How much do valley fills influence headwater streams?

Valley fill mining has the potential to alter headwater stream habitat in many areas in the eastern United States. In valley fill mining, overburden is removed to expose underlying coal seams. The overburden is then deposited in the adjacent valley. The deposited overburden from mining increases sedimentation, increases stream conductivity, and alters hydrologic regimes downstream of the fill. Changes in downstream communities are not well documented. However, it was suspected the increased sedimentation and conductivity would have deleterious effects upon the downstream macroinvertebrate communities. In southern West Virginia, four pairs of streams, each consisting of a fill and a reference stream, were selected as representative of watersheds experiencing valley fill mining. Stream pairs were selected for similar environmental conditions, with one stream having a valley fill in its headwaters. Each stream was sampled by replicate Surber samples (n=9 per stream). Water chemistry and sediment measurements also were taken at each location. Valley fill streams experienced significantly higher specific conductance (p < 0.01), but did not have elevated levels of fine sediment. Fills exhibited elevated levels of Na, K, Mn, Mg, Ca, Ni and Fe relative to reference streams. Additionally, valley fill streams demonstrated significantly lower densities of Ephemeroptera, Coleoptera, Odonata, Non-insects, Scrapers, and Shredders (p < 0.03) than reference streams. Further, Ephemeroptera richness was negatively related to specific conductivity and many of the richness metrics were negatively related to metals, both of which were generally elevated in fill streams. It appears that at the minimum, valley fills increase specific conductance and metals in streams and this or some other unqualified factors structure the macroinvertebrate community downstream of the valley fill. However, given the level of disturbance in valley fills, it is surprising how little differences existed between fills and reference stream biota.     

How are arthopod communities structured and why are they so diverse? Answers from Mediterranean mountains using hierarchical additive partitioning

Mountains are complex ecosystems supporting a great variety of taxa. Here, we explored the diversity patterns of arthropods in two mountains, pinpointing the spatial scale that accounts most for overall diversity variation, using an additive partitioning framework. Butterflies and Orthoptera were sampled in Rodopi (2012) and Grammos (2013) mountains. Diversity was partitioned into five hierarchical levels (mountain, elevational zone, habitat, transect and plot). We compared the estimated diversity values for each level to the respective permuted values expected by chance, for all species, as well as for species identified as “rare” or “common”. At broader spatial levels, the variation in total diversity was attributed to the beta diversity component: mountains accounted for 20.94 and 26.25% of butterfly and Orthoptera diversity, and elevational zones accounted for 28.94 and 35.87% respectively. At finer spatial scales, beta diversity was higher than expected by chance in terms of the Shannon index. The type of habitat was found to play a significant role only for rare orthopterans. Finally, common species were recognized for shaping overall species diversity. We highlight the importance of the spatial levels of elevation zone and then mountain position in conservation planning, due to the greater beta diversity recorded at this scale as compared to habitat or more finite scales. Monitoring programs might need to adapt different strategies with respect to the focal organisms, and consider patterns of common rather than rare species that found to drive the patterns of the entire community.     

How many fish populations in Finland are affected by acid precipitation?

Synopsis The number of fish populations affected or lost from small lakes in southern and central Finland due to acid precipitation is estimated. Tolerance limits (pH and labile aluminum) of common fish species were obtained from a fish status survey of 80 lakes. These tolerance values were used to estimate the proportion of affected lakes from the water chemistry data of 783 statistically selected lakes. The proportion of anthropogenically acidified lakes was estimated by calculating pre-acidification pH and aluminum concentrations of the lakes, using a steady-state model based on water chemistry. The number of fish populations for which acidification has affected growth or population structure was estimated at between 2200 and 4400. Out of these, the number of fish populations that have disappeared due to acid precipitation would be about 1000–2000. Almost 60% of the affected or lost populations are roach, Rutilus rutilus, the most sensitive of the common fish species in small lakes in southern and central Finland. Less than 15% of the damaged population is European perch, Perca fluviatilis, the most common species. This is due to the substantially higher tolerance of perch to acidified water in comparison with roach.     

How do chronic nutrient loading and the duration of nutrient pulses affect nutrient uptake in headwater streams?

Our study aimed to analyze the effects of chronic nutrient loading on the capacity of headwater streams to retain phosphorus and ammonium pulses of different duration. For this purpose, we selected nine headwater streams located across a gradient of increasing agricultural land use and eutrophication. In each stream, we performed sequential plateau additions with increasing nutrient concentrations in summer 2015 and instantaneous slug additions in summer 2016 under similar hydrological conditions. We modelled kinetic uptake curves from the slug additions via the Tracer Additions for Spiraling Curve Characterization method and calculated ambient uptake parameters. Ambient uptake rates generally increased (1.4–20.8 µg m^?2 s^?1 for NH₄–N and 0.3–10.3 µg m^?2 s^?1 for SRP, respectively), while ambient uptake velocities decreased from oligotrophic to polytrophic streams (1.8–14.0 mm min^?1 for NH₄–N and 1.6–9.9 mm min^?1 for SRP, respectively). However, correlations between ambient uptake parameters and background concentrations were weak. Concentration-dependent uptake rates followed either a linear or a Michaelis–Menten saturation model, regardless of the degree of nutrient loading. Uptake rate curves showed counter-clockwise hysteresis in oligotrophic streams and clockwise hysteresis in streams of higher trophic states, indicating a reduced significance of hyporheic uptake with increasing nutrient loading. Comparisons of slug and plateau additions revealed that oligotrophic streams were most efficient in uptake during short nutrient pulses, while eutrophic streams profited from longer pulse duration. The results indicate that nutrient uptake is increasingly transport-controlled in polluted streams where increased biofilm thickness and clogging of sediments restrict nutrient transport to reactive sites.     

Do changes in rainfall patterns affect semiarid annual plant communities?

Question: Climate change models forecast a reduction in annual precipitation and more extreme events (less rainy days and longer drought periods between rainfall events), which may have profound effects on terrestrial ecosystems. Plant growth, population and community dynamics in dry environments are likely to be affected by these changes since productivity is already limited by water availability. We tested the effects of reduced precipitation and fewer rain events on three semiarid plant communities dominated by annual species. Location: Three semiarid plant communities from Almería province (SE Spain). Methods: Rain‐out shelters were set up in each community and watering quantity and frequency were manipulated from autumn to early summer. Plant productivity, cover and diversity were measured at the end of the experimental period. Results: We found that a 50% reduction in watering reduced productivity, plant cover and diversity in all three communities. However, neither the 25% reduction in watering nor changes in the frequency of watering events affected these parameters. Conclusions: The lack of response to small reductions in water could be due to the identity and resistance of the plant communities involved, which are adapted to rainfall variability characteristic of arid environments. Therefore, a rainfall reduction of 25% or less may not affect these plant communities in the short term, although higher reductions or long‐term changes in water availability would probably reduce productivity and diversity in these communities.     

Can bryophytes be used to characterize hydrologic permanence in forested headwater streams?

Recent court cases have questioned whether all headwater streams, particularly those that are not perennial, fall within the protective boundaries of the Clean Water Act. Rapid field-based indicators of hydrologic permanence are critically needed for jurisdictional determinations. The study objective was to determine whether characteristics of bryophyte assemblages in forested headwater streams can be useful indicators of hydrologic permanence. We sampled bryophytes from the channel thalweg at 113 sites across 10 forests in the U.S. which varied in hydrologic permanence. Hydrologic permanence was based on hydrologic status during spring (wet season) and late summer (dry season) visits. Perennial sites had flow (surface or visibly interstitial) during both spring and summer visits. Intermittent sites flowed during spring but were either dry or had surface water limited to isolated pools in summer. Ephemeral sites did not have flow during either visit. The species composition significantly differed between ephemeral and perennial stream reaches. The species composition of intermittent streams overlapped with both ephemeral and perennial streams. Three species and six families were identified as indicators of particular permanence classes. Liverworts occurred more frequently at perennial than at intermittent or ephemeral sites. Bryophytes with cushion and turf growth forms were common to ephemeral sites, whereas mat and weft forms were more common to perennial sites. Acrocarpous mosses were more frequent at ephemeral than at perennial sites. The frequencies of bryophyte-moisture associations (derived from the literature) across the three permanence categories were significantly different. Species associated with dry habitats were more frequently encountered at ephemeral than perennial sites, whereas species associated with wet habitats were more frequent at perennial than ephemeral sites. Species richness varied among forests and permanence classes. Ephemeral sites tended to have higher species richness than perennial sites; however, this pattern did not hold for all forests. Bryophytes should be included among the tools for jurisdictional determinations, much like vascular plants are used to help delineate wetlands. Growth forms and higher level taxonomy of bryophytes (i.e., phyla) can be particularly useful for making jurisdictional determinations because of their ability to discriminate ephemeral from intermittent and perennial reaches and they are measures that can be rapidly trained and routinely used by agencies making such determinations.     

How do grazers affect periphyton heterogeneity in streams?

The effects of grazing by stream invertebrates on algal biomass and spatial heterogeneity were tested experimentally in flow-through microcosms with natural substrates (rocks). One experiment tested the effects of fixed densities of three species of grazers (the caddisfly Allomyia sp. and two mayflies, Epeorus deceptivus and Baetis bicaudatus) on periphyton. Baetis was tested with and without chemical cues from fish predators, which reduced grazer foraging activity to levels similar to the less mobile mayfly (Epeorus). Mean algal biomass (chlorophyll a; chl a) was reduced in grazer treatments compared to ungrazed controls, but there were no differences among grazer treatments. Algal heterogeneity (Morisita index) increased with grazer mobility, with the highest heterogeneity occurring in the Baetis-no fish treatment (most mobile grazer) and the lowest in the caddisfly treatment (most sedentary grazer). A second experiment used a three factorial design, and tested whether initial resource distribution (homogeneous vs. heterogeneous), Baetis density (high vs. low) and fish odor (present vs. absent) affected grazer impact on algal resources. Abundances of Baetis and chl a on individual rocks were recorded to explore the mechanisms responsible for the observed distributions of algae. Initial resource heterogeneity was maintained despite being subjected to grazing. Mean chl a was highest in controls, as in experiment I, and effects of Baetis on algal biomass increased with grazer density. There were no fish effects on algal biomass and no effects of grazer density or fish on algal heterogeneity. At the scale of individual rocks Baetis was unselective when food was homogeneously distributed, but chose high-food rocks when it was heterogeneously distributed. Results of these mechanistic experiments showed that Baetis can track resources at the scale of single rocks; and at moderate densities mobile grazers could potentially maintain periphyton distributions observed in natural streams.     

Do fish regulate phytoplankton in shallow eutrophic Northeast Brazilian reservoirs?

SUMMARY 1. In a comparative study, we examined the potential for fish to structure planktonic food webs in shallow mesotrophic to hypereutrophic Northeast Brazilian reservoirs. The food webs were dominated by three guilds of fish (facultative piscivores, generalist planktivores and omnivores), small herbivorous zooplankton and bloom‐forming cyanobacteria, with few littoral macrophytes. 2. A principal component's analysis on data from 13 reservoirs (27 sampling dates in 1995–99) revealed that euphotic depth, the relative density of phytoplankton (i.e. the percentage of overall phytoplankton density) represented by cyanobacteria, and the relative biomass of fish (i.e. percentage of overall biomass) represented by omnivores and facultative piscivores, explained most of the variance in the data. Physico‐chemical conditions, lake morphometry and rainfall were secondary factors. 3. Phytoplankton was related to fish guild structure. Chlorophyll concentration increased with total phosphorus and the relative biomass of omnivorous fish, decreased with the relative biomass of facultative piscivores, but was unrelated to the biomass and mean body size of herbivorous zooplankton. Chlorophyll concentration and the densities of filamentous and colonial cyanobacteria decreased with the ratio of the biomass of facultative piscivores to that of omnivores (FP : OM). 4. We propose two complementary mechanisms for the observed relationships between fish and phytoplankton. At a low biomass of facultative piscivores, juvenile zooplanktivorous fishes may induce a trophic cascade on zooplankton in the littoral zone. Regardless of piscivore biomass, piscivores and omnivores may regulate phytoplankton via multichannel omnivory because of the predominance of omnivorous or detritivorous foraging behaviour. 5. Manipulative experiments are needed to explore further whether, depending on priorities in the use of the reservoir, fisheries management could alter the FP : OM ratio either to enhance fish yields or to reduce phytoplankton densities and cyanobacterial blooms.     

Are biological classifications of headwater streams concordant across multiple taxonomic groups?

1. Studies assessing human impacts on freshwater ecosystems are typically based on a single taxonomic group, often macroinvertebrates or fish. Unfortunately, the degree to which such macroinvertebrate or fish‐based surveys can be generalised across other taxonomic groups remains largely unknown. A prerequisite for useful generalisations is that different taxonomic groups exhibit concordant patterns of community structure across sites. 2. We examined the concordance among fish, benthic macroinvertebrates and bryophytes in 32 streams in a boreal catchment in Finland. Our goal was to test how consistently different taxonomic groups classify stream sites; for example, can site groupings based on macroinvertebrates be used as a surrogate for bryophyte or fish assemblage classification? 3. Our results show that community classifications in headwater streams are not concordant across taxonomic groups, at least not at the within‐river system scale. The lack of concordance reflected the fact that all three groups responded to different environmental factors. Macroinvertebrate community structure was mainly correlated with stream size and pH, whereas bryophytes were related to water colour, nutrient content and in‐stream habitat variability. Fish community structure was best described by stream depth, substrate size and water oxygen concentration. 4. Our results suggest that great care should be taken when typologies based on benthic macroinvertebrates, or any other taxonomic group, are extrapolated to other groups in creating typologies of lotic environments.     

How objective are genera in euascomycetes?

Intra- and intergeneric distances derived from maximum-likelihood phylogenetic trees inferred from 254 nuclear ITS rDNA sequences were examined for seven families of euascomycetes, representing five classes. The intra- and intergeneric distances were well separated in most cases, but the distances varied between families. The analysis of the distance distributions provides a powerful tool for identifying certain taxa with highly deviating distances and thus cases of excessive lumping or splitting. Some cases of lumping and splitting found in different families are briefly discussed. The results of the analysis show that the generic concepts differ between the families. The consequences for nomenclature are discussed and a method abandoning binomial nomenclature while keeping the style of species names is recommended to ensure nomenclatural stability.     

Brazilian Cerrado Folivore and Florivore Caterpillars: How Different are They?

Because of the large differences between flower buds and leaves as food resources we predict that florivore caterpillars compared with folivores would present: (1) higher degree of dietary specificity; (2) higher abundance; and (3) lower species richness. Based on emerged adults from caterpillars collected on leaves and inflorescences our results showed that the florivore fauna was strongly dominated by a few species with higher abundance and dietary specificity than the folivore fauna.     

How resilient are Andean montane forest bird communities to habitat degradation?

The threatened forest habitats of the tropical Andes are reportedly being modified and destroyed 30% faster than their lowland tropical counterparts, but impacts on the hyper-diverse resident avifauna have received little systematic study. We present a baseline analysis of the effects of habitat modification on birds in a lower montane forest landscape in Ecuador, comparing avian community composition in landscape elements subjected to different levels of human modification: primary forest, secondary forest, edge habitat and agricultural land. We use data from a point count survey of 300 counts at 150 sites to test whether community composition and density of birds with different reported habitat preferences and foraging strategies change among landscape elements. Species richness and diversity were lowest in agricultural land, but on some measures, equally low in primary forest. Richness and diversity peaked in secondary forest and edge habitat, but ordination and density analysis revealed clear differences in their species composition. While secondary forest contained mostly forest-preferring species, edge habitat harboured a mix of forest and open-land birds. There was a clearly structured gradient in species composition across landscape elements, with densities of habitat specialists, foraging guilds and families varying considerably from primary forest to agricultural land. Agricultural land was characterised by an assemblage of widespread, abundant species very different from that in core forest habitats. As such, while the majority of montane forest birds appear resilient to a certain level of habitat modification, they cannot persist, and are displaced, where forest has been cleared outright. We argue that, for Andean montane forests, preservation of mature secondary forest offers flexibility in supplementing preserved primary forest areas to provide sufficient habitat for the persistence of this incredibly diverse but severely threatened bird community.     

How heterogeneous are the cloud forest communities in the mountains of central Veracruz, Mexico?

The montane forest in central Veracruz, Mexico is a hotspot of biodiversity. We asked whether lower and upper montane forests could be distinguished in this ecoregion. Variables of vegetation and seasonality in precipitation were tested across 14 sites between 1,250- and 2,550-m elevations. A total of 1,639 individuals and 128 tree species was recorded. There was a unimodal pattern in the richness of species, genera, and families; their richness was positively correlated with precipitation in the wettest quarter of the year, though there were no differences in the basal area and density. Rarefaction, species turnover, nonmetric multidimensional scaling, and a cluster histogram suggest two major groups: lower elevation forests that are less diverse, have low beta diversity and are more similar in composition, with Clethra macrophylla, Liquidambar styraciflua, and Quercus lancifolia as indicator species; and higher elevation forests that are more diverse, have high species turnover, and include forests with Quercus corrugata and Prunus rhamnoides, and forests with Fagus grandifolia, Persea americana, and Ternstroemia sylvatica as indicator species. However, other communities (an Oreomunnea mexicana at the upper site, and a limestone site in the lower forests), exemplify the high regional heterogeneity. We conclude that elevation and seasonality in precipitation produce a directional change in richness and indicator species, but not in vegetation structure. Lower montane forests differed from cloud forests at upper elevations. However, other factors should be included—mainly biogeographic affinities, historic and recent anthropogenic disturbance—to conclusively distinguish them. Montane forest can still be considered very heterogeneous and very high in beta diversity.     

How local is the local field potential?

Local field potentials (LFPs) are of growing importance in neurophysiological investigations. LFPs supplement action potential recordings by indexing activity relevant to EEG, magnetoencephalographic, and hemodynamic (fMRI) signals. Recent reports suggest that LFPs reflect activity within very small domains of several hundred micrometers. We examined this conclusion by comparing LFP, current source density (CSD), and multiunit activity (MUA) signals in macaque auditory cortex. Estimated by frequency tuning bandwidths, these signals' "listening areas" differ systematically with an order of MUA?< CSD?< LFP. Computational analyses confirm that observed LFPs receive local contributions. Direct measurements indicate passive spread of LFPs to sites more than a centimeter from their origins. These findings appear to be independent of the frequency content of the LFP. Our results challenge the idea that LFP recordings typically integrate over extremely circumscribed local domains. Rather, LFPs appear as a mixture of local potentials with "volume conducted" potentials from distant sites.     

Why are there so few fish in the sea?

The most dramatic gradient in global biodiversity is between marine and terrestrial environments. Terrestrial environments contain approximately 75-85% of all estimated species, but occupy only 30 per cent of the Earth's surface (and only approx. 1-10% by volume), whereas marine environments occupy a larger area and volume, but have a smaller fraction of Earth's estimated diversity. Many hypotheses have been proposed to explain this disparity, but there have been few large-scale quantitative tests. Here, we analyse patterns of diversity in actinopterygian (ray-finned) fishes, the most species-rich clade of marine vertebrates, containing 96 per cent of fish species. Despite the much greater area and productivity of marine environments, actinopterygian richness is similar in freshwater and marine habitats (15 150 versus 14 740 species). Net diversification rates (speciation-extinction) are similar in predominantly freshwater and saltwater clades. Both habitats are dominated by two hyperdiverse but relatively recent clades (Ostariophysi and Percomorpha). Remarkably, trait reconstructions (for both living and fossil taxa) suggest that all extant marine actinopterygians were derived from a freshwater ancestor, indicating a role for ancient extinction in explaining low marine richness. Finally, by analysing an entirely aquatic group, we are able to better sort among potential hypotheses for explaining the paradoxically low diversity of marine environments.     

Evaluating detectability of freshwater fish assemblages in tropical streams: Is hand-seining sufficient?

Unprecedented threats to natural ecosystems mean that accurate quantification of biodiversity is a priority, particularly in the tropics which are underrepresented in monitoring schemes. Data from a freshwater fish assemblage in Trinidad were used to evaluate the effectiveness of hand-seining as a survey method in tropical streams. We uncovered large differences in species detectability when hand-seining was used alone, in comparison with when hand-seining and electrofishing were used together. The addition of electrofishing increased the number of individuals caught threefold, and increased the biomass fivefold. Some species were never detected using hand-seining, resulting in significant underestimates of species richness; rarefaction curves suggest that even when hand-seining effort increases, species richness is still underestimated. Diversity indices (Shannon and Simpson index) reveal that diversity was also significantly lower for hand-seined samples. Furthermore, the results of multivariate analyses investigating assemblage structure also differed significantly depending on whether they were based on hand-seined data alone, or a combination of hand-seining and electrofishing. Despite the extra equipment and maintenance required, these findings underline the value of including electrofishing when sampling tropical freshwater streams.     

How will coral reef fish communities respond to climate-driven disturbances? Insight from landscape-scale perturbations

Global climate change is rapidly altering disturbance regimes in many ecosystems including coral reefs, yet the long-term impacts of these changes on ecosystem structure and function are difficult to predict. A major ecosystem service provided by coral reefs is the provisioning of physical habitat for other organisms, and consequently, many of the effects of climate change on coral reefs will be mediated by their impacts on habitat structure. Therefore, there is an urgent need to understand the independent and combined effects of coral mortality and loss of physical habitat on reef-associated biota. Here, we use a unique series of events affecting the coral reefs around the Pacific island of Moorea, French Polynesia to differentiate between the impacts of coral mortality and the degradation of physical habitat on the structure of reef fish communities. We found that, by removing large amounts of physical habitat, a tropical cyclone had larger impacts on reef fish communities than an outbreak of coral-eating sea stars that caused widespread coral mortality but left the physical structure intact. In addition, the impacts of declining structural complexity on reef fish assemblages accelerated as structure became increasingly rare. Structure provided by dead coral colonies can take up to decades to erode following coral mortality, and, consequently, our results suggest that predictions based on short-term studies are likely to grossly underestimate the long-term impacts of coral decline on reef fish communities.     

Are native cyprinids or introduced salmonids stronger regulators of benthic invertebrates in South African headwater streams?

The introduction of nonnative salmonids in the Southern Hemisphere generally leads to a reduction in invertebrate abundance and changes in assemblage composition. In the Cape Floristic Region of South Africa, introduced rainbow trout Oncorhynchus mykiss is the dominant predator in many headwater streams, where they have replaced small‐bodied native fishes such as Breede River redfin Pseudobarbus burchelli. To examine the consequences of this species replacement on food web structure, we used a month‐long field experiment to compare the top‐down effects of Breede River redfin and rainbow trout on benthic invertebrate assemblages (abundance and composition) and basal resources (periphyton and particulate organic matter) in 1 × 1.5 m of plastic cages. Benthic invertebrate abundance was more strongly depleted in the cages with redfin than in the cages with trout, and redfin and trout had distinct effects on invertebrate assemblage composition. On the other hand, neither redfin nor trout had a significant influence over standing stocks of periphyton or organic matter, implying that their differential effects on benthic invertebrates did not cascade down to the base of the stream food web in our experiment. Gut content analysis showed that aquatic invertebrates contributed more to the diet of redfin, while terrestrial invertebrates contributed more to the diet of trout, which may be responsible for the relatively weak effect of trout on aquatic invertebrates. This pattern contrasts with nonnative salmonid impacts elsewhere in the Southern Hemisphere. That trout can strongly alter the structure of benthic invertebrate assemblages, in addition to severely depleting native fish abundance, in Cape Floristic Region headwater streams should be weighed into management decisions, and our findings highlight the need for a detailed understanding of species‐specific top‐down effects where native predators are replaced by invasive predators.     

Are populations of mayflies living in adjacent fish and fishless streams genetically differentiated?

1. Conspecific populations living in habitats with different risks of predation often show phenotypic variation in defensive traits. Traits of two species of mayflies (Baetidae: Baetis bicaudatus and Baetis sp. nov.) differ between populations living in fish and fishless streams in a high altitude drainage basin in western Colorado, U.S.A. We tested for genetic differentiation between mayfly populations in these two habitat types, assuming that lack of genetic differentiation would be consistent with the hypothesis that those traits are phenotypically plastic. 2. Previous work has shown that larvae of both species behave differently and undergo different developmental pathways in adjacent fish and fishless streams. These phenotypic differences in behaviour and development have been induced experimentally, suggesting that populations from fishless streams have the genetic capability to respond to fish. 3. During summer 2001 we collected Baetis larvae from several fish and fishless streams, and from fish and fishless sections of the same streams. We used allozymes and a fragment of the cytochrome oxidase subunit 1 mitochondrial gene to examine genetic variation of Baetis individuals within and among streams. 4. Results showed that genetic variation exists among populations of the same species of Baetis from different streams, but none of that variation was associated with the presence or absence of fish. These data confirm that populations of Baetis living in fish and fishless streams are not genetically distinct, and are consistent with the hypothesis that traits associated with environments of different risk are phenotypically plastic.