Showing papers in "Ecological Monographs in 2008"

Measuring bee diversity in different european habitats and biogeographical regions

Abstract: Bee pollinators are currently recorded with many different sampling methods. However, the relative performances of these methods have not been systematically evaluated and compared. In response to the strong need to record ongoing shifts in pollinator diversity and abundance, global and regional pollinator initiatives must adopt standardized sampling protocols when developing large-scale and long-term monitoring schemes. We systematically evaluated the performance of six sampling methods (observation plots, pan traps, standardized and variable transect walks, trap nests with reed internodes or paper tubes) that are commonly used across a wide range of geographical regions in Europe and in two habitat types (agricultural and seminatural). We focused on bees since they represent the most important pollinator group worldwide. Several characteristics of the methods were considered in order to evaluate their performance in assessing bee diversity: sample coverage, observed species richness, species richness estimators, collector biases (identified by subunit-based rarefaction curves), species composition of the samples, and the indication of overall bee species richness (estimated from combined total samples). The most efficient method in all geographical regions, in both the agricultural and seminatural habitats, was the pan trap method. It had the highest sample coverage, collected the highest number of species, showed negligible collector bias, detected similar species as the transect methods, and was the best indicator of overall bee species richness. The transect methods were also relatively efficient, but they had a significant collector bias. The observation plots showed poor performance. As trap nests are restricted to cavity-nesting bee species, they had a naturally low sample coverage. However, both trap nest types detected additional species that were not recorded by any of the other methods. For large-scale and long-term monitoring schemes with surveyors with different experience levels, we recommend pan traps as the most efficient, unbiased, and cost-effective method for sampling bee diversity. Trap nests with reed internodes could be used as a complementary sampling method to maximize the numbers of collected species. Transect walks are the principal method for detailed studies focusing on plant-pollinator associations. Moreover, they can be used in monitoring schemes after training the surveyors to standardize their collection skills.

A multi-scale test of the forage maturation hypothesis in a partially migratory ungulate population

Abstract: The forage maturation hypothesis (FMH) proposes that ungulate migration is driven by selection for high forage quality. Because quality declines with plant maturation, but intake declines at low biomass, ungulates are predicted to select for intermediate forage biomass to maximize energy intake by following phenological gradients during the growing season. We tested the FMH in the Canadian Rocky Mountains by comparing forage availability and selection by both migrant and nonmigratory resident elk (Cervus elaphus) during three growing seasons from 2002-2004. First, we confirmed that the expected trade-off between forage quality and quantity occurred across vegetation communities. Next, we modeled forage biomass and phenology during the growing season by combining ground and remote-sensing approaches. The growing season started 2.2 days earlier every 1 km east of the continental divide, was delayed by 50 days for every 1000-m increase in elevation, and occurred 8 days earlier on south aspects. Migrant and resident selection for forage biomass was then compared across three spatial scales (across the study area, within summer home ranges, and along movement paths) using VHF and GPS telemetry locations from 119 female elk. Migrant home ranges occurred closer to the continental divide in areas of higher topographical diversity, resulting in migrants consistently selecting for intermediate biomass at the two largest scales, but not at the finest scale along movement paths. In contrast, residents selected maximum forage biomass across all spatial scales. To evaluate the consequences of selection, we compared exposure at telemetry locations of migrant and resident elk to expected forage biomass and digestibility. The expected digestibility for migrant elk in summer was 6.5% higher than for residents, which was corroborated with higher fecal nitrogen levels for migrants. The observed differences in digestibility should increase migrant elk body mass, pregnancy rates, and adult and calf survival rates. Whether bottom-up effects of improved forage quality are realized will ultimately depend on trade-offs between forage and predation. Nevertheless, this study provides comprehensive evidence that montane ungulate migration leads to greater access to higher-quality forage relative to nonmigratory congeners, as predicted by the forage maturation hypothesis, resulting primarily from large-scale selection patterns.

Polar bear diets and arctic marine food webs: insights from fatty acid analysis

Abstract: We used quantitative fatty acid signature analysis (QFASA) to examine the diets of 1738 individual polar bears (Ursus maritimus) sampled across the Canadian Arctic over a 30-year span. Polar bear foraging varied over large and small spatial and temporal scales, and between demographic groups. Diets in every subpopulation were dominated by ringed seals (Phoca hispida) and, in the eastern Arctic, secondarily by harp seals (Pagophilus groenlandica). Beluga whales (Delphinapterus leucas) were an important food source for bears in the High Arctic, which is consistent with previous anecdotal reports. Foraging patterns were most similar among neighboring subpopulations with similar prey assemblages, but also differed geographically within Western Hudson Bay. The sexual size dimorphism of polar bears had an important effect on foraging, as large bearded seals (Erignathus barbatus) and walruses (Odobenus rosmarus) were consumed most often by older, male bears, whereas ringed seals and, where available, harbor seals (P. vitulina) were most important to younger age classes. Larger, older bears also had the greatest dietary diversity, apparently because of their ability to include larger-bodied prey. During spring and summer, polar bears in some areas increased predation on migratory harp seals and beluga whales. In Western Hudson Bay, bearded seal consumption declined between 1995 and 2001 for both male and female bears and continued to decline among females up to the most recent sampling (2004). Ringed seal consumption in Western Hudson Bay increased between 1998 and 2001, perhaps in response to increased ringed seal productivity, but was not significantly affected by date of sea-ice breakup. Overall, our data indicate that polar bears are capable of opportunistically altering their foraging to take advantage of locally abundant prey, or to some degree compensating for a decline in a dominant prey species. However, in other areas polar bears are dependent on the availability of ringed and bearded seals. Recent population data suggest that polar bears with the most specialized diets may be most vulnerable to climate-related changes in ice conditions. The results of this large-scale, ecosystem-based study indicate a complex relationship between sea-ice conditions, prey population dynamics, and polar bear foraging.

Seasonal reassembly of a river food web: floods, droughts, and impacts of fish

Abstract: Eighteen years of field observations and five summer field experiments in a coastal California river suggest that hydrologic regimes influence algal blooms and the impacts of fish on algae, cyanobacteria, invertebrates, and small vertebrates. In this Mediterranean climate, rainy winters precede the biologically active summer low-flow season. Cladophora glomerata, the filamentous green alga that dominates primary producer biomass during summer, reaches peak biomass during late spring or early summer. Cladophora blooms are larger if floods during the preceding winter attained or exceeded “bankfull discharge” (sufficient to mobilize much of the river bed, estimated at 120 m3/s). In 9 out of 12 summers preceded by large bed-scouring floods, the average peak height of attached Cladophora turfs equaled or exceeded 50 cm. In five out of six years when flows remained below bankfull, Cladophora biomass peaked at lower levels. Flood effects on algae were partially mediated through impacts on consumers in food webs....

Fire, logging, and overstory affect understory abundance, diversity, and composition in boreal forest

Abstract: Understory vegetation plays a critical role in boreal ecosystems. Despite this, quantitative evaluation of the factors controlling understory vegetation abundance, diversity, and composition in the most diverse boreal forest region in North America is lacking. This study examined the dynamics of understory vegetation of stands of fire origin and tested effects of overstory composition and logging vs. fire on the understory vegetation dynamics in Ontario, Canada. Understory vegetation communities were sampled in 68 stands of conifer, mixed-wood, and deciduous overstory type ranging from 7 to 201 years postfire for stands of fire origin, and from 7 to 31 years for stands of logging origin. For stands of fire origin, total cover and species richness followed similar trends for the three overstory types and were highest in the intermediate-aged stands (72-90 years). Trends in cover and richness, however, differed significantly for vascular and nonvascular plant groups. Vascular cover and species richness were generally higher under deciduous stands, and lower on older stands, while nonvascular species richness was highest under conifer stands and increased with time since fire. Neither species richness nor compositional turnover was higher under mixed-wood stands; mixed-wood stands were compositionally intermediate to conifer and deciduous stands. Multivariate analysis using multiple-response permutation procedures indicated that understory communities were compositionally distinct for all overstory types and showed no convergence with increasing time since fire. Compared with postfire stands of similar ages, post-logged stands had similar total understory cover and richness. Vascular cover and richness, however, were higher on post-logged stands, and nonvascular cover and richness were lower. Stands of logging and fire origin were compositionally distinct for all overstory types and ages. Compositional differences appeared to be driven by higher preestablished rhizomatous species and few pyrophilic species on post-logged sites. Understory vegetation communities in the central boreal shield appear to support the intermediate disturbance hypothesis. Understory richness, however, was not negatively associated with high cover values as predicted by the intermediate disturbance hypothesis. Moreover, richness appears to be highest on sites with high light availability, suggesting that boreal understory communities are influenced more by plant tolerances for low resources, rather than by competition.

Douglas-fir growth in mountain ecosystems. water limits tree growth from stand to region

Abstract: The purpose of this work is to understand the nature of growth-climate relationships for Douglas-fir (Pseudotsuga menziesii) across the climatic dimensions of its niche. We used a combination of biophysically informed sampling (to identify sample sites) and dendroclimatology (to identify growth-climate relationships) along a climate gradient in northwestern United States mountain ecosystems from the western Olympic Peninsula, Washington to the eastern Rocky Mountain Front, Montana. We used a multi-scale sampling strategy that accounted for continentality, physiography, and topography as non-climatic factors that could influence climate and alter tree growth. We developed a network of 124 Douglas-fir tree-ring chronologies and explored growth-climate correlations across the sampled gradients. We considered two different spatial scales of monthly and seasonal climate variables as potential controlling factors on tree growth. Annual radial growth in 60-65% of the plots across the region is significantly correlated with precipitation, drought, or water balance during the late summer prior to growth and the early summer the year of growth. In a few plots, growth is positively correlated with cool-season temperature or negatively correlated with snowpack. Water availability is therefore more commonly limiting to Douglas-fir growth than energy limitations on growing season length. The first principal component derived from the chronologies is significantly correlated with independent drought reconstructions. The sensitivity of Douglas-fir to summer water balance deficit (potential evapotranspiration minus actual evapotranspiration) indicates that increases in April to September temperature without increases in summer precipitation or soil moisture reserves are likely to cause decreases in growth over much of the sampled area, especially east of the Cascade crest. In contrast, Douglas-fir may exhibit growth increases at some higher elevation sites where seasonal photosynthesis is currently limited by growing-season length or low growing-season temperature. Life-history processes such as establishment, growth, and mortality are precursors to changes in biogeography, and measurements of climate effects on those processes can provide early indications of climate-change effects on ecosystems.

Scaling from trees to forests: tractable macroscopic equations for forest dynamics

Abstract: Individual-based forest simulators, such as TASS and SORTIE, are spatial stochastic processes that predict properties of populations and communities by simulating the fate of every plant throughout its life cycle. Although they are used for forest management and are able to predict dynamics of real forests, they are also analytically intractable, which limits their usefulness to basic scientists. We have developed a new spatial individual-based forest model that includes a perfect plasticity formulation for crown shape. Its structure allows us to derive an accurate approximation for the individual-based model that predicts mean densities and size structures using the same parameter values and functional forms, and also it is analytically tractable. The approximation is represented by a system of von Foerster partial differential equations coupled with an integral equation that we call the perfect plasticity approximation (PPA). We have derived a series of analytical results including equilibrium abundances for trees of different crown shapes, stability conditions, transient behaviors, such as the constant yield law and self-thinning exponents, and two species coexistence conditions.

Spatial and temporal patterns of invertebrate recruitment along the west coast of the united states

Abstract: Patterns of recruitment in marine ecosystems can reflect the distribution of adults, dispersal by ocean currents, or patterns of mortality after settlement. In turn, patterns of recruitment can play an important role in determining patterns of adult abundance and community dynamics. Here we examine the biogeographic structure of recruitment variability along the U.S. West Coast and examine its association with temperature variability. From 1997 to 2004 we monitored monthly recruitment rates of dominant intertidal invertebrates, mussels and barnacles, at 26 rocky shore sites on the West Coast of the United States, from northern Oregon to southern California, a span of 1750 km of coastline. We examined spatial variation in the dynamics of recruitment rates and their relationship to coastal oceanography using satellite-derived time series of monthly sea surface temperature (SST). Recruitment rates showed a biogeographic structure with large regions under similar dynamics delimited by abrupt transitions. The seasonal peak in recruitment rates for both mussels and barnacles changed from a late summer-early fall peak in Oregon to winter or early spring in northern California, and then back toward summer in southern California. Recruitment rates varied greatly in magnitude across the latitudinal range. The barnacle Balanus glandula and mussels (Mytilus spp.) showed a decline of two orders of magnitude south of Oregon. In contrast, recruitment rates of barnacles of the genus Chthamalus showed a variable pattern across the region examined. The spatial distribution of associations between raw SST and recruitment rates for all species showed positive associations, indicating recruitment during warm months, for all species in Oregon, northern California, and several sites in south-central California. By considerably extending the spatial and temporal scales beyond that of previous studies on larval recruitment rates in this system, our study has identified major biogeographic breaks around Cape Blanco and Point Conception despite considerable spatial and temporal variation within each region and among species. These large differences in recruitment rates across biogeographic scales highlight the need for a better understanding of larval responses to ocean circulation patterns in the conservation and management of coastal ecosystems.

Changing the gap dynamics paradigm: vegetative regeneration control on forest response to disturbance

Abstract: Understanding the manner in which changes in disturbance regimes will affect forest biodiversity is an important goal of global change research. Prevailing theories of recruitment after disturbance center on the role of pioneer species; predictions of forest biodiversity focus almost exclusively on dispersal and shade tolerance while vegetative reproduction is virtually omitted from models and serious discussions of the topic. However, the persistence of live damaged trees increases understory shade, generates fine-scale environmental heterogeneity, and moderates ecosystem responses to damage, while the sprouting of damaged trees offers a shortcut to reestablishment of the canopy. While a number of studies document snapshots of post-disturbance vegetative reproduction, we lack an understanding of the underlying demographic processes needed in order to both comprehend and predict observed patterns. In this study we quantify the abundance, competitive ability, and interspecific variability of vegetative reproduction in 18 replicated experimental gaps in the southern Appalachians and Carolina Piedmont, USA, in order to assess the potential role of sprouting in driving gap dynamics. Annual rates of damaged adult survival, sprout initiation, growth, and mortality were monitored over four years and compared to the performance of gap-regenerating saplings. Recruitment from sprouts was found to constitute 26-87% of early gap regeneration and forms the dominant pathway of regeneration for some species. Sprouts from recently damaged trees also grow significantly faster than the saplings with which they compete. For all measured demographic rates (damaged tree survival, sprout initiation, number, growth, and survival) differences among species are large and consistent across sites, suggesting that vegetative reproduction is an important and non-neutral process in shaping community composition. Sprouting ability does not correlate strongly with other life-history trade-offs, thus sprouting potentially provides an alternate trait axis in promoting diversity.

No evidence for trade‐offs: centaurea plants from america are better competitors and defenders

Abstract: The natural enemies hypothesis has led to a number of ideas by which invaders might evolve superior competitive ability. In this context, we compared growth, reproduction, competitive effect, competitive response, and defense capabilities between invasive North American populations of Centaurea maculosa and populations in Europe, where the species is native. We found that Centaurea from North America were larger than plants from European populations. North American Centaurea also demonstrated stronger competitive effects and responses than European Centaurea. However, competitive superiority did not come at a cost to herbivore defense. North American plants were much better defended against generalist insect herbivores and slightly better defended against specialists. North Americans showed a stronger inhibitory effect on the consumers (resistance) and a better ability to regrow after attack by herbivores (tolerance). Better defense by North Americans corresponded with higher constitutive levels of a biochemical defense compound precursor, tougher leaves, and more leaf trichomes than Europeans. North American F1 progeny of field collected lines retained the traits of larger size and greater leaf toughness suggesting that genetic differences, rather than maternal effects, may be the cause of intercontinental differences, but these sample sizes were small. Our results suggest that the evolution of increased competitive ability may not always be driven by physiological trade-offs between the allocation of energy or resources to growth or to defense. Instead, we hypothesize that Centaurea maculosa experiences strong directional selection on novel competitive and defense traits in its new range.

Intertidal mussels exhibit energetic trade-offs between reproduction and stress resistance

Abstract: Life-history theory suggests that trade-offs exist between fitness components, with organisms balancing investment in reproduction against survival and future reproduction. This study examined the influence of stress on physiological trade-offs in the dominant rocky intertidal mussel Mytilus californianus on the central Oregon coast, USA. The intertidal zone is a highly heterogeneous thermal environment that could lead to intrapopulation variation in stress responses. Stress increases along a vertical gradient, with higher physical stress occurring in the higher intertidal zone, both due to reduced feeding time and longer exposure to aerial conditions. Reproduction and carotenoid content were compared in mussels from the low and high vertical edges of the mussel bed. High-edge mussels invested less relative energy in reproduction and also spawned all of their gametes in the early summer, whereas low-edge mussels continuously spawned small batches of gametes throughout the year. High-edge mussels accumulated high concentrations of carotenoid pigments into their gonadal tissues, potentially to protect gametes from damaging oxidative stress experienced during aerial exposure. A reciprocal transplant experiment revealed plastic responses in growth and reproduction to increased stress. In contrast, carotenoid content did not increase in response to stress, suggesting that carotenoids may not change rapidly or may not be easily lost or gained. Our results indicate that mussels exhibit physiological trade-offs and, under increased stress predicted from climate change scenarios, may allocate energy away from reproduction toward costly physiological defenses.

Quantifying periodic, stochastic, and catastrophic environmental variation

Abstract: Environmental variation plays a central role in regulating processes at all levels of ecological organization. Environmental data (e.g., temperature, rainfall, stream discharge, water chemistry) are typically easy to collect in large quantity, a requirement for many data-hungry time series tools. Unfortunately, these data are very rarely used effectively in ecology. Here we address this problem by outlining a suite of tools that can be used to quantify periodic, stochastic, and catastrophic variation in environmental conditions. We illustrate the application of these tools using long-term records of average daily discharge in 105 streams and rivers maintained by the U.S. Geological Survey on the NWIS (National Water Information System) web site. Specifically, we apply Fourier analysis to estimate the periodic (seasonal) and stochastic (interannual) components of variation in discharge. We then estimate the temporal autocorrelation structure of stochastic variation (i.e., noise color) in daily flows for ea...

Neighborhood models of the effects of invasive tree species on ecosystem processes

Abstract: Changes in the composition of a community due to the invasion by exotic plant species can lead to modification of ecosystem function that, in turn, produces feedbacks that drive further changes in community composition. The development of predictive models of this process requires an understanding of the spatial extent of the impacts of the exotic species, particularly during early stages of invasion. The main objective of this study was to characterize the neighborhood dynamics of ecosystem transformations by the invasive tree species Norway maple (Acer platanoides L.) and tree of heaven (Ailanthus altissima Mill.) in temperate forests of the northeastern United States. By using a neighborhood approach in multiple sites, we sought to determine (1) the magnitude and spatial extent of the effects of the two invasive species on soil properties and processes; (2) whether the effects of the invasive species were site dependent; and (3) the differences in the effects of invasive versus native tree species on ecosystem processes. Our results showed that Norway maple and tree of heaven alter the functioning of temperate forest ecosystems even at relatively low densities by increasing cycling rates (i.e., net N mineralization, net nitrification, Ca mineralization) and nutrient availability (i.e., pH, Ca, Mg, K, N). At the neighborhood scale, the spatial extent of the impact of the two species varied strikingly among soil properties. Moreover, the neighborhood effects of the two invasive species were site dependent, with the magnitude of the impact increasing with soil fertility. At the community level, Norway maple and to a lesser extent tree of heaven had stronger effect on soils than any of the dominant native tree species considered. We conclude that the invasion of northeastern forests by Norway maple and tree of heaven is characterized by predictable, neighborhood-specific acceleration of nutrient cycling rates and localized increases in nutrient pools. These ecosystem alterations have enormous potential for the modification of competitive hierarchies in forest communities. In particular, Norway maple and tree of heaven may change relative abundances within the native community to the benefit of native species that are more competitive on nutrient rich sites.

Animal habitat quality and ecosystem functioning: exploring seasonal patterns using ndvi

Abstract: Many animal species have developed specific evolutionary adaptations to survive prolonged periods of low energy availability that characterize seasonal environments. The seasonal course of primary production, a major aspect of ecosystem functioning, should therefore be an important factor determining the habitat quality of such species. We tested this hypothesis by analyzing the relationship between habitat quality and ecosystem functioning for brown bears (Ursus arctos), a species showing hyperphagia and hibernation as evolutionary adaptation to seasonal peaks and bottlenecks in ecosystem productivity, respectively. Our unique long-term data set comprised data from two brown bear populations in northern Spain on historical presence, current presence, and reproduction. The data were classified on a grid of 5 3 5 km pixels into five classes: frequent reproduction, sporadic reproduction, frequent presence, sporadic presence, and recent extinction. We used the long- term average of the seasonal course of NDVI (normalized difference vegetation index) as a proxy for ecosystem functioning and investigated the relationship between habitat quality and ecosystem functioning with methods borrowed from statistical point-pattern analysis. We found that brown bears indeed selected habitat with specific ecosystem functioning (i.e., the variance in all habitat classes was smaller than in the landscape overall) and the relationship between habitat quality and ecosystem functioning was ordered. First, the average distance in ecosystem functioning between two habitat classes was larger if the difference in habitat quality was larger. Second, habitat for which there was the greatest need (i.e., breeding habitat) occupied the narrowest niche regarding ecosystem functioning and showed the most pronounced seasonality. Progressively poorer classes occupied wider niches that partly overlapped those of better classes. This indicated that nonbreeding animals are less selective. Our methodology provided new insight into the relationship between ecosystem functioning and habitat quality and could be widely applied to animal species living in seasonal environments. Because NDVI data are continuously collected, our methodology allows for continuous monitoring of changes in habitat quality due to global change.

Multiple regime shifts in a subtropical peatland: community‐specific thresholds to eutrophication

Abstract: Ecosystems have a natural resilience to perturbations, where resilience is the magnitude of a disturbance that an ecosystem can resist before changes in structure, function, and services result in a regime shift. The Everglades region of Florida, USA, has been detrimentally impacted by phosphorus (P) enrichment and a regime shift from Cladium (sawgrass) to Typha (cattail) marsh has been described. We examine another facet of the low nutrient Everglades stability regime, open-water sloughs, to determine if eutrophication leads to similar regime shifts. We analyzed surface water P and soil P as controlling variables that, once a critical threshold is surpassed, alter ecosystem state variables. Nonlinear relationships between P and vegetation were observed along a northern Everglades eutrophication gradient. In addition to the Cladium-Typha regime shift, a second independent regime shift, slough-Typha, was identified. Synoptic surveys of 49 sloughs within the boundary between the slough and Typha regime revealed that surface water total phosphorus (TP) and the benthic algal floe layer (BAFL) were the controlling variables, with critical thresholds of 11 ug/L and 412 mg/kg, respectively. The slough regime below these thresholds was characterized by calcareous periphyton (BAFL TP = 298 mg P/kg; BAFL calcium = 149 g Ca/kg). Above the TP thresholds, vegetation composition shifted to open-marsh species with significantly higher BAFL TP (700 mg P/kg) and total organic carbon (TOC) (350 g C/kg). A second BAFL TP threshold occurred at 712 mg P/kg, above which Nymphaea dominated and BAFL TP (1034 mg P/kg) and TOC (417 g C/kg) significantly increased. Nymphaea sloughs transitioned to the Typha regime. The boundary reflects the loss of ecosystem resilience due to eutrophication. Both low-nutrient stability regimes (slough and Cladium) lie precariously close to the P critical threshold but differ in how eutrophication is absorbed and resisted. The slough regime transitions rapidly through a series of ecosystem state changes linked to positive feedback loops that affect P dynamics, whereas the Cladium regime does not. An adaptive management strategy has been implemented to address the surface water TP threshold; however, to ensure successful restoration of the Everglades, the BAFL and soil TP thresholds also need to be considered.

Tests of the mid-domain hypothesis: a review of the evidence

Abstract: Geographic variation of species richness is strongly correlated with environmental gradients. However, random arrangement of species distributions within a bounded domain can also theoretically produce richness gradients without underlying environmental gradients. This mid-domain effect (MDE) could serve as the null hypothesis against which to test effects of environmental variables, or as a component of a multivariate explanation of species-richness patterns. Recent reviews have concluded that there is a substantial MDE signature in observed geographical patterns of richness, based on correlations between observed patterns of richness and the predictions of mid-domain models. However, the mid-domain hypothesis makes additional powerful predictions about how richness should vary through space, and about the slope of the relationship between predicted and observed richness. Very few studies have tested these more powerful MDE predictions. Here, we reexamine the published mid-domain literature for agreement between observed patterns of richness and MDE predictions. We find that 50 of 53 published studies of MDEs showed significant deviations from the predictions of mid-domain models. When observed richness is correlated with MDE predictions, there are nearly always strongly collinear environmental gradients (e.g., in the Americas, climatic favorability and MDE-predicted richness are both maximal in the middle). Interpolation in sparsely sampled data can also give rise to spurious, apparently strong, mid-domain effects (e.g., the classic study of the Madagascan rain forest). We conclude that observed broad-scale patterns of species richness are not consistent with the mid-domain hypothesis.

Evolution of adaptive plasticity: risk‐sensitive hatching in neotropical leaf‐breeding treefrogs

Abstract: Adaptive plasticity at switch points in complex life cycles (e.g., hatching, metamorphosis) is well known, but the evolutionary history of such plasticity is not. Particularly unclear is how a single plastic response (e.g., shifts in hatching timing) evolves to respond to different threats and cues (e.g., abiotic and biotic). We conducted a comparative phylogenetic study of hatching plasticity in a group of frogs with arboreal embryos to determine when risk-accelerated hatching evolved in the clade, whether responses to two common egg-stage risks (snake predation and flooding) evolved independently, and whether the overall capacity for hatching plasticity was evolutionarily more conservative than responses to specific cues. Red-eyed treefrogs (Agalychnis callidryas) hatch early to escape from several egg-stage risks but otherwise hatch later, improving larval survival with predators. We reconstructed a phylogeny for Agalychnis and related genera based on three mitochondrial and four nuclear genes. We quantified onset of hatching competence, spontaneous hatching timing, responses to egg-stage risks, and costs of premature hatching in Agalychnis and Pachymedusa. We also assessed hatching plasticity in a basal phyllomedusine, Cruziohyla calcarifer. The capacity to hatch ;30% before the spontaneous hatching age appears ancestral for phyllomedusines, with little change over ;34-50 million years among the species examined. A strong hatching response to flooding, with no mortality of hatching-competent eggs, is similarly ancient and conserved. Premature hatchlings of Agalychnis and Pachymedusa are more vulnerable to fish predation than are full-term hatchlings, indicating a conserved risk trade-off across hatching that would make plasticity advantageous. In contrast, the hatching response to snake attack has undergone major changes at least twice in the Agalychnis-Pachymedusa clade, with two species showing substantially lower escape success than the others. Responses to different threats have thus evolved independently. The capacity for switch point plasticity may be evolutionarily more stable than the response to individual stage-specific threats.

Crown development of coastal pseudotsuga menziesii, including a conceptual model for tall conifers

Abstract: Seventy trees from seven stands 50–650 years old were selected for this investigation of crown structural development in Pseudotsuga menziesii. All branches, limbs, and trunks were nondestructively measured for size, structure, and location while climbing the trees with ropes. These data were used to generate a computer model of each tree's crown that was error-checked trigonometrically. Leaves, bark, cambium, and wood were quantified by using limited destructive sampling to develop predictive equations that were applied to the complete inventory of structures in each tree's crown. Summations of these values yielded whole-tree estimates of several structural variables. A second set of equations was then developed to predict these whole-tree parameters from simple, ground-based measurements. Principal components analysis of 24 tree-level variables revealed two orthogonal dimensions of structure that accounted for 71.3% and 12.4% of total variation in the 70 trees. The first dimension represented a gradient...

Plant and microbe contribution to community resilience in a directionally changing environment

Abstract: To understand the role biota play in resilience or vulnerability to environmental change, we investigated soil, plant, and microbial responses to a widespread environmental change, increased nitrogen (N). Our aim was to test the plant-soil threshold hypothesis: that changed biotic structure influences resilience to accumulated changes in N. For six years, we removed one of two codominant species, Geum rossii and Deschampsia caespitosa, in moist- meadow alpine tundra in Colorado, USA. We also manipulated nutrient availability by adding carbon (C) or N, separately and in combination with the species removals. Consistent with our hypothesis, Geum was associated with soil feedbacks that slowed rates of N cycling and Deschampsia with feedbacks that increased rates of N cycling. After a four- year initial resilience period, Geum dramatically declined (by almost 70%) due to increasing N availability. In contrast, Deschampsia abundance did not respond to changes in N supply; it only responded to the removal of Geum. Forbs and graminoids responded more positively to Deschampsia removal than to Geum removal, indicating stronger competitive effects by Deschampsia. The changed biotic interactions appear to have community-level consequences: after six years of Geum (but not Deschampsia) removal, evenness of the community declined by over 35%. Increased N affected the soil-microbial feedbacks, particularly in association with Geum. Microbial biomass N declined at higher N, as did the activities of two C-acquiring and one N- acquiring extracellular microbial enzymes. In the presence of Geum, N fertilization slowed the activity of phenol oxidase, a tannin-degrading enzyme, suggesting that microbes shift from degrading Geum-derived compounds. In the absence of Geum, acid phosphatase activity increased, suggesting increased phosphorus limitation in association with Deschampsia. With continued N deposition forecast for this system, these results suggest that initial resilience of Geum to increased N will be overwhelmed through elimination of microbial feedbacks. Once Geum declines, the loss will indirectly facilitate Deschampsia via competitive release. Because Deschampsia exerts strong competitive effects on subordinate species, increased Deschampsia abundance may be accompanied by a community-wide drop in diversity. We conclude that plant-soil feedbacks through the microbial community can influence vulnerability to exogenous changes in N and contribute to threshold dynamics.

Scale‐dependent habitat use by fall migratory birds: vegetation structure, floristics, and geography

Abstract: Animal habitat selection is a central focus of ecology and conservation biology. Understanding habitat associations in migratory animals is particularly complicated because individuals have variable habitat requirements during the annual cycle, across their geographic range, along migratory routes, and at multiple spatial scales. We studied habitat associations of 16 fall Nearctic-Neotropical migratory land birds at two spatial scales at a stopover site along the northern Yucatan coast to examine scale-dependent habitat use, identify proximate cues shaping birds' distributions, and evaluate similarities in habitat use between our tropical stopover site and temperate sites. We addressed scale-dependent habitat associations in two ways, by (1) quantifying species' distributions among and within broad vegetation types and (2) comparing migrants' associations with architectural gradients between the two spatial scales. We also evaluated the relative importance of vegetation architecture and floristics in explaining migrants' distributions within broad coastal vegetation types. Bird species were nonrandomly distributed among broad- and fine-scale vegetation types, and patterns of habitat use varied between the two scales. Moreover, birds had different preferences for vegetation architecture at the two scales, which may reflect trade-offs between competing demands and/or reduced variation in vegetation at the small scale in our study. These findings illustrate the manner in which spatial scale and range of vegetation variation influence perceptions of animal-habitat associations. Within broad coastal vegetation types birds refined their distributions in relation to architectural and floristic attributes, which provided them with redundant and/or complementary information regarding the distribution of suitable habitat. We suggest that the relative importance of architecture and floristics is likely scale-dependent. Habitat use at our site was similar to that observed at temperate stopover sites for almost all species, indicating that habitat affinities are maintained along the migratory route for these eastern populations despite latitudinal changes in environmental factors. We highlight examples of similar patterns observed in other migratory and nonmigratory animals to illustrate the generality of these patterns beyond en route land birds and our tropical site. We also indicate where we expect to see differences and outline areas of research that merit greater attention in order to advance our understanding of animal habitat selection.

Temporal changes in vertebrates during landscape transformation: a large-scale ''natural experiment''

Abstract: Plantation development is a significant form of landscape change worldwide. We report findings from a large-scale longitudinal natural experiment that quantified changes in Australian vertebrates as a former grazing landscape was transformed to one dominated by a radiata pine (Pinus radiata) plantation. The study included four main “treatments”: woodland remnants surrounded by emerging radiata pine (52 sites, termed “woodland treatments”), stands of radiata pine (10 sites, “pine controls”), woodland remnants where the surrounding landscape remained unchanged (56 sites, “woodland controls”), and paddocks with scattered woodland trees that surrounded the 56 woodland remnants (10 sites, “paddock controls”). In our study region, woodland is distinguished from forest by differences in tree height, tree spacing, bole length, and canopy development. Between 1998 and 2006, occupancy rates of “woodland treatments” by most mammals and reptiles increased linearly. Similar trends occurred in the “woodland controls,” suggesting that species had increased landscape-wide, rather than displaying year × treatment interaction effects. We cross-classified birds according to the statistical significance and nature of time trajectories. Groups included those that: (1) declined in woodland treatments in comparison with woodland controls, (2) decreased within woodland treatments but increased in woodland controls, (3) declined across the entire study area, (4) increased within woodland treatments in comparison with woodland controls, (5) increased within woodland treatments but declined in woodland controls, and (6) increased across the entire study area. Attributes of woodland treatments significantly associated with temporal changes in bird occupancy included: (1) age of surrounding pine stands; (2) number of boundaries with surrounding pines; (3) size of the woodland patches; (4) dominant vegetation type of woodland patches; and (5) temporal changes in vegetation structure in the woodland treatments. Bird species associated with open country and woodland environments were disadvantaged by landscape transformation, whereas those that benefited were forest taxa and/or habitat generalists capable of inhabiting pine stands and adjacent woodland patches. Beyond this generalization, an unanticipated finding was a lack of association between life history attributes and landscape transformation. We suggest that several key processes are likely drivers of change at multiple spatial scales. Recognition of such processes is important for conservation in landscapes transformed by plantation expansion.

Stand and landscape-level disturbance dynamics in old-growth forests in western massachusetts

Abstract: Natural disturbances strongly influence the dynamics and developmental patterns of forest ecosystems; however, relatively little is known about the historic patterns of natural disturbance for many portions of eastern North America, such as southern New England, where human disturbance has predominated for centuries For these regions, much of our understanding of natural disturbance dynamics comes from studies of younger second- growth forests or isolated old-growth stands, thus limiting the temporal and spatial resolution of our knowledge of dynamics in these regions To address these limitations, we analyzed dendroecological data from the 18 largest remaining old-growth stands in western Massachusetts, ranging in proximity from 1-60 km apart, in an effort to characterize the historic stand and landscape-level patterns of natural disturbance Our results indicate that disturbance regimes for these systems were dominated by relatively frequent, low-intensity disturbances (average 50% 6 02% canopy area disturbed per decade) operating somewhat randomly on the landscape Across the study areas, most decadal disturbances (862%) involved ,10% canopy loss There was no evidence of stand-replacing disturbances during the period examined (1700-1989), and the maximum canopy area disturbed in any given decade was 263% Nonmetric multidimensional scaling demonstrated that several forests shared similar disturbance histories despite being separated by 50 km Comparisons of these patterns with model simulations of past hurricane events and historical documents suggest that broadscale disturbances, such as hurricanes and ice storms, resulted in common disturbance peaks and subsequent recruitment peaks at spatially disparate areas in the 1790s, 1870s, 1900s, and 1920s Conversely, the lack of synchrony in proximate areas during these events highlights the patchy nature of these disturbances on the landscape Compositional and physiographic factors influenced disturbance patterns, as stands located on northwest-facing slopes or containing significant Picea rubens components in the forest overstory experienced the highest levels of disturbance Our results highlight the utility of incorporating dendroecological reconstructions across numerous old-growth stands to interpret the historic stand and landscape-level disturbance dynamics in areas devoid of large, contiguous old- growth landscapes

Alteration of seagrass species composition and function over two decades

Abstract: Changes in the species composition and structural characteristics of marine vegetated habitats in response to climate change or local anthropogenic impacts may alter their quality as habitat for associated fish and invertebrates. Summer densities and biomass of the eelgrass, Zostera marina, declined significantly between 1985 and 2004 in Bogue Sound, North Carolina, USA, within the present-day zone of biogeographic overlap in the distribution of this subtidal temperate species and the intertidal subtropical seagrass, Halodule wrightii. Zostera decline was associated with increased spring water temperatures and water nutrient concentrations. In contrast, Halodule did not exhibit a consistent trend of temporal change. Experimental seagrass transplants indicated that Halodule has the capacity to grow at depths greater than it currently occupies, suggesting that Halodule might, over time, replace Zostera. The abundance and diversity of infaunal invertebrates were lower in seagrass beds dominated by Halodule than in those dominated by Zostera or mixtures of the two species, suggesting that changing seagrass species composition affects associated faunal assemblages. Experimental deployment of artificial seagrass patches mimicking the structure of the two species showed that both depth and structural characteristics of seagrass explain differences in faunal assemblages. Epifaunal community structure differed significantly between structurally identical seagrass mimics deployed in intertidal and subtidal habitat, and invertebrates and fishes were significantly more abundant in artificial Zostera than Halodule patches at one of the two study sites. Synthesis of these results suggests that, in the event of continued Zostera loss, with or without replacement by Halodule, important habitat functions will be lost, and secondary productivity of these lagoonal ecosystems may significantly decrease.

Spatial environmental correlates of intertidal recruitment: a test using barnacles in northern chile

Abstract: Nearshore circulation processes and local geomorphological patterns are thought to be important correlates of spatial variation in larval recruitment of benthic organisms. However, few studies have attempted to quantitatively separate their relative influences upon recruitment dynamics. Here we use 12 study sites spanning 250 km of the northern Chile upwelling ecosystem and the intertidal barnacles Jehlius cirratus and Notochthamalus scabrosus to examine the extent to which spatial variation in larval recruitment is related to environmental variability and how these relationships depend on spatial autocorrelation. We find that upwelling intensity and wind velocity are negatively related to larval recruitment while surface slicks showed a positive correlation. However, the pure effect of each environmental variable was lower than its combined effects. Taking spatial autocorrelation into account, we find that the spatially structured variation of upwelling dynamics, distance to upwelling fronts, wind velocity, and slick occurrence explained most of the variation in barnacle recruitment. Spatial variation in recruitment rates of barnacles showed a characteristic length scale of 60-70 km, similar to the scale estimated for the spatially structured variables (upwelling, wind velocity, and surface slicks). We find that conditions for the occurrence of surface slicks (as surrogates of internal waves activity) seem to operate across several sites, suggesting a meso- instead local-scale influence over spatial variation in barnacle recruitment. Our results suggest that spatial variation in barnacle recruitment is modulated by the combined influence of several spatially structured nearshore processes operating at scales of 60-70 km and that spatial autocorrelation must be taken into account in the study of the recruitment-environment relationship. However, the influence of local and small-scale factors on recruitment dynamics of benthic invertebrates should be interpreted cautiously.

White spruce meets black spruce: dispersal, postfire establishment, and growth in a warming climate

Abstract: Local distributions of black spruce (Picea mariana) and white spruce (Picea glauca) are largely determined by edaphic and topographic factors in the interior of Alaska, with black spruce dominant on moist permafrost sites and white spruce dominant on drier upland sites. Given the recent evidence for climate warming and permafrost degradation, the distribution of white spruce is expected to expand, but the transition from black to white spruce may be dispersal limited: unlike the semi-serotinous black spruce, postfire regeneration of white spruce relies on seed dispersal from unburned areas. To determine the relative roles of dispersal, establishment, and growth in recruitment of white and black spruce, we studied postfire spruce regeneration in a 21-year-old burn across a white spruce-black spruce transition in the interior of Alaska. Although prefire spatial distributions of adults of the two species were well separated along the topographic sequence from upland to floodplain sites, the spatial distributions of recruits overlapped considerably. Even .700 m away from its seed source, white spruce sapling density on typical black spruce sites was high enough to form fully stocked stands. In contrast, black spruce regeneration was sparse on typical white spruce upland sites. Establishment rates of both species, estimated from a statistical model, were highest in mossy, wet depressions, which tended to have a thick residual postfire organic layer (;10 cm). On all site types, height growth rates inferred from age-height relationships were comparable for recruits of both species. On typical black spruce sites � 300 m into the burn, white spruce was younger (and, therefore, shorter) than black spruce due to the timing of masting events following the fire. There was no indication that dispersal, establishment, or edaphic constraints on juvenile growth limit white spruce's capacity to invade typical black spruce stands during the recruitment stage in our study area. It is unlikely that white spruce recruits would persist to the adult stage if the permafrost returned to the original prefire levels during future postfire succession. However, if permafrost continues to degrade under climate warming, transition to a white spruce-dominated landscape could be rapid.

Anatomy of a bottleneck: diagnosing factors limiting population growth in the puerto rican parrot

Abstract: The relative importance of genetic, demographic, environmental, and catastrophic processes that maintain population bottlenecks has received little consideration. We evaluate the role of these factors in maintaining the Puerto Rican Parrot (Amazona vittata) in a prolonged bottleneck from 1973 through 2000 despite intensive conservation efforts. We first conduct a risk analysis, then examine evidence for the importance of specific processes maintaining the bottleneck using the multiple competing hypotheses approach, and finally integrate these results through a sensitivity analysis of a demographic model using life-stage simulation analysis (LSA) to determine the relative importance of genetic, demographic, environmental, and catastrophic processes on population growth. Annual population growth has been slow and variable (1.0 ± 5.2 parrots per year, or an average λ = 1.05 ± 0.19) from 16 parrots (1973) to a high of 40–42 birds (1997–1998). A risk analysis based on population prediction intervals (PPI) indi...

Large-herbivore distribution and abundance: intra-and interspecific niche variation in the tropics

Abstract: Determining the biological and environmental factors that limit the distribution and abundance of organisms is central to our understanding of the niche concept and crucial for predicting how species may respond to large-scale environmental change, such as global warming. However, detailed ecological information for the majority of species has been collected only at a local scale, and insufficient consideration has been given to geographical variation in intraspecific niche requirements. To evaluate the influence of environmental and biological factors on patterns of species distribution and abundance, we conducted a detailed, broadscale study across the tropical savannas of northern Australia on the ecology of three large, sympatric marsupial herbivores (family Macropodidae): the antilopine wallaroo (Macropus antilopinus), common wallaroo (M. robustus), and eastern grey kangaroo (M. giganteus). Using information on species abundance, climate, fire history, habitat, and resource availability, we constructed species' habitat models varying from the level of the complete distribution to smaller regional areas. Multiple factors affected macropod abundance, and the importance of these factors was dependent on the spatial scale of analyses. Fire regimes, water availability, geology, and soil type and climate were most important at the large scale, whereas aspects of habitat structure and interspecific species abundance were important at smaller scales. The distribution and abundance of eastern grey kangaroos and common wallaroos were strongly influenced by climate. Our results suggest that interspecific competition between antilopine wallaroos and eastern grey kangaroos may occur. The antilopine wallaroo and eastern grey kangaroo (grazers) preferred more nutrient-rich soils than the common wallaroo (grazer/browser), which we relate to differences in feeding modes. The abundance of antilopine wallaroos was higher on sites that were burned, whereas the abundance of common wallaroos was higher on unburned sites. Future climate change predicted for Australia has the capacity to seriously affect the abundance and conservation of macropod species in tropical savannas. The results of our models suggest that, in particular, the effects of changing climatic conditions on fire regimes, habitat structure, and water availability may lead to species declines and marked changes in macropod communities.

Intra-individual plant response to moose browsing: feedback loops and impacts on multiple consumers

Abstract: Salix phylicifolia responds to moose browsing with an intra-individual plant response. This plant-animal system is characterized by complex changes in plant chemistry and plant morphology which in turn lead to differences in feeding strategies of future consumers. I postulate that the willow Salix phylicifolia has developed a two-level strategy which results in two different types of twigs (new annual shoots) after moose browsing in winter. This strategy enables the plant to protect reproductive organs by offering two chemically different twigs to herbivores, with catkins concentrated on the less preferred twigs. In addition, it may enable plants to grow out of the reach of browsers. New growth of browsed and unbrowsed twigs on the same individual willow differed in morphology and chemical composition (intra individual response). New growth from browsed twigs had lower concentrations of individual phenolics, a higher biomass (twigs and leaves), and lower concentrations of nitrogen in leaves than the new growth of unbrowsed twigs. This response resulted in a positive feedback loop for moose, which typically rebrowsed the new growth of browsed twigs, whereas the new growth of unbrowsed twigs was avoided. This feeding behavior was consistent for leaf stripping by moose in summer and pruning of twigs in winter. I found no differences in the feeding behavior of herbivorous insects on the leaves growing on the new annual shoots of either browsed or unbrowsed twigs. The new growth of unbrowsed twigs, which were protected by different specific phenolics, had more catkins than the new growth of browsed twigs. In an interindividual comparison, the number of catkins on browsed willows was reduced. Hence, browsing has an impact on willow reproduction and pollinators. Consequently, large herbivores such as moose are likely to be a driving force in shaping patterns of plant response and plant and consumer communities.

Greenfall links groundwater to aboveground food webs in desert river floodplains

Abstract: Groundwater makes up nearly 99% of unfrozen freshwater worldwide and sustains riparian trees rooted in shallow aquifers, especially in arid and semiarid climates. The goal of this paper is to root animals in the regional water cycle by quantifying the significance of groundwater to riparian animals. We focused our efforts on the cricket, Gryllus alogus: a common primary consumer found in floodplain forests along the San Pedro River, in southeast Arizona, USA. Cottonwood trees make groundwater available to G. alogus as dislodged, groundwater-laden leaves (greenfall). We hypothesized that groundwater fluxes mediated by greenfall sustain G. allogus through the prolonged dry season and link these aboveground consumers to belowground aquifers. To test this hypothesis, we first characterized gradients in absolute humidity (air) and water stress in field-collected G. alogus. Absolute humidity declined with distance from river across wide stands of floodplain cottonwood forest during the dry season, but not durin...

Hybrid vigor in a tropical pacific soft-coral community

Abstract: Although hybridization is a relatively widespread phenomenon in marine ecosystems, the ecological importance of hybrids is poorly understood. As crosses between two distinct genotypes, hybrids might express fitness characteristics similar to either parent species, or they might produce unique phenotypic attributes that make them more or less fit than either parent species. We identified a potential hybrid zone among soft corals in a back-reef community on Guam, where the broadcast-spawning species Sinularia maxima and S. polydactyla co-occur. Morphological and chemical traits confirmed the intermediate and unique status of the putative hybrid. Laboratory cross-fertilization experiments using S. maxima and S. polydactyla gametes demonstrated that barriers to hybridization are absent and that the laboratory-reared hybrids developed specific characteristics identified in adult hybrid field populations. Changes in the populations of each parent species and the hybrid were monitored over a decade. While popula...