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Toward a scientifically rigorous basis for developing mapped ecological regions.

Despite the wide use of ecological regions in conservation and resource-management evaluations and assessments, a commonly accepted theoretical basis for ecological regionalization does not exist. This fact, along with the paucity of focus on ecological regionalization by professional associations, journals, and faculties, has inhibited the advancement of a broadly acceptable scientific basis for the development, use, and verification of ecological regions. The central contention of this article is that ecological regions should improve our understanding of geographic and ecological phenomena associated with biotic and abiotic processes occurring in individual regions and also of processes characteristic of interactions and dependencies among multiple regions. Research associated with any ecoregional framework should facilitate development of hypotheses about ecological phenomena and dominant landscape elements associated with these phenomena, how these phenomena are structured in space, and how they function in a hierarchy. Success in addressing the research recommendations outlined in this article cannot occur within an ad hoc, largely uncoordinated research environment. Successful implementation of this plan will require activities--coordination, funding, and education--that are both scientific and administrative in nature. Perhaps the most important element of an infrastructure to support the scientific work of ecoregionalization would be a national or international authority similar to the Water and Science Technology Board of the National Academy of Sciences.

Environmental Management↗

Egg size variation among tropical and temperate songbirds: An embryonic temperature hypothesis

Species with 'slow' life history strategies (long life, low fecundity) are thought to produce high-quality offspring by investing in larger, but fewer, young. Larger eggs are indeed associated with fewer eggs across taxa and can yield higher-quality offspring. Tropical passerines appear to follow theory because they commonly exhibit slow life history strategies and produce larger, but fewer, eggs compared with northern species. Yet, I show here that relative egg mass (corrected for adult mass) varies extensively in the tropics and subtropics for the same clutch size, and this variation is unexplained. I propose a hypothesis to explain egg size variation both within the tropics and between latitudes: Relative egg mass increases in species with cooler egg temperatures and longer embryonic periods to offset associated increases in energetic requirements of embryos. Egg temperatures of birds are determined by parental incubation behavior and are often cooler among tropical passerines because of reduced parental attentiveness of eggs. Here, I show that cooler egg temperatures and longer embryonic periods explained the enigmatic variation in egg mass within and among regions, based on field studies in tropical Venezuela (36 species), subtropical Argentina (16 species), and north temperate Arizona (20 species). Alternative explanations are not supported. Thus, large egg sizes may reflect compensation for increased energetic requirements of cool egg temperatures and long embryonic periods that result from reduced parental attentiveness in tropical birds. ?? 2008 by The National Academy of Sciences of the USA.

Proceedings of the National Academy of Sciences of↗

Research in the United States relative to geochemistry and health

Increasing concern regarding the effects of the geochemical environment on health in the United States has fostered research studies in a number of universities and government agencies. The necessity to evaluate the effects of natural and man-made elemental excesses in the environment on health requires the establishment of requirements and tolerance limits for the various elements in water and crops. Maps of the geographic distribution of these elements in rocks, surficial materials and ground and surface waters are also essential for comparison with the occurrence of disease. Funding support for research projects that relate to various parameters of these problems emanates largely from a few federal agencies, and much of the work is conducted at government, university and private facilities. An example of the latter is the National Academy of Sciences-National Research Council, which has several components that are addressing a variety of comparative studies of the geochemical environment related to health; studies involve specific trace elements (like selenium and magnesium), diseases (like cancer, urolithiasis and cardiovascular disease), other health factors (like aging and nutrition) and links with timely major problems (like the health effects of greatly increasing the use of coal).

Physics and Chemistry of the Earth↗

Discovery of a unique Ig heavy-chain (IgT) in rainbow trout: Implications for a distinctive B cell developmental pathway in teleost fish

During the analysis of Ig superfamily members within the available rainbow trout (Oncorhynchus mykiss) EST gene index, we identified a unique Ig heavy-chain (IgH) isotype. cDNAs encoding this isotype are composed of a typical IgH leader sequence and a VDJ rearranged segment followed by four Ig superfamily C-1 domains represented as either membrane-bound or secretory versions. Because teleost fish were previously thought to encode and express only two IgH isotypes (IgM and IgD) for their humoral immune repertoire, we isolated all three cDNA isotypes from a single homozygous trout (OSU-142) to confirm that all three are indeed independent isotypes. Bioinformatic and phylogenetic analysis indicates that this previously undescribed divergent isotype is restricted to bony fish, thus we have named this isotype "IgT" (??) for teleost fish. Genomic sequence analysis of an OSU-142 bacterial artificial chromosome (BAC) clone positive for all three IgH isotypes revealed that IgT utilizes the standard rainbow trout VH families, but surprisingly, the IgT isotype possesses its own exclusive set of DH and JH elements for the generation of diversity. The IgT D and J segments and ?? constant (C) region genes are located upstream of the D and J elements for IgM, representing a genomic IgH architecture that has not been observed in any other vertebrate class. All three isotypes are primarily expressed in the spleen and pronephros (bone marrow equivalent), and ontogenically, expression of IgT is present 4 d before hatching in developing embryos. ?? 2005 by The National Academy of Sciences of the USA.

Proceedings of the National Academy of Sciences of↗

Food limitation leads to behavioral diversification and dietary specialization in sea otters

Dietary diversity often varies inversely with prey resource abundance. This pattern, although typically measured at the population level, is usually assumed to also characterize the behavior of individual animals within the population. However, the pattern might also be produced by changes in the degree of variation among individuals. Here we report on dietary and associated behavioral changes that occurred with the experimental translocation of sea otters from a food-poor to a food-rich environment. Although the diets of all individuals were broadly similar in the food-rich environment, a behaviorally based dietary polymorphism existed in the food-poor environment. Higher dietary diversity under low resource abundance was largely driven by greater variation among individuals. We further show that the dietary polymorphism in the food-poor environment included a broad suite of correlated behavioral variables and that the individuals that comprised specific behavioral clusters benefited from improved foraging efficiency on their individually preferred prey. Our findings add to the growing list of examples of extreme individuality in behavior and prey choice within populations and suggest that this phenomenon can emerge as a behavioral manifestation of increased population density. Individuality in foraging behavior adds complexity to both the fitness consequences of prey selection and food web dynamics, and it may figure prominently as a diversifying process over evolutionary timescales. ?? 2008 by The National Academy of Sciences of the USA.

Proceedings of the National Academy of Sciences of↗

An introduced and a native vertebrate hybridize to form a genetic bridge to a second native species

The genetic impacts of hybridization between native and introduced species are of considerable conservation concern, while the possibility of reticulate evolution affects our basic understanding of how species arise and shapes how we use genetic data to understand evolutionary diversification. By using mitochondrial NADH dehydrogenase subunit 2 (ND2) sequences and 467 amplified fragment-length polymorphism nuclear DNA markers, we show that the introduced white sucker (Catostomus commersoni) has hybridized with two species native to the Colorado River Basin - the flannelmouth sucker (Catostomus latipinnis) and the bluehead sucker (Catostomus discobolus). Hybrids between the flannelmouth sucker and white sucker have facilitated introgression between the two native species, previously isolated by reproductive barriers, such that individuals exist with contributions from all three genomes. Most hybrids had the mitochondrial haplotype of the introduced white sucker, emphasizing its pivotal role in this three-way hybridization. Our findings highlight how introduced species can threaten the genetic integrity of not only one species but also multiple previously reproductively isolated species. Furthermore, this complex three-way reticulate (as opposed to strictly bifurcating) evolution suggests that seeking examples in other vertebrate systems might be productive. Although the present study involved an introduced species, similar patterns of hybridization could result from natural processes, including stream capture or geological formations (e.g., the Bering land bridge). ?? 2008 by The National Academy of Sciences of the USA.

Proceedings of the National Academy of Sciences of↗

Abrupt climate change and collapse of deep-sea ecosystems

We investigated the deep-sea fossil record of benthic ostracodes during periods of rapid climate and oceanographic change over the past 20,000 years in a core from intermediate depth in the northwestern Atlantic. Results show that deep-sea benthic community "collapses" occur with faunal turnover of up to 50% during major climatically driven oceanographic changes. Species diversity as measured by the Shannon-Wiener index falls from 3 to as low as 1.6 during these events. Major disruptions in the benthic communities commenced with Heinrich Event 1, the Inter-Aller??d Cold Period (IACP: 13.1 ka), the Younger Dryas (YD: 12.9-11.5 ka), and several Holocene Bond events when changes in deep-water circulation occurred. The largest collapse is associated with the YD/IACP and is characterized by an abrupt two-step decrease in both the upper North Atlantic Deep Water assemblage and species diversity at 13.1 ka and at 12.2 ka. The ostracode fauna at this site did not fully recover until ???8 ka, with the establishment of Labrador Sea Water ventilation. Ecologically opportunistic slope species prospered during this community collapse. Other abrupt community collapses during the past 20 ka generally correspond to millennial climate events. These results indicate that deep-sea ecosystems are not immune to the effects of rapid climate changes occurring over centuries or less. ?? 2008 by The National Academy of Sciences of the USA.

Proceedings of the National Academy of Sciences of↗

Competitive release and facilitation of drug-resistant parasites after therapeutic chemotherapy in a rodent malaria model

Malaria infections frequently consist of mixtures of drug-resistant and drug-sensitive parasites. If crowding occurs, where clonal population densities are suppressed by the presence of coinfecting clones, removal of susceptible clones by drug treatment could allow resistant clones to expand into the newly vacated niche space within a host. Theoretical models show that, if such competitive release occurs, it can be a potent contributor to the strength of selection, greatly accelerating the rate at which resistance spreads in a population. A variety of correlational field data suggest that competitive release could occur in human malaria populations, but direct evidence cannot be ethically obtained from human infections. Here we show competitive release after pyrimethamine curative chemotherapy of acute infections of the rodent malaria Plasmodium chabaudi in laboratory mice. The expansion of resistant parasite numbers after treatment resulted in enhanced transmission-stage densities. After the elimination or near-elimination of sensitive parasites, the number of resistant parasites increased beyond that achieved when a competitor had never been present. Thus, a substantial competitive release occurred, markedly elevating the fitness advantages of drug resistance above those arising from survival alone. This finding may explain the rapid spread of drug resistance and the subsequently brief useful lifespans of some antimalarial drugs. In a second experiment, where subcurative chemotherapy was administered, the resistant clone was only partly released from competitive suppression and experienced a restriction in the size of its expansion after treatment. This finding raises the prospect of harnessing in-host ecology to slow the spread of drug resistance. ?? 2007 by The National Academy of Sciences of the USA.

Proceedings of the National Academy of Sciences of↗

Aquatic insect ecophysiological traits reveal phylogenetically based differences in dissolved cadmium susceptibility

We used a phylogenetically based comparative approach to evaluate the potential for physiological studies to reveal patterns of diversity in traits related to susceptibility to an environmental stressor, the trace metal cadmium (Cd). Physiological traits related to Cd bioaccumulation, compartmentalization, and ultimately susceptibility were measured in 21 aquatic insect species representing the orders Ephemeroptera, Plecoptera, and Trichoptera. We mapped these experimentally derived physiological traits onto a phylogeny and quantified the tendency for related species to be similar (phylogenetic signal). All traits related to Cd bioaccumulation and susceptibility exhibited statistically significant phylogenetic signal, although the signal strength varied among traits. Conventional and phylogenetically based regression models were compared, revealing great variability within orders but consistent, strong differences among insect families. Uptake and elimination rate constants were positively correlated among species, but only when effects of body size and phylogeny were incorporated in the analysis. Together, uptake and elimination rates predicted dramatic Cd bioaccumulation differences among species that agreed with field-based measurements. We discovered a potential tradeoff between the ability to eliminate Cd and the ability to detoxify it across species, particularly mayflies. The best-fit regression models were driven by phylogenetic parameters (especially differences among families) rather than functional traits, suggesting that it may eventually be possible to predict a taxon's physiological performance based on its phylogenetic position, provided adequate physiological information is available for close relatives. There appears to be great potential for evolutionary physiological approaches to augment our understanding of insect responses to environmental stressors in nature. ?? 2008 by The National Academy of Sciences of the USA.

Proceedings of the National Academy of Sciences of↗

Geographic Research in the USGS Western Region

The two geography research programs of the U.S. Geological Survey (USGS), Land Remote Sensing and Geographic Analysis and Monitoring, have very strong relevance to the USGS mission and science strategy. In the western United States, the particular niche of these geography programs is in connecting USGS science to people and communities. Reports from the National Academy of Sciences and other organizations invariably encourage the U.S. Geological Survey (USGS) to ensure the quality of its science while finding ways to make it more relevant to important societal issues. Much of the geography research conducted in the USGS Western Region does exactly that. In Menlo Park, California, the geography research team is focused on developing tools and techniques to help people assess risk from natural hazards and environmental impacts. In Flagstaff and Tucson, Arizona, geography scientists explore new ways to use remote sensing to help communities deal with environmental issues.

Fact Sheet↗

Research and development program; Conservation Division Outer Continental Shelf oil and gas operations; Technical report 1981

As a result of recommendations several years ago from the National Academy of Sciences, the University of Oklahoma, and the National Aeronautics and Space Administration (NASA), the U.S. Geological Survey has embarked upon a program of research and development to provide the technological insights needed for its regulatory operations offshore—operations which provide assurances to the public for safety and for the prevention of pollution in oil and gas drilling and production. These clear objectives are, therefore, those of the research program, not the economics of operations, which are of concern to industry. The Program is a contract research program and is an integral part of the Conservation Division. It is a focal point for deriving possible solutions from the university community, private industry, and the Federal laboratory system for identified offshore operational problems. This vast interdisciplinary body of science and technology provides the kind of research needed by the Division in its Outer Continental Shelf (OCS) operations which involve such problematic areas as structural dynamics, fluid flow, and geotechnology. The Program encourages innovation and creativity which can be accomplished only by talented scientists and engineers who are dedicated to man's endeavor to make breakthroughs in science and technology. Because the Division's mission is operational, in a sense like the U.S. Navy, the R&D Program must progress in a timely manner even though technological advances cannot really be scheduled. As the Navy's Office of Naval Research (ONR) has so successfully coped with the seeming dichotomy of anticipating the occurrence of innovations, so must the Conservation Division. Thus, USGS, like ONR, makes use of the unsolicited proposal and the best effort contract to accomplish its objectives. Good science and technology can only be accomplished when several variables converge: a talented investigator doing his own research, which happens to coincide with our needs, availability of resources, time scales, etc. Our task is to "beat the bushes" so that these people come to us with their innovative ideas and concepts. They have done so by learning of our interests through announcements of the Program in the Federal Register, reading reports which emanate from the Program, and by attending our seminars.

Open-File Report↗

Geochemical proxies of North American freshwater routing during the Younger Dryas cold event

The Younger Dryas cold interval represents a time when much of the Northern Hemisphere cooled from ???12.9 to 11.5 kiloyears B.P. The cause of this event, which has long been viewed as the canonical example of abrupt climate change, was initially attributed to the routing of freshwater to the St. Lawrence River with an attendant reduction in Atlantic meridional overturning circulation. However, this mechanism has recently been questioned because current proxies and dating techniques have been unable to confirm that eastward routing with an increase in freshwater flux occurred during the Younger Dryas. Here we use new geochemical proxies (??Mg/Ca, U/Ca, and 87Sr/86Sr) measured in planktonic foraminifera at the mouth of the St. Lawrence estuary as tracers of freshwater sources to further evaluate this question. Our proxies, combined with planktonic ??18Oseawater and ??13C, confirm that routing of runoff from western Canada to the St. Lawrence River occurred at the start of the Younger Dryas, with an attendant increase in freshwater flux of 0.06 ?? 0.02 Sverdrup (1 Sverdrup = 106 m3??s-1). This base discharge increase is sufficient to have reduced Atlantic meridional overturning circulation and caused the Younger Dryas cold interval. In addition, our data indicate subsequent fluctuations in the freshwater flux to the St. Lawrence River of ???0.06-0.12 Sverdrup, thus explaining the variability in the overturning circulation and climate during the Younger Dryas. ?? 2007 by The National Academy of Sciences of the USA.

Proceedings of the National Academy of Sciences of↗

Regulating continent growth and composition by chemical weathering

Continents ride high above the ocean floor because they are underlain by thick, low-density, Si-rich, and Mg-poor crust. However, the parental magmas of continents were basaltic, which means they must have lost Mg relative to Si during their maturation into continents. Igneous differentiation followed by lower crustal delamination and chemical weathering followed by subduction recycling are possible solutions, but the relative magnitudes of each process have never been quantitatively constrained because of the lack of appropriate data. Here, we show that the relative contributions of these processes can be obtained by simultaneous examination of Mg and Li (an analog for Mg) on the regional and global scales in arcs, delaminated lower crust, and river waters. At least 20% of Mg is lost from continents by weathering, which translates into >20% of continental mass lost by weathering (40% by delamination). Chemical weathering leaves behind a more Si-rich and Mg-poor crust, which is less dense and hence decreases the probability of crustal recycling by subduction. Net continental growth is thus modulated by chemical weathering and likely influenced by secular changes in weathering mechanisms. ?? 2008 by The National Academy of Sciences of the USA.

Proceedings of the National Academy of Sciences of↗

The evolution of vertebrate Toll-like receptors

The complete sequences of Takifugu Toll-like receptor (TLR) loci and gene predictions from many draft genomes enable comprehensive molecular phylogenetic analysis. Strong selective pressure for recognition of and response to pathogen-associated molecular patterns has maintained a largely unchanging TLR recognition in all vertebrates. There are six major families of vertebrate TLRs. This repertoire is distinct from that of invertebrates. TLRs within a family recognize a general class of pathogen-associated molecular patterns. Most vertebrates have exactly one gene ortholog for each TLR family. The family including TLR1 has more species-specific adaptations than other families. A major family including TLR11 is represented in humans only by a pseudogene. Coincidental evolution plays a minor role in TLR evolution. The sequencing phase of this study produced finished genomic sequences for the 12 Takifugu rubripes TLRs. In addition, we have produced > 70 gene models, including sequences from the opossum, chicken, frog, dog, sea urchin, and sea squirt. ?? 2005 by The National Academy of Sciences of the USA.

Proceedings of the National Academy of Sciences of↗

Increased salinization of fresh water in the Northeastern United States

Chloride concentrations are increasing at a rate that threatens the availability of fresh water in the northeastern United States. Increases in roadways and deicer use are now salinizing fresh waters, degrading habitat for aquatic organisms, and impacting large supplies of drinking water for humans throughout the region. We observed chloride concentrations of up to 25% of the concentration of seawater in streams of Maryland, New York, and New Hampshire during winters, and chloride concentrations remaining up to 100 times greater than unimpacted forest streams during summers. Mean annual chloride concentration increased as a function of impervious surface and exceeded tolerance for freshwater life in suburban and urban watersheds. Our analysis shows that if salinity were to continue to increase at its present rate due to changes in impervious surface coverage and current management practices, many surface waters in the northeastern United States would not be potable for human consumption and would become toxic to freshwater life within the next century. ?? 2005 by The National Academy of Sciences of the USA.

Proceedings of the National Academy of Sciences of↗

Warming of the Indian Ocean threatens eastern and southern African food security but could be mitigated by agricultural development

Since 1980, the number of undernourished people in eastern and southern Africa has more than doubled. Rural development stalled and rural poverty expanded during the 1990s. Population growth remains very high, and declining per-capita agricultural capacity retards progress toward Millennium Development goals. Analyses of in situ station data and satellite observations of precipitation have identified another problematic trend: main growing-season rainfall receipts have diminished by ???15% in food-insecure countries clustered along the western rim of the Indian Ocean. Occurring during the main growing seasons in poor countries dependent on rain-fed agriculture, these declines are societally dangerous. Will they persist or intensify? Tracing moisture deficits upstream to an anthropogenically warming Indian Ocean leads us to conclude that further rainfall declines are likely. We present analyses suggesting that warming in the central Indian Ocean disrupts onshore moisture transports, reducing continental rainfall. Thus, late 20th-century anthropogenic Indian Ocean warming has probably already produced societally dangerous climate change by creating drought and social disruption in some of the world's most fragile food economies. We quantify the potential impacts of the observed precipitation and agricultural capacity trends by modeling 'millions of undernourished people' as a function of rainfall, population, cultivated area, seed, and fertilizer use. Persistence of current tendencies may result in a 50% increase in undernourished people by 2030. On the other hand, modest increases in per-capita agricultural productivity could more than offset the observed precipitation declines. Investing in agricultural development can help mitigate climate change while decreasing rural poverty and vulnerability. ?? 2008 by The National Academy of Sciences of the USA.

Proceedings of the National Academy of Sciences of↗

Soil nutrients influence spatial distributions of tropical trees species

The importance of niche vs. neutral assembly mechanisms in structuring tropical tree communities remains an important unsettled question in community ecology [Bell G (2005) Ecology 86:1757-1770]. There is ample evidence that species distributions are determined by soils and habitat factors at landscape (<104 km2) and regional scales. At local scales (<1 km2), however, habitat factors and species distributions show comparable spatial aggregation, making it difficult to disentangle the importance of niche and dispersal processes. In this article, we test soil resource-based niche assembly at a local scale, using species and soil nutrient distributions obtained at high spatial resolution in three diverse neotropical forest plots in Colombia (La Planada), Ecuador (Yasuni), and Panama (Barro Colorado Island). Using spatial distribution maps of >0.5 million individual trees of 1,400 species and 10 essential plant nutrients, we used Monte Carlo simulations of species distributions to test plant-soil associations against null expectations based on dispersal assembly. We found that the spatial distributions of 36-51% of tree species at these sites show strong associations to soil nutrient distributions. Neutral dispersal assembly cannot account for these plant-soil associations or the observed niche breadths of these species. These results indicate that belowground resource availability plays an important role in the assembly of tropical tree communities at local scales and provide the basis for future investigations on the mechanisms of resource competition among tropical tree species. ?? 2007 by The National Academy of Sciences of the USA.

Proceedings of the National Academy of Sciences of↗

U.S. Geological Survey scientific activities in the exploration of Antarctica: 2002-03 field season

The U.S. Geological Survey (USGS) mapping program in Antarctica is one of the longest continuously funded projects in the United States Antarctic Program (USAP). This is the 53rd U.S. expedition to Antarctica in which USGS scientists have participated. The financial support from the National Science Foundation, which extends back to the time of the International Geophysical Year (IGY) in 1956–57, can be attributed to the need for accurate maps of specific field areas or regions where NSF-funded science projects were planned. The epoch of Antarctic exploration during the IGY was being driven by science, and, in a spirit of peaceful cooperation, the international scientific community wanted to limit military activities on the continent to logistical support. The USGS, a Federal civilian science agency in the Department of the Interior, had, since its founding in 1879, carried out numerous field-based national (and some international) programs in biology, geology, hydrology, and mapping. Therefore, the USGS was the obvious choice for these tasks, because it already had a professional staff of experienced mapmakers and program managers with the foresight, dedication, and understanding of the need for accurate maps to support the science programs in Antarctica when asked to do so by the U.S. National Academy of Sciences. Public Laws 85-743 and 87-626, signed in August 1958 and in September 1962, respectively, authorized the Secretary, U.S. Department of the Interior, through the USGS, to support mapping and scientific work in Antarctica. The USGS mapping and science programs still play a significant role in the advancement of science in Antarctica today. Antarctica is the planet's 5th largest continent [13.2 million km 2 (5.1 million mi 2 )], it contains the world's largest (of two) remaining ice sheets, and it is considered to be one of the most important scientific laboratories on Earth. This report provides documentation of USGS scientific activities in the exploration of Antarctica during the 2002–03 field season (Mullins, 2002).

Open-File Report↗