Variability in grazer-mediated defensive responses of green and red macroalgae on the south coast of South Africa

Variabilities in the responses of several South African red and green macroalgae to direct grazing and the responses of one green alga to cues from grazers were tested. We used two feeding experiments: (1) testing the induced responses of three red and one green algae to direct grazing by mesograzers and (2) a multi-treatment experiment, in which the direct and indirect effects of one macrograzer species on the green alga Codium platylobium were assessed. Consumption rates were assessed in feeding assays with intact algal pieces and with agar pellets containing non-polar extracts of the test algae. Defensive responses were induced for intact pieces of Galaxaura diessingiana, but were not induced in pellets, suggesting either morphological defence or chemical defence using polar compounds other than polyphenols. In contrast, exposure to grazing stimulated consumption of Gracilaria capensis and Hypnea spicifera by another grazing species. In the multi-treatment experiment, waterborne cues from both grazing and non-grazing snails induced defensive algal traits in C. platylobium. We suggest that inducible defences among macroalgae are not restricted to brown algae, but that both the responses of algae to grazers and of grazers to the defences of macroalgae are intrinsically variable and complex.     

Extrinsic and intrinsic determinants of winter foraging and breeding phenology in a temperate seabird

In temperate regions, winter presents animals with a number of challenges including depressed food abundance, increased daily energy requirements, higher frequency of extreme weather events and shortened day length. Overcoming these constraints is critical for overwintering survival and scheduling of future breeding of long-lived species and is likely to be state dependent, associated with intrinsic abilities such as food acquisition rates. We examined the relationship between environmental and intrinsic factors on overwintering foraging and subsequent breeding phenology of the European shag Phalacrocorax aristotelis, a diurnal marine predator. We tested a range of hypotheses relating to overwintering foraging time and location. We found that individuals greatly increased their foraging time in winter to a peak of more than 90% of available daylight at the winter solstice. The seasonal patterns of foraging time appear to be driven by a combination of light levels and weather conditions and may be linked to the availability of the shag's principal prey, the lesser sandeel Ammodytes marinus. There was no evidence that shags dispersed south in winter to increase potential foraging time. Foraging time decreased after the winter solstice and, crucially, was correlated with subsequent breeding phenology, such that individuals that spent less time foraging in February bred earlier. The relationship was much stronger in females than males, in line with their more direct control of timing of breeding. Our results demonstrate that pre-breeding intrinsic foraging ability is critical in determining breeding phenology.     

A trait-based test for habitat filtering: convex hull volume

Community assembly theory suggests that two processes affect the distribution of trait values within communities: competition and habitat filtering. Within a local community, competition leads to ecological differentiation of coexisting species, while habitat filtering reduces the spread of trait values, reflecting shared ecological tolerances. Many statistical tests for the effects of competition exist in the literature, but measures of habitat filtering are less well-developed. Here, we present convex hull volume, a construct from computational geometry, which provides an n-dimensional measure of the volume of trait space occupied by species in a community. Combined with ecological null models, this measure offers a useful test for habitat filtering. We use convex hull volume and a null model to analyze California woody-plant trait and community data. Our results show that observed plant communities occupy less trait space than expected from random assembly, a result consistent with habitat filtering.     

Jellyfish aggregations and leatherback turtle foraging patterns in a temperate coastal environment

Leatherback turtles (Dermochelys coriacea) are obligate predators of gelatinous zooplankton. However, the spatial relationship between predator and prey remains poorly understood beyond sporadic and localized reports. To examine how jellyfish (Phylum Cnidaria: Orders Semaeostomeae and Rhizostomeae) might drive the broad-scale distribution of this wide ranging species, we employed aerial surveys to map jellyfish throughout a temperate coastal shelf area bordering the northeast Atlantic. Previously unknown, consistent aggregations of Rhizostoma octopus extending over tens of square kilometers were identified in distinct coastal "hotspots" during consecutive years (2003-2005). Examination of retrospective sightings data (>50 yr) suggested that 22.5% of leatherback distribution could be explained by these hotspots, with the inference that these coastal features may be sufficiently consistent in space and time to drive long-term foraging associations.     

Whole-system nutrient enrichment increases secondary production in a detritus-based ecosystem

Although the effects of nutrient enrichment on consumer-resource dynamics are relatively well studied in ecosystems based on living plants, little is known about the manner in which enrichment influences the dynamics and productivity of consumers and resources in detritus-based ecosystems. Because nutrients can stimulate loss of carbon at the base of detrital food webs, effects on higher consumers may be fundamentally different than what is expected for living-plant-based food webs in which nutrients typically increase basal carbon. We experimentally enriched a detritus-based headwater stream for two years to examine the effects of nutrient-induced changes at the base of the food web on higher metazoan (predominantly invertebrate) consumers. Our paired-catchment design was aimed at quantifying organic matter and invertebrate dynamics in the enriched stream and an adjacent reference stream for two years prior to enrichment and two years during enrichment. Enrichment had a strong negative effect on standing crop of leaf litter, but no apparent effect on that of fine benthic organic matter. Despite large nutrient-induced reductions in the quantity of leaf litter, invertebrate secondary production during the enrichment was the highest ever reported for headwater streams at this Long Term Ecological Research site and was 1.2-3.3 times higher than predicted based on 15 years of data from these streams. Abundance, biomass, and secondary production of invertebrate consumers increased significantly in response to enrichment, and the response was greater among taxa with larval life spans < or = 1 yr than among those with larval life spans >1 yr. Production of invertebrate predators closely tracked the increased production of their prey. The response of invertebrates was largely habitat-specific with little effect of enrichment on food webs inhabiting bedrock outcrops. Our results demonstrate that positive nutrient-induced changes to food quality likely override negative changes to food quantity for consumers during the initial years of enrichment of detritus-based stream ecosystems. Longer-term enrichment may impact consumers through eventual reductions in the quantity of detritus.     

Cicada emergence in Southwestern riparian forest: influences of wildfire and vegetation composition.

Annually emerging cicadas are a numerically and ecologically dominant species in Southwestern riparian forests. Humans have altered disturbance regimes that structure these forests such that floods are less common and wildfires occur more frequently than was historically the case. Impacts of these changes on primary consumers such as riparian cicadas are unknown. Because cicadas are consumed by a variety of animal species, disturbances that alter timing of their emergence or abundance could have consequences for species at higher trophic levels. We trapped emerging cicadas (Tibicen dealbatus) in burned and unburned riparian forest plots along the Middle Rio Grande in central New Mexico (USA) to determine effects of wildfire and vegetation structure on their density and phenology. We measured vegetation variables and soil temperature at cicada traps and related these variables to variation in emergence density and phenology. We also experimentally heated soil under emergence traps to examine the relationship between soil temperature and emergence phenology. Emergence density was similar in wildfire and unburned plots, though emergence date averaged earlier in wildfire plots and experimentally heated traps. We identified models containing cottonwood proximity (distance from the nearest cottonwood tree) and cottonwood canopy coverage as the most parsimonious explanations of emergence density at each trap. Model selection results were consistent with the literature and field observations that showed that cottonwood trees are an essential resource for T. dealbatus. Cottonwood canopy was also correlated with low soil temperatures, which are associated with later emergence dates. Failure of cottonwoods to reestablish following wildfire could result in cicadas emerging at lower densities and at earlier dates. For cicadas to emerge at densities and times that provide the greatest benefits to birds and other riparian-obligate secondary consumers, riparian forests should be protected from fire, and native vegetation in wildfire sites should be restored.     

Mechanisms generating modification of benthos following tidal flat invasion by a Spartina hybrid.

Many coastal habitats are being substantially altered by introduced plants. In San Francisco Bay, California, USA, a hybrid form of the eastern cordgrass Spartina alterniflora is rapidly invading open mudflats in southern and central sections of the Bay, altering habitat, reducing macrofaunal densities, and shifting species composition. The invasion has resulted in significant losses of surface-feeding amphipods, bivalves, and cirratulid polychaetes, while subsurface feeding groups such as tubificid oligochaetes and capitellid polychaetes have been unaffected. In the present paper, we document the causes and mechanisms underlying the changes observed. Through a series of in situ manipulative experiments we examined the influence of hybrid Spartina canopy on a range of physical, chemical, and biological properties. The hybrid Spartina canopy exerted a strong influence on the hydrodynamic regime, triggering a series of physical, chemical, and biological changes in the benthic system. Relative to tidal flats, water velocity was reduced in hybrid patches, promoting deposition of fine-grained, organic-rich particles. The resulting changes in the sediment environment included increased porewater sulfide concentrations and anoxia, which led to poor survivorship of surface feeders such as bivalves, amphipods, and polychaetes. These are key taxa that support higher trophic levels including migratory shorebirds that feed on tidal flats. Altered flow in the Spartina canopy further contributed to changes in barnacle recruitment and resuspension of adult benthic invertebrates. Increased crab-induced predation pressure associated with Spartina invasion also contributed to changes in benthic invertebrate communities. Our results suggest that multiple physical, chemical, biotic, and trophic impacts of the Spartina invasion have resulted in substantial changes in benthic communities that are likely to have important effects on the entire ecosystem.     

Multi-model analysis of tree competition along environmental gradients in southern New England forests.

Robust predictions of competitive interactions among canopy trees and variation in tree growth along environmental gradients represent key challenges for the management of mixed-species, uneven-aged forests. We analyzed the effects of competition on tree growth along environmental gradients for eight of the most common tree species in southern New England and southeastern New York using forest inventory and analysis (FIA) data, information theoretic decision criteria, and multi-model inference to evaluate models. The suite of models estimated growth of individual trees as a species-specific function of average potential diameter growth, tree diameter at breast height, local environmental conditions, and crowding by neighboring trees. We used ordination based on the relative basal area of species to generate a measure of site conditions in each plot. Two ordination axes were consistent with variation in species abundance along moisture and fertility gradients. Estimated potential growth varied along at least one of these axes for six of the eight species; peak relative abundance of less shade-tolerant species was in all cases displaced away from sites where they showed maximum potential growth. Our crowding functions estimate the strength of competitive effects of neighbors; only one species showed support for the hypothesis that all species of competitors have equivalent effects on growth. The relative weight of evidence (Akaike weights) for the best models varied from a low of 0.207 for Fraxinus americana to 0.747 for Quercus rubra. In such cases, model averaging provides a more robust platform for prediction than that based solely on the best model. We show that predictions based on the selected best models dramatically overestimated differences between species relative to predictions based on the averaged set of models.     

Modeling species-habitat relationships with spatially autocorrelated observation data.

Spatial autocorrelation in wildlife observation data arises when extrinsic environmental processes and patterns that influence the spatial distribution of wildlife are themselves spatially structured, or when species are subject to intrinsic population processes, causing contagion or dispersion effects. Territoriality, Allee effects, dispersal limitations, and social clustering are examples of intrinsic processes. Both forms of autocorrelation can violate the assumptions of generalized linear regression models, resulting in biased estimation of model coefficients and diminished predictive performance. Such consequences may be avoided for extrinsic autocorrelation when autocorrelated environmental variables are available for use as model covariates, whereas intrinsic spatial autocorrelation requires an alternative modeling approach. The autologistic model provides an approach suited to the binary observations often obtained in wildlife surveys, but its performance has not been tested across widely varying sampling intensities or strengths of intrinsic spatial structure. Here we use simulated data to test the autologistic model under a range of sampling conditions. The autologistic model obtains better fits and substantially better predictive performance than the standard logistic regression model over the full range of sampling designs and intensities tested. We provide a simple Bayesian implementation of the autologistic model, which until now has not been achieved with standard statistical software alone. A step-by-step procedure is given for characterizing and modeling spatial autocorrelation in binary observation data, along with computer code for fitting autologistic models in WinBUGS, a freeware Bayesian analysis package. This approach avoids normal approximations to the pseudo-likelihood, in contrast to previous Bayesian applications of the autologistic model. We provide a sample application of the autologistic model, fitted to survey data for a gliding marsupial in southeastern Australia.