Intrusive volcanic rocks in western Pacific forearcs
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We elaborate on the well-known spatial association between axc volcanoes and Wadati Benioff zones and explore in detail their genetic relationships as dual physical expressions of slab metamorphism of the oceanic crust and uppermost mantle. At hypocentral depths less than 200 km intra slab Wadati-Benioff earthquakes tend to occur near the top surfaces of slabs. Subduction of very young lithosphere (age < 15-25 Ma) with high heat flow (> 75 mW/m 2) produces mainly shallow earthquakes and spaxse or absent arc volcanism. Subduction of older crust with normal heat flow (50-65 mW/m 2) produces markedly deeper intraslab earthquakes and generally normal volcanic vigor. Seismological observations show that the low-seismic-velocity gabbroic mineralogy of the crust may persist to depths of as much as 150 km in old, cold lithosphere but only to depths of 50-60 km in young, warm slabs. Metamorphic processes in the crust and shallow upper mantle of subducting slabs and the reactivation of faults originally created at shallow depths in the ocean basins probably control the occurrence of intraslab earthquakes to depths of as much as 350 kin. A conceptual model for this metamorphism incorporates the likely effects of water liberated by dehydration. Such dehydration facilitates both brittle faulting by fault reactivation and promotes the kinetics of the transformation of the anhydrous gabbro component of the crust to eclogite. Finite-element modelling shows that densification to eclogite is expected to produce extensional stresses in transformed crust and a smaller compression in the underlying mantle. This model helps explain why most intermediate-depth intraslab earthquakes occur just below the top surfaces of slabs and why many have focal mechanisms indicating down-dip extension. Young, warm slabs have mostly shallow intraslab earthquakes and sparse axc volcanoes because dehydration and eclogite formation largely cease before such slabs axe in contact with asthenosphere. These processes are evidently delayed by kinetic hindrance and the high pressure stability of hydrous phazes at low temperatures in older, colder subducting crust, and thus earthquake activity and asthenospheric-wedge melting tend to be focused at depths of 100-170 kin. Anomalous behavior correlated with the subduction of island-and seamount chains appears to be associated with anomalou shallow intraplate faulting and with perturbations of slab metamorphism by these chains.
The Canada Basin (CB) formed during a short period of seafloor spreading inferred to be Early Cretaceous in age. Brookian strata of inferred Early Cretaceous–Holocene age comprise the sedimentary fill of the Canada Basin Tectono-Sedimentary Element (CB TSE). Although the CB has remained tectonically quiet since seafloor spreading ceased, both proximal and distal tectonism (Alpha Ridge magmatism, and the Cordilleran, Brooks Range and Eurekan orogenies) have influenced sediment source areas, dispersal paths and thicknesses in the basin. In the Neogene, the dominant source of sediments was the Mackenzie River, which drains northern portions of the Cordilleran orogen. The CB TSE is one of the most remote and challenging places on Earth to explore. Although regional seismic reflection and refraction data exist, there are no boreholes to constrain interpretations. Existing published estimates of hydrocarbon potential range from limited to moderate to significant.
The first part of this synthesis summarizes the morphology of the jelly layers surrounding an amphibian ovum. We propose a standard terminology and discuss the evolution of jelly layers. The second part reviews the morphological diversity and arrangement of deposited eggs?the ovipositional mode; we recognize 5 morphological classes including 14 modes. We discuss some of the oviductal, ovipositional, and postovipositional events that contribute to these morphologies. We have incorporated data from taxa from throughout the world but recognize that other types will be discovered that may modify understanding of these modes. Finally, we discuss the evolutionary context of the diversity of clutch structure and present a first estimate of its evolution.
Numerous factors have contributed to declines in populations of the federally threatened Agassiz's Desert Tortoise ( Gopherus agassizii ) and continue to limit recovery. In 2010, we surveyed a low-density population on a military test facility in the northwestern Mojave Desert of California, USA, to evaluate population status and identify potential factors contributing to distribution and low densities. Estimated densities of live tortoises ranged spatially from 1.2/km 2 to 15.1/km 2 . Although only one death of a breeding-age tortoise was recorded for the 4-yr period prior to the survey, remains of 16 juvenile and immature tortoises were found, and most showed signs of predation by Common Ravens ( Corvus corax ) and mammals. Predation may have limited recruitment of young tortoises into the adult size classes. To evaluate the relative importance of different types of impacts to tortoises, we developed predictive models for spatially explicit densities of tortoise sign and live tortoises using topography (i.e., slope), predators (Common Raven, signs of mammalian predators), and anthropogenic impacts (distances from paved road and denuded areas, density of ordnance fragments) as covariates. Models suggest that densities of tortoise sign increased with slope and signs of mammalian predators and decreased with Common Ravens, while also varying based on interaction effects involving these predictors as well as distances from paved roads, denuded areas, and ordnance. Similarly, densities of live tortoises varied by interaction effects among distances to denuded areas and paved roads, density of ordnance fragments, and slope. Thus multiple factors predict the densities and distribution of this population.
We surveyed an area of ∼260 km 2 in the western Mojave Desert to evaluate relationships between condition of Agassiz's Desert Tortoise populations ( Gopherus agassizii ) and habitat on lands that have experienced three different levels of management and protection. We established 240 1-ha plots using random sampling, with 80 plots on each of the three types of managed lands. We conducted surveys in spring 2011 and collected data on live tortoises, shell-skeletal remains, other signs of tortoises, perennial vegetation, predators, and evidence of human use. Throughout the study area and regardless of management area, tortoise abundance was positively associated with one of the more diverse associations of perennial vegetation. The management area with the longest history of protection, a fence, and legal exclusion of livestock and vehicles had significantly more live tortoises and lower death rates than the other two areas. Tortoise presence and abundance in this protected area had no significant positive or negative associations with predators or human-related impacts. In contrast, the management area with a more recent exclusion of livestock, limited vehicular traffic, and with a recent, partial fence had lower tortoise densities and high death rates. Tortoise abundance here was negatively associated with vehicle tracks and positively associated with mammalian predators and debris from firearms. The management area with the least protection—unfenced, with uncontrolled vehicle use, sheep grazing, and high trash counts—also had low tortoise densities and high death rates. Tortoise abundance was negatively associated with sheep grazing and positively associated with trash and mammalian predator scat. cat.
Eastern Kingsnakes ( Lampropeltis getula ) are an important component and predator in herpetofaunal communities, but many Eastern Kingsnake populations have declined precipitously in the last few decades, particularly in the southeastern United States. Here, we describe an intensive capture–mark–recapture study of L. getula conducted during 1974–1978 in a canal bank–Water Hyacinth ( Eichhornia crassipes ) community at Rainey Slough in southern Florida, where annual capture probabilities of adults ranged from 0.662–0.787. Population size and structure, seasonal activity, movements, microhabitat use, behavior, thermal ecology, and predator–prey relationships are described. At this site kingsnakes were susceptible to capture mostly in winter and spring, were diurnal, used rodent ( Sigmodon hispidus ) burrows on canal banks as nocturnal retreats, and emerged from burrows on 13–26% of the sampling days. Overlap of burrow use by both sexes was extensive with no evidence of territoriality. Kingsnakes readily entered the Water Hyacinths to bask, pursue mates, and forage. At Rainey Slough only snakes were detected in the diet of kingsnakes. Concurrent sampling of potential snake prey in the hyacinths and on canal banks revealed 10 species that varied in use of the two sampled habitats and in body size. A range-wide analysis confirmed that in descending order snakes, reptile eggs, and lizards dominate the diet of L. getula in Florida (94.8%) and remain important prey types elsewhere (80.2%). At Rainey Slough the density of six species of semiaquatic snakes in Water Hyacinths averaged 3534 individuals/ha with a mean annual biomass of 135.8 kg/ha, and kingsnake biomass was only 2.2–3.9% of prey snake biomass. We estimated that the kingsnake population consumed 36.82–63.58 kg/yr, or about 10.0–17.2% of the standing crop of snakes in the Water Hyacinth community. Adult male L. getula lost on average 39.3% of their body mass associated with the spring reproductive season, whereas females lost only 3.4% in the same period. Body condition indices for both sexes improved substantially thereafter. In follow-up surveys at Rainey Slough during 2006–2010 no kingsnakes were found. Semiaquatic snake densities in the Water Hyacinths were 77.2% lower (807.4/ha) than in the 1970s and consisted of only three species. Compared to the enigmatic declines and extirpation of L. getula populations elsewhere, at Rainey Slough the primary cause likely was unsustainable mortality from road reconstruction and paving in the winter–spring of 1979 and subsequent roadkill. Other potentially causative agents of extirpation of L. getula in this system are discussed.
Our study represents the first attempt to describe biogeographic provinces for North American (México, United States, and Canada) turtles. We analyzed three nested data sets separately: (1) all turtles, (2) freshwater turtles, and (3) aquatic turtles. We georeferenced North American turtle distributions, then we created presence–absence matrices for each of the three data sets. We used watershed unit as biogeographic units. We conducted an unweighted pair-group method with arithmetic mean clustering analysis on each Jaccard index distance matrix from our watershed species matrices to delineate biogeographic provinces. Provinces were then tested for significant differences in species compositions in a global model with the use of a one-way analysis of similarity. We conducted a best subset of environmental variables with maximum (rank) correlation with community dissimilarities that determined the best model of abiotic variables explaining province delineation (i.e., climate, topography, and stream channel). To identify which species contributed the most to province delineations, we conducted an indicator species analysis and a similarity-percentage analysis. There were 16 all-turtle provinces, 15 freshwater provinces, and 13 aquatic provinces. Species compositions delineating the provinces were explained by abiotic variables, including mean annual precipitation, mean precipitation seasonality, and diversity of streams. Province delineations correspond closely with geographical boundaries, many of which have Pleistocene origins. For example, rivers with a history of carrying glacial runoff (e.g., Arkansas, Mississippi) sometimes dissect upland provinces, especially for aquatic and semiaquatic turtles. Compared with freshwater fishes, turtles show greater sensitivity to decreased temperature with restriction of most taxa south of the last permafrost maximum. Turtles also exhibit higher sensitivity to climatic, geomorphic, and tectonic instability, with richness and endemism concentrated along the more stable Gulf of México and Atlantic (south of the last permafrost maximum) coasts. Although distribution data indicate two aquatic turtles are most cold tolerant (i.e., Chrysemys picta , Chelydra serpentina ), aquatic turtles overall show the most restriction to warmer, wetter climates. Sequential addition of semiaquatic and terrestrial turtles into analyses shows, as expected, that these taxa flesh out turtle faunas in climatically harsh (e.g., grasslands) or remote (e.g., California, Sonoran Desert) regions. The turtle assemblages of southwestern versus southeastern North America are distinct. But there is a transition zone across the semiarid plains of the Texas Gulf Coast, High Plains, and Chihuahuan Desert, including a strong boundary congruent with the Cochise Filter-Barrier. This is not a simple subdivision of Neotropical versus Nearctic taxa, as some lineages from both realms span the transition zone.
During a mark–recapture study of Townsend's ground squirrels ( Spermophilus townsendii ) on 20 sites in the Snake River Birds of Prey National Conservation Area, Idaho, in 1991 through 1994, 4407 animals were marked in 17639 capture events. This study of differences in population dynamics of Townsend's ground squirrels among habitats spanned a drought near the extreme of the 130-yr record, followed by prolonged winter conditions. Townsend's ground squirrels have a short active season (≈4 mo) in which to reproduce and store fat for overwintering. Their food consists largely of succulent grasses and forbs in this dry shrubsteppe and grassland habitat. The drought in the latter half of the 1992 active season produced early drying of Sandberg's bluegrass ( Poa secunda ) and was associated with low adult and juvenile body masses prior to immergence into estivation/hibernation. The following prolonged winter was associated with late emergence of females in 1993. Early-season body masses of adults were low in 1993 relative to 1992, whereas percentage of body fat in males was relatively high. These weather patterns in spring 1992 and winter 1993 also resulted in reduced adult persistence through the ≈7-mo inactive period, especially for adult females, and near-zero persistence of >1200 juveniles. Consequently, densities of Townsend's ground squirrels across the 20 livetrap sites declined. The demographic effects of drought and prolonged winter lasted at least through the subsequent breeding season. Adult females that survived these weather extremes produced fewer emergent young per female than did adult females prior to the event. Prior to the drought/prolonged winter, yearling female body masses were higher than, or indistinguishable from, those of adults. Females produced in 1993 had lower body masses as yearlings than did adult females. Demographic response to the drought and prolonged winter varied with habitat; ground squirrels in sagebrush habitat showed less decline in persistence and density and produced more young per female during the next active season following the drought (1993) than did ground squirrels in grassland habitat, where densities had been significantly higher prior to the drought and prolonged winter. Studies involving habitat comparisons of animal demography should always be placed in the context of long-term weather patterns, because habitat quality rankings based on density, reproduction, and survival may differ with environmental conditions. Physiological effects of environmental “crunches” on consumers may persist beyond the period of influence on food resources, reducing reproductive success and growth rates of future offspring.
We assessed the relative importance of environmental variation, interspecific competition for space, and predator abundance on assemblage structure and microhabitat use in a stream fish assemblage inhabiting Coweeta Creek, North Carolina, USA. Our study encompassed a 10–yr time span (1983–1992) and included some of the highest and lowest flows in the last 58 years. We collected 16 seasonal samples which included data on: (1) habitat availability (total and microhabitat) and microhabitat diversity, (2) assemblage structure (i.e., the number and abundances of species comprising a subset of the community), and (3) microhabitat use and overlap. We classified habitat availability data on the basis of year, season, and hydrologic period. Hydrologic period (i.e., pre–drought [PR], drought [D], and post–drought [PO]) represented the temporal location of a sample with respect to a four–year drought that occurred during the study. Hydrologic period explained a greater amount of variance in habitat availability data than either season or year. Total habitat availability was significantly greater during PO than in PR or D, although microhabitat diversity did not differ among either seasons or hydrologic periods. There were significantly fewer high–flow events (i.e., ≥2.1 m 3 /s) during D than in either PR or PO periods. We observed a total of 16 species during our investigation, and the total number of species was significantly higher in D than in PR samples. Correlation analyses between the number of species present (total and abundant species) and environmental data yielded limited results, although the total number of species was inversely correlated with total habitat availability. A cluster analysis grouped assemblage structure samples by hydrologic period rather than season or year, supporting the contention that variation in annual flow had a strong impact on this assemblage. The drought had little effect on the numerical abundance of benthic species in this assemblage; however, a majority of water–column species increased in abundance. The increased abundances of water–column species may have been related to the decrease in high-flow events observed during the drought. Such high–flow events are known to cause mortality in stream fishes. Microhabitat use data showed that species belonged to one of three microhabitat guilds: benthic, lower water column, and mid water column. In general, species within the same guild did not exhibit statistically distinguishable patterns of microhabitat use, and most significant differences occurred between members of different guilds. However, lower water–column guild species frequentlywere not separable from all members of either benthic or mid–water–column species. Variations in the abundance of potential competitors or predators did not produce strong shifts in microhabitat use by assemblage members. Predators were present in the site in only 9 of 16 seasonal samples and never were abundant (maximum number observed per day was 2). In conclusion, our results demonstrate that variability in both mean and peak flows had a much stronger effect on the structure and use of spatial resources within this assemblage then either interspecific competition for space or predation. Consequently, we suspect that the patterns in both assemblage structure and resource use displayed by fishes in Coweeta Creek arose from the interaction between environmental variation and species–specific evolutionary constraints on behavior, morphology, and physiology.
Monte Carlo simulations were conducted to evaluate robustness of four tests to detect density dependence, from series of population abundances, to the addition of sampling variance. Population abundances were generated from random walk, stochastic exponential growth, and density-dependent population models. Population abundance estimates were generated with sampling variances distributed as lognormal and constant coefficients of variation ( cv ) from 0.00 to 1.00. In general, when data were generated under a random walk, Type I error rates increased rapidly for Bulmer's R, Pollard et al.'s, and Dennis and Taper's tests with increasing magnitude of sampling variance for n > 5 yr and all values of process variation. Bulmer's R * test maintained a constant 5% Type I error rate for n > 5 yr and all magnitudes of sampling variance in the population abundance estimates. When abundances were generated from two stochastic exponential growth models ( R = 0.05 and R = 0.10), Type I errors again increased with increasing sampling variance; magnitude of Type I error rates were higher for the slower growing population. Therefore, sampling error inflated Type I error rates, invalidating the tests, for all except Bulmer's R * test. Comparable simulations for abundance estimates generated from a density-dependent growth rate model were conducted to estimate power of the tests. Type II error rates were influenced by the relationship of initial population size to carrying capacity ( K ), length of time series, as well as sampling error. Given the inflated Type I error rates for all but Bulmer's R *, power was overestimated for the remaining tests, resulting in density dependence being detected more often than it existed. Population abundances of natural populations are almost exclusively estimated rather than censused, assuring sampling error. Therefore, because these tests have been shown to be either invalid when only sampling variance occurs in the population abundances (Bulmer's R, Pollard et al.'s, and Dennis and Taper's tests) or lack power (Bulmer's R * test), little justification exists for use of such tests to support or refute the hypothesis of density dependence.
Some theories and experimental studies suggest that areas of low plant species richness may be invaded more easily than areas of high plant species richness. We gathered nested-scale vegetation data on plant species richness, foliar cover, and frequency from 200 1-m 2 subplots (20 1000-m 2 modified-Whittaker plots) in the Colorado Rockies (USA), and 160 1-m 2 subplots (16 1000-m 2 plots) in the Central Grasslands in Colorado, Wyoming, South Dakota, and Minnesota (USA) to test the generality of this paradigm. At the 1-m 2 scale, the paradigm was supported in four prairie types in the Central Grasslands, where exotic species richness declined with increasing plant species richness and cover. At the 1-m 2 scale, five forest and meadow vegetation types in the Colorado Rockies contradicted the paradigm; exotic species richness increased with native-plant species richness and foliar cover. At the 1000-m 2 plot scale (among vegetation types), 83% of the variance in exotic species richness in the Central Grasslands was explained by the total percentage of nitrogen in the soil and the cover of native plant species. In the Colorado Rockies, 69% of the variance in exotic species richness in 1000-m 2 plots was explained by the number of native plant species and the total percentage of soil carbon. At landscape and biome scales, exotic species primarily invaded areas of high species richness in the four Central Grasslands sites and in the five Colorado Rockies vegetation types. For the nine vegetation types in both biomes, exotic species cover was positively correlated with mean foliar cover, mean soil percentage N, and the total number of exotic species. These patterns of invasibility depend on spatial scale, biome and vegetation type, spatial autocorrelation effects, availability of resources, and species-specific responses to grazing and other disturbances. We conclude that: (1) sites high in herbaceous foliar cover and soil fertility, and hot spots of plant diversity (and biodiversity), are invasible in many landscapes; and (2) this pattern may be more closely related to the degree resources are available in native plant communities, independent of species richness. Exotic plant invasions in rare habitats and distinctive plant communities pose a significant challenge to land managers and conservation biologists.
Parasites and pathogens can influence the survivorship, behavior, and very structure of their host species. For example, experimental studies have shown that trematode parasites can cause high frequencies of severe limb malformations in amphibians. In a broad-scale field survey covering parts of California, Oregon, Washington, Idaho, and Montana, we examined relationships between the frequency and types of morphological abnormalities in amphibians and the abundance of trematode parasite infection, pH, concentrations of 61 pesticides, and levels of orthophosphate and total nitrate. We recorded severe malformations at frequencies ranging from 1% to 90% in nine amphibian species from 53 aquatic systems. Infection of larvae by the trematode Ribeiroia ondatrae was associated with, and functionally related to, higher frequencies of amphibian limb malformations than found in uninfected populations (≤5%). Parasites were concentrated around the basal tissue of hind limbs in infected anurans, and malformations associated with infection included skin webbings, supernumerary limbs and digits, and missing or malformed hind limbs. In the absence of Ribeiroia, amphibian populations exhibited low (0-5%) frequencies of abnormalities involving missing digits or distal portions of a hind limb. Species were affected differentially by the parasite, and Ambystoma macrodactylum, Hyla regilla, Rand aurora, R. luteiventris, and Taricha torosa typically exhibited the highest frequencies of abnormalities. None of the water-quality variables measured was associated with malformed amphibians, but aquatic snail hosts (Planorbella spp.) were significant predictors of the presence and abundance of Ribeiroia infection. Morphological comparisons of adult specimens of Ribeiroia collected from different sites and raised in experimental definitive hosts suggested that all samples represented the same species - R. ondatrae. These field results, coupled with experimental research on the effects of Ribeiroia on amphibians, demonstrate that Ribeiroia infection is an important and widespread cause of amphibian limb malformations in the western United States. The relevance of trematode infection to declines of amphibian populations and the influence of habitat modification on the pathology and life cycle of Ribeiroia are emphasized as areas requiring further research.
The geomorphic effectiveness of extreme floods increases with aridity and decreasing watershed size. Therefore, in small dry watersheds extreme floods should control the age structure and spatial distribution of populations of disturbance-dependent riparian trees. We examined the influence of extreme floods on the bottomland morphology and forest of ephemeral streams in a semiarid region. Along six stream reaches on the Colorado Piedmont we examined channel changes by analyzing a rectified sequence of aerial photographs spanning 56 yr, and we investigated the spatial distribution of different-aged patches of forest by aging 189 randomly sampled cottonwood trees. Channel change in these ephemeral sand-bed streams is dominated by widening, which occurs over a span of hours during infrequent floods, and postflood narrowing, which occurs over decades between floods. Narrowing is accelerated where reliable moisture increases the density and growth rate of vegetation on the former bed. Reproduction of cottonwood trees has occurred mostly in former channel bed during periods of channel narrowing beginning after floods in 1935 and 1965 and continuing for as long as two decades. Thus cottonwood establishment is related to low flows at the time scale of a year, but to high flows at the time scale of decades. At sites that have not experienced major floods in the last 80 yr, little channel change has occurred, cottonwood reproduction has been limited, tree density has declined, and succession to grassland is occurring. Because channel change and tree reproduction in this region are driven by infrequent local events, channel width and tree age distributions vary greatly over time and among sites. For the same reason, riparian forests along these ephemeral streams can be as wide as forests along perennial rivers with much higher mean discharge.