), but only weakly over the earlier Cenozoic, suggesting increasingly strong linkage of diatom and climate evolution in the Neogene. Our results suggest that many living marine planktonic diatom species may be at risk of extinction in future warm oceans, with an unknown but potentially substantial negative impact on the ocean biologic pump and oceanic carbon sequestration. We cannot however extrapolate our my-scale correlations with generic climate proxies to anthropogenic time-scales of warming without additional species-specific information on proximate ecologic controls.

PLoS ONE

Introduced species as evolutionary traps

Invasive species can alter environments in such a way that normal behavioural decision-making rules of native species are no longer adaptive. The evolutionary trap concept provides a useful framework for predicting and managing the impact of harmful invasive species. We discuss how native species can respond to changes in their selective regime via evolution or learning. We also propose novel management strategies to promote the long-term co-existence of native and introduced species in cases where the eradication of the latter is either economically or biologically unrealistic.

Ecology Letters

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

Screening of duplicated loci reveals hidden divergence patterns in a complex salmonid genome

A whole-genome duplication (WGD) doubles the entire genomic content of a species and is thought to have catalysed adaptive radiation in some polyploid-origin lineages. However, little is known about general consequences of a WGD because gene duplicates (i.e., paralogs) are commonly filtered in genomic studies; such filtering may remove substantial portions of the genome in data sets from polyploid-origin species. We demonstrate a new method that enables genome-wide scans for signatures of selection at both nonduplicated and duplicated loci by taking locus-specific copy number into account. We apply this method to RAD sequence data from different ecotypes of a polyploid-origin salmonid ( Oncorhynchus nerka ) and reveal signatures of divergent selection that would have been missed if duplicated loci were filtered. We also find conserved signatures of elevated divergence at pairs of homeologous chromosomes with residual tetrasomic inheritance, suggesting that joint evolution of some nondiverged gene duplicates may affect the adaptive potential of these genes. These findings illustrate that including duplicated loci in genomic analyses enables novel insights into the evolutionary consequences of WGDs and local segmental gene duplications.

Molecular Ecology

The epidemiology of avian pox and interaction with avian malaria in Hawaiian forest birds

Despite the purported role of avian pox ( Avipoxvirus spp.) in the decline of endemic Hawaiian birds, few studies have been conducted on the dynamics of this disease, its impact on free‐living avian populations, or its interactions with avian malaria ( Plasmodium relictum ). We conducted four longitudinal studies of 3–7 yr in length and used generalized linear models to evaluate cross‐sectional prevalence of active pox infection and individuals with healed deformities that had recovered from pox. Our goal was to understand how species, season, elevation, malaria infection, and other biological characteristics influenced pox infection in ʻApapane, Hawaiʻi ʻAmakihi, ʻIʻiwi, and Japanese White‐eye across low‐, mid‐, and high‐elevation forests on the island of Hawaiʻi. We also used multi‐state capture‐recapture (longitudinal) models to estimate pox infection rates, recovery rates, and potential pox‐related mortality. Pox infection rates were typically highest in low‐elevation forests, followed by mid‐elevation forests, and lowest in high‐elevation forests. We also found seasonal changes in pox prevalence throughout the annual cycle; typically increasing from spring through summer, peaking in fall, and declining in winter. These seasonal changes occurred in low‐ and mid‐elevation forests, but not in high elevations where pox infection was low. Seasonal and elevation patterns of pox infection are like those for avian malaria, strongly implicating mosquito vectors, rather than other biting arthropods or contact transmission, as the primary source of transmitting both diseases. Most native Hawaiian birds recovered from pox infection within 6 months; frequently without permanent lesions. Contrary to our expectations, we found no direct evidence that pox is a substantial mortality factor in any of the three native bird species we studied. Birds with chronic malaria infection were more likely to have both active pox infection and healed pox lesions suggesting a synergistic interaction that may influence the evolution of pox virulence. Because pox infection can be assessed visually, and birds have a high recovery rate, this disease may be a sensitive indicator of the seasonal and annual risk of transmission of malaria in Hawaiʻi.

Ecological Monographs

The "terminal Triassic catastrophic extinction event" in perspective: A review of carboniferous through Early Jurassic terrestrial vertebrate extinction patterns

A catastrophic terminal Triassic extinction event among terrestrial vertebrates is not supported by available evidence. The current model for such an extinction is based on at least eight weak or untenable assumptions: (1) a terminal Triassic extinction-inducing asteroid impact occurred, (2) a terminal Triassic synchronous mass extinction of terrestrial vertebrates occurred, (3) a concurrent terminal Triassic marine extinction occurred, (4) all terrestrial vertebrate families have similar diversities and ecologies, (5) changes in familial diversity can be gauged accurately from the known fossil record, (6) extinction of families can be compared through time without normalizing for changes in familial diversity through time, (7) extinction rates can be compared without normalizing for differing lengths of geologic stages, and (8) catastrophic mass extinctions do not select for small size. These assumptions have resulted in unsupportable and (or) erroneous conclusions. Carboniferous through Early Jurassic terrestrial vertebrate families mostly have evolution and extinction patterns unlike the vertebrate evolution and extinction patterns during the terminal Cretaceous event. Only the Serpukhovian (mid Carboniferous) extinction event shows strong analogy to the terminal Cretaceous event. Available data suggest no terminal Triassic extinction anomaly, but rather a prolonged and nearly steady decline in the global terrestrial vertebrate extinction rate throughout the Triassic and earliest Jurassic.

Palaeogeography, Palaeoclimatology, Palaeoecology

Geological sampling data and benthic biota classification: Buzzards Bay and Vineyard Sound, Massachusetts

Sea-floor sample collection is an important component of a statewide cooperative mapping effort between the U.S. Geological Survey (USGS) and the Massachusetts Office of Coastal Zone Management (CZM). Sediment grab samples, bottom photographs, and video transects were collected within Vineyard Sound and Buzzards Bay in 2010 aboard the research vessel Connecticut . This report contains sample data and related information, including analyses of surficial-sediment grab samples, locations and images of sea-floor photography, survey lines along which sea-floor video was collected, and a classification of benthic biota observed in sea-floor photographs and based on the Coastal and Marine Ecological Classification Standard (CMECS). These sample data and analyses information are used to verify interpretations of geophysical data and are an essential part of geologic maps of the sea floor. These data also provide a valuable inventory of benthic habitat and resources. Geographic information system (GIS) data, maps, and interpretations, produced through the USGS and CZM mapping cooperative, are intended to aid efforts to manage coastal and marine resources and to provide baseline information for research focused on coastal evolution and environmental change.

Massachusetts

Group composition effects on aggressive interpack interactions of gray wolves in Yellowstone National Park

Knowledge of characteristics that promote group success during intraspecific encounters is key to understanding the adaptive advantages of sociality for many group-living species. In addition, some individuals in a group may be more likely than others to influence intergroup conflicts, a relatively neglected idea in research on social animals. Here we use observations of aggressive interactions between wolf ( Canis lupus ) packs over an extended period and use pack characteristics to determine which groups had an advantage over their opponents. During 16 years of observation in Yellowstone National Park from 1995 to 2010, we documented 121 interpack aggressive interactions. We recorded pack sizes, compositions, and spatial orientation related to residency to determine their effects on the outcomes of interactions between packs. Relative pack size (RPS) improved the odds of a pack displacing its opponent. However, pack composition moderated the effect of RPS as packs with relatively more old members (>6.0 years old) or adult males had higher odds of winning despite a numerical disadvantage. The location of the interaction with respect to pack territories had no effect on the outcome of interpack interactions. Although the importance of RPS in successful territorial defense suggests the evolution and maintenance of group living may be at least partly due to larger packs&rsquo; success during interpack interactions, group composition is also an important factor, highlighting that some individuals are more valuable than others during interpack conflicts.

Wyoming

Processes governing phytoplankton blooms in estuaries. II: The role of horizontal transport

The development and distribution of phytoplankton blooms in estuaries are functions of both local conditions (i.e. the production-loss balance for a water column at a particular spatial location) and large-scale horizontal transport. In this study, the second of a 2-paper series, we use a depth-averaged hydrodynamic-biological model to identify transport-related mechanisms impacting phytoplankton biomass accumulation and distribution on a system level. We chose South San Francisco Bay as a model domain, since its combination of a deep channel surrounded by broad shoals is typical of drowned-river estuaries. Five general mechanisms involving interaction of horizontal transport with variability in local conditions are discussed. Residual (on the order of days to weeks) transport mechanisms affecting bloom development and location include residence time/export, import, and the role of deep channel regions as conduits for mass transport. Interactions occurring on tidal time scales, i.e. on the order of hours) include the phasing of lateral oscillatory tidal flow relative to temporal changes in local net phytoplankton growth rates, as well as lateral sloshing of shoal-derived biomass into deep channel regions during ebb and back into shallow regions during flood tide. Based on these results, we conclude that: (1) while local conditions control whether a bloom is possible, the combination of transport and spatial-temporal variability in local conditions determines if and where a bloom will actually occur; (2) tidal-time-scale physical-biological interactions provide important mechanisms for bloom development and evolution. As a result of both subtidal and tidal-time-scale transport processes, peak biomass may not be observed where local conditions are most favorable to phytoplankton production, and inherently unproductive areas may be regions of high biomass accumulation.

Marine Ecology Progress Series

Terrain, vegetation, and landscape evolution of the R4D research site, Brooks Range Foothills, Alaska

Maps of the vegetation and terrain of a 22 km2 area centered on the Department of Energy (DOE) R4D (Response, Resistance, Resilience to and Recovery from Disturbance in Arctic Ecosystems) study site in the Southern Foothills Physiographic Province of Alaska were made using integrated geobotanical mapping procedures and a geographic-information system. Typical land forms and surface f orms include hillslope water tracks, Sagavanirktok-age till deposits, nonsorted stone stripes, and colluvial-basin deposits. Thirty-two plant communities are described; the dominant vegetation (51% of the mapped area) is moist tussock-sedge, dwarf-shrub tundra dominated by Eriophorum vaginatum or Carex bigelowii. Much of the spatial variation in the mapped geobotanical characters reflects different-aged glaciated surfaces. Shannon-Wienerin dices indicate that the more mature landscapes, represented by retransported hillslope deposits and basin colluvium, are less heterogeneous than newer landscapes such as surficial till deposits and floodplains. A typical toposequence on a mid-Pleistocene-age surface is discussed with respect to evolution of the landscape. Thick Sphagnum moss layers occur on lower hillslopes, and the patterns of moss-layer development, heat flux, active layer thickness, and ground-ice are seen as keys to developing thermokarst-susceptibility maps.

Alaska

Near-surface stratigraphy and morphology, Mississippi Inner Shelf, northern Gulf of Mexico

Over the past decade, the Mississippi Barrier Islands have been the focus of a comprehensive geologic investigation by the U.S. Geological Survey (USGS), in collaboration with the U.S. Army Corps of Engineers (USACE) and the National Park Service (NPS). The islands (Dauphin, Petite Bois, Horn, East Ship, West Ship, and Cat) are part of the Gulf Islands National Seashore (GUIS), and provide a diverse ecological habitat, protect the mainland from storm waves, and help maintain estuarine conditions within Mississippi Sound. Over the past century, the islands have been in a state of decline with respect to elevation and land-area loss. In 2005, the islands were severely impacted by Hurricane Katrina, which inundated them with a storm surge of 8 meters, causing severe shoreface erosion and widening breaches in Dauphin, West Ship, and Cat Islands. To evaluate the impact and fate of the islands, understanding their evolution and resiliency became a priority for the USGS under the Northern Gulf of Mexico Ecosystem Change and Hazard Susceptibility Project. The project formed the basis for collaboration with the USACE Mississippi Coastal Improvement Project, which is intended to restore portions of coastal Mississippi and GUIS affected by storm impact. Since then, many studies have contributed to our understanding of the islands&rsquo; morphology and nearshore stratigraphy. This report expands upon the nearshore component to provide a stratigraphic and morphologic assessment offshore of Petit Bois Island. In June 2013, as part of the MsCIP project, the USGS conducted a geophysical survey consisting of about 650 line-kilometers (km), encompassing an area of approximately 212 square kilometers (km 2 ). The survey area extended from 1 to 13 km offshore of Petite Bois Island. The geophysical investigation included interferometric swath bathymetry, sidescan sonar, and chirp subbottom profiling. The intent of the survey was to provide geologic information that would assist the USACE in developing a sediment sampling strategy for identifying deposits suitable for shoreline restoration operations. The data from the geophysical survey would also further our understanding of the geologic framework along the inner shelf. Numerous seafloor and subbottom features were identified. At the surface, shoals and shelf sand sheets of various sizes and orientations are the predominant morphology. In the subsurface, Holocene- and Pleistocene-age features include marine transgressive deposits infilling older fluvia distributary systems. These interpretations from the geophysical research were integrated with sediment cores collected by the USGS and USACE to provide textural and volumetric information.

Mississippi-Alabama shelf

Mediterranean-type ecosystems: the influence of biodiversity on their functioning

Ecosystems in the Mediterranean-climate regions of the world have served as a unit for comparative ecological studies for over two decades. The cohesiveness of research in this set of widely distributed regions rests on the similarity of the climates where they occur, and the identifiable convergence in elements of their vegetation structure (Di Castri and Mooney 1973). In this chapter we review functional aspects of what have come to be known as Mediterranean-type ecosystems (METs) in the context of a concerned global interest in the sustainability of the human environment and its dependence on biological diversity. The approach we adopt here is to look for evidence that this biodiversity, for which some MTEs are renowned (Cowling, 1992; Hobbs, 1992), has an influence on processes which are important both for the maintenance of natural systems, and for providing "ecosystem services" with human utility. Almost a century ago, Schimper (1903) recognized the biological similarities between five widely separated regions characterized by Mediterranean-type climates, and much comparative work has been done on that basis since. These regions comprise the Mediterranean basin itself, a major portion of California, central Chile, the southwestern and southern extremities of South Africa, and parts of southwestern and southern Australia (Figure 7.1). The first attention paid to MTEs in terms of quantitative ecological research arose out of the International Biological Programme (IBP) of the 1960s and 1970s. Those efforts focused on comparisons between the Chilean and Californian systems (Mooney 1977), and dealt with parallel models of ecosystem processes, especially water flux (Fuentes et al 1995). Because of the already perceived similarities between vegetation in these and the other three regions, the project was soon extended to include all five regions. The first broad comparative overview was published as an anthology which considered the origins and the convergent evolution of MTE components (Di Castri and Mooney 1973). Although the currently accepted classifications of the MTEs is to some extent artificial, it does provide a basis for comparative work, as well as placing mild, temperate winter rainfall regions in perspective with other system types, such as forests, arid lands and even savannas. It is against this backdrop that the MTE research collegium has grown, giving rise to the organizational structure known as ISOMED (the International Society of Mediterranean Ecologists), which has convened regular conferences under the label MEDECOS, plus a number of extra meetings on specific topics (Table 7.1). One of the more recent in this series of MTE meetings was convened under the auspices of ICSU's Scientific Committee on Problems of the Environment (SCOPE) (see Table 7.1), and dealt with the questions about the functional value of biodiversity. This chapter is based on that meeting and its proceedings (Richardson and Cowling 1993); David and Richardson 1995), and is a distillation of input by teams of ecologists from each of the five regions.

Book chapter