Ryan S. King

Two statistical methods for the detection of environmental thresholds

A nonparametric method and a Bayesian hierarchical modeling method are proposed in this paper for the detection of environmental thresholds. The nonparametric method is based on the reduction of deviance, while the Bayesian method is based on the change in the response variable distribution parameters. Both methods are tested using macroinvertebrate composition data from a mesocosm experiment conducted in the Everglades wetlands, where phosphorus is the limiting nutrient. Using the percent of phosphorus tolerant species and a dissimilarity index as the response variables, both methods resulted in a similar and well-defined TP concentration threshold, with a distribution function that can be used to determine the probability of exceeding the threshold.

Evaluating Subsampling Approaches and Macroinvertebrate Taxonomic Resolution for Wetland Bioassessment

Methods for wetland bioassessment using macroinvertebrates are not well developed. Two of the most controversial issues in stream bioassessment, subsampling and taxonomic resolution, have yet to be quantitatively addressed for wetlands. Using a multivariate approach, we evaluated the efficacy of family-, genus-, and species-level assemblage data in reflecting the environment and distinguishing impaired sites from the reference condition. We used 5 basic levels of subsampling (100-, 200-, and 300-organism fixed counts; 10% and 25% fixed areas), an integrated subsample requiring a minimum fixed count and fixed area (100&10%), and 100-count and 10%-area subsamples coupled with a supplementary large-rare (LR) search. Data were obtained from 1.5-m2 composite samples collected from 126 plots along a 10-km-long eutrophication gradient in the Florida Everglades. Our results suggest that effectiveness of subsampling depended more upon the minimum number of individuals retained than minimum area or proportion of the sample picked. Fixed-area subsamples were generally less efficient than fixed counts, with 200- and 300-individual fixed counts resulting in significantly greater assemblage-environment relationships and much higher accuracy in detecting impairment than 10% fixed area, despite averaging similar numbers of individuals. The greatest improvement with increasing subsample size was observed between fixed counts of 100 and 200 individuals; detecting impairment, in particular, was not markedly improved with subsample sizes >200 individuals. Supplementing subsamples with a LR search resulted in only very slight improvements in assemblage-environment relationships, but was effective in improving prediction accuracy, particularly for family-level data. However, family-level assemblage-environment relationships and abilities to detect impairment were inferior to genus- and species-level data, regardless of subsample size. Species-level data performed best, primarily because of the large proportion (>20%) of total species belonging to Chironomidae. The potential importance of Chironomidae to wetland bioassessment was further revealed through an evaluation of a tiered-taxonomic approach, which showed that non-Chironomidae family-level data tiered with species-level Chironomidae data produced results very similar to those obtained using genus- or species-level data exclusively. Our results suggest that fixed counts ≥200 or integrated fixed-area/fixed-count approaches that consistently obtain a minimum of 200 individuals should be considered as minimum subsample sizes for wetlands. We additionally advocate LR searches and recommend genus- or species-level taxonomy, particularly for the Chironomidae.

Thresholds, breakpoints, and nonlinearity in freshwaters as related to management

Abstract Nonlinear ecological responses to anthropogenic forcing are common, and in some cases, the ecosystem responds by assuming a new stable state. This article is an overview and serves as the introduction to several articles in this BRIDGES cluster that are directed toward managers interested in dealing with nonlinear responses in freshwaters, particularly streams. A threshold or breakpoint occurs where the system responds rapidly to a relatively small change in a driver. The existence of a threshold can signal a change in system configuration to an alternative stable state, although such a change does not occur with all thresholds. In general, a mechanistic understanding of ecological dynamics is required to predict thresholds, where they will occur, and if they are associated with the occurrence of alternative stable states. Thresholds are difficult to predict, although a variety of univariate methods has been used to indicate thresholds in ecological data. When we applied several methods to one type of response variable, the resulting threshold values varied 3-fold, indicating that more research on detection methods is necessary. Numerous case studies suggest that the threshold concept is important in all ecosystems. Managers should be aware that human actions might result in undesirable rapid changes and potentially an unwanted alternative stable state, and that recovery from that state might require far more resources and time than avoiding entering the state in the first place would have required. Given the difficulties in predicting thresholds and alternative states, the precautionary approach to ecosystem management is probably the most prudent.

How Many Mountains Can We Mine? Assessing the Regional Degradation of Central Appalachian Rivers by Surface Coal Mining

Surface coal mining is the dominant form of land cover change in Central Appalachia, yet the extent to which surface coal mine runoff is polluting regional rivers is currently unknown. We mapped surface mining from 1976 to 2005 for a 19,581 km(2) area of southern West Virginia and linked these maps with water quality and biological data for 223 streams. The extent of surface mining within catchments is highly correlated with the ionic strength and sulfate concentrations of receiving streams. Generalized additive models were used to estimate the amount of watershed mining, stream ionic strength, or sulfate concentrations beyond which biological impairment (based on state biocriteria) is likely. We find this threshold is reached once surface coal mines occupy >5.4% of their contributing watershed area, ionic strength exceeds 308 μS cm(-1), or sulfate concentrations exceed 50 mg L(-1). Significant losses of many intolerant macroinvertebrate taxa occur when as little as 2.2% of contributing catchments are mined. As of 2005, 5% of the land area of southern WV was converted to surface mines, 6% of regional streams were buried in valley fills, and 22% of the regional stream network length drained watersheds with >5.4% of their surface area converted to mines.

Threshold effects of coastal urbanization onPhragmites australis (common reed) abundance and foliar nitrogen in Chesapeake Bay

The invasion of North American tidal marshes byPhragmites australis, or common reed, is a large-scale ecological problem that has been primarily studied at small spatial scales. Previous local-scale studies have provided evidence that the expansion ofPhragmites is facilitated by disturbance and increased nitrogen (N) associated with agricultural and urban-suburban (developed) land uses along wetland-upland borders. We tested the generality of previous findings across a larger spatial scale and wider range of environmental conditions in Chesapeake Bay, the largest estuarine ecosystem in the USA. We sampled 90 tidal wetlands nested within 30 distinct subestuarine watersheds and examined the relationship between land use andPhragmites abundance and foliar N, an indicator of nitrogen availability. We estimated land use adjacent to wetland borders and within subestuary watersheds and explored the importance of spatial proximity by weighting land use by its distance from the wetland border or subestuary shoreline, respectively. Regression tree and changepoint analyses revealed thatPhragmites abundance sharply increased in almost every wetland where development adjacent to borders exceeded 15%. Where development was <15% but natural land cover at the near the subestuary shoreline was low (<∼35%),Phragmites was abundant, suggesting that wetlands in highly modified watersheds also were susceptible to invasion, regardless of land use adjacent to wetlands.Phragmites foliar N was markedly elevated in watersheds with >14–22% shoreline development, the same level of development that corresponded to high levels of invasion. Our results suggest that development near wetlands is at least partially responsible for patterns of invasion across Chesapeake Bay. Larger-scale phenomena, such as nitrogen pollution at the watershed-subestuary scale, also may be facilitating invasion. Urbanization near coastlines appears to play an important role in the invasion success ofPhragmites in coastal wetlands of Chesapeake Bay and probably much of eastern North America.

How novel is too novel? Stream community thresholds at exceptionally low levels of catchment urbanization

Novel physical and chemical conditions of many modern ecosystems increasingly diverge from the environments known to have existed at any time in the history of Earth. The loss of natural land to urbanization is one of the most prevalent drivers of novel environments in freshwaters. However, current understanding of aquatic community response to urbanization is based heavily upon aggregate indicators of community structure and linear or wedge-shaped community response models that challenge ecological community theory. We applied a new analytical method, threshold indicator taxa analysis (TITAN), to a stream biomonitoring data set from Maryland to explicitly evaluate linear community response models to urbanization that implicitly assume individual taxa decline or increase at incrementally different levels of urbanization. We used TITAN (1) to identify the location and magnitude of greatest change in the frequency and abundance of individual taxa and (2) to assess synchrony in the location of change points as evidence for stream community thresholds in response to percent impervious cover in catchments. We documented clear and synchronous threshold declines of 110 of 238 macroinvertebrate taxa in response to low levels of impervious cover. Approximately 80% of the declining taxa did so between 0.5% and 2% impervious cover, whereas the last 20% declined sporadically from 2% to 25% impervious cover. Synchrony of individual responses resulted in distinct community-level thresholds ranging from < or = 0.68% (mountains), 1.28% (piedmont), and 0.96% (coastal plain) impervious cover. Upper limits (95% confidence intervals) of community thresholds were < 2% cover in all regions. Within distinct physiographic classes, higher-gradient, smaller catchments required less impervious cover than lower gradient, larger catchments to elicit community thresholds. Relatively few taxa showed positive responses to increasing impervious cover, and those that did gradually increased in frequency and abundance, approximating a linear cumulative distribution. The sharp, synchronous declines of numerous taxa established a consistent threshold response at exceptionally low levels of catchment urbanization, and uncertainty regarding the estimation of impervious cover from satellite data was mitigated by several corroborating lines of evidence. We suggest that threshold responses of communities to urban and other novel environmental gradients may be more prevalent than currently recognized.

Spatial Dependency of Vegetation- Environment Linkages in an Anthropogenically Influenced Wetland Ecosystem

AbstractManagement and restoration of vegetation patterns in ecosystems depends on an understanding of allogenic environmental factors that organize species assemblages and autogenic processes linked to assemblages. However, our ability to make strong inferences about vegetation–environment linkages in field studies is often limited due to correlations among environmental variables, spatial autocorrelation, and scale dependency of observations. This is particularly true in large, heterogeneous ecosystems such as the Everglades. Here, an extensive canal-and-levee system has modified historical fire regimes and hydropatterns while contributing large inputs of surface-water phosphorus (P), nitrogen (N) and cations such as sodium (Na). Some of these anthropogenic influences have been implicated as factors leading to the shift of sawgrass (Cladium jamaicense Crantz) and slough communities to an assemblage of weedy species such as cattail (Typha domingensis Pers.). To untangle the independent effect of multiple variables, we used a spatially explicit, multivariate approach to identify linkages among spatial patterns, environmental factors, and vegetation composition along a 10-km gradient of anthropogenic influence in the Everglades, an area immediately downstream from canal inflow structures. Clusters of plots were stratified among three zones (Impacted, Transition, and Reference), a design that allowed us to contrast vegetation–environment linkages and spatial patterns at multiple scales and degrees of ecosystem alteration. Along the 10-km gradient, partial Mantel tests showed that nutrients (phosphorus, nitrogen, and potassium) and hydropattern (frequency of dryness) were independently linked to patterns in fine-scale vegetation composition, but phosphorus was the only environmental variable linked to patterns of coarse-scale composition. Regardless of scale, the effect of distance from canal inflows accounted for variation in vegetation that could not be explained by other variables. A significant residual effect of spatial proximity among sampling locations also was detected and was highly suggestive of dispersal or other spatial determinants of vegetation pattern. However, this pure spatial effect was significantly stronger in the Transition and Impacted zones than in the Reference zone—fine-scale environmental variables explained all of the spatial structure in vegetation in the Reference zone. A further examination of spatial patterns in vegetation by using Mantel correlograms revealed significant heterogeneity at fine, local scales in the Reference zone, but this pattern progressively degraded toward homogeneity among closely neighboring locations in the Impacted zone. However, the fine-scale vegetation pattern in the Reference zone was hierarchically nested at a broader scale and yielded a similar coarse pattern across the landscape, whereas the coarse pattern in the Transition and Impacted zones was relatively heterogeneous and fragmented. Collectively, these results indicate that allogenic spatial and environmental factors related to the canal system have disrupted the coupling between pattern and process by altering fine-scale vegetation–environment linkages and spatial patterns characteristic of the natural Everglades ecosystem.

Considerations for analyzing ecological community thresholds in response to anthropogenic environmental gradients

Abstract The goal of this paper is to help managers better understand implications of using aggregate community metrics, such as taxon richness or Indices of Biotic Integrity (IBI), for detecting threshold responses to anthropogenic environmental gradients. To illustrate, we offer an alternative analytical approach, Threshold Indicator Taxa ANalysis (TITAN), geared toward identifying synchronous changes in the distribution of multiple taxa as evidence of an ecological community threshold. Our approach underscores the fundamental reality that which taxa are affected by stressors is important, both from a conservation standpoint and because taxon-specific life-history traits help us understand relevant mechanisms. First, we examine macroinvertebrate community response to an impervious cover gradient using a well-studied biomonitoring data set to show that representative community metrics are relatively insensitive to synchronous threshold declines of numerous individual taxa. We then reproduce these response relationships using a simulated community data set with similar properties to demonstrate that linear or wedge-shaped responses of community metrics to anthropogenic gradients can occur as an artifact of aggregating multiple taxa into a single value per sampling unit, despite strong nonlinearity in community response. Our findings do not repudiate the use of community metrics or multimetric indices, but they challenge assumptions that such metrics are capable of accurately reflecting community thresholds across a broad range of anthropogenic gradients. We recommend an alternative analysis framework that begins with characterization of the responses of individual taxa and uses aggregation only after distinguishing the magnitude, direction, and uncertainty in the responses of individual members of the community.

Does nutrient enrichment decouple algal–bacterial production in periphyton?

Abstract Coupled production between algae and bacteria in stream epilithon was assessed along a nutrient-enrichment gradient in 8 Texas streams with open canopies. Photosynthesis (PS) and bacterial biomass production (BBP) were measured simultaneously using a dual-label radioassay (14C-HCO3– uptake and 3H-L-leucine incorporation into protein) on multiple samples within a stream reach. PS and BBP were measured after light (1200–1500 μmol m−2 s−1) and dark incubations. The degree of coupled production between algae and bacteria within a stream was estimated as the covariation (i.e., correlation or covariance) between PS and BBP derived from unshaded replicates in each stream. Streamwater nutrients ranged from 0.18 to 8.1 mg/L total N and 0.009 to 2.0 mg/L total P. Epilithon N and P content (as % dry mass) and C:N:P ratios varied widely among streams and were positively correlated with streamwater nutrient concentrations. Mean BBP measured in light incubations (BBPL) was greater than mean BBP measured in dark incubations (BBPD), and the difference between the 2 means (BBPL – BBPD) was positively correlated with mean PS among streams (R2 = 0.53). Covariance between PS and BBPL within streams (COVPS–BBP) decreased as epilithon nutrient content increased. COVPS–BBP was positively correlated with both epilithon C:N (R2 = 0.78) and C:P (R2 = 0.77) among streams. These results suggest that algal and bacterial production are decoupled by nutrient enrichment, and that algae might rely more heavily on bacterial-regenerated nutrients than on streamwater nutrients to support production in nutrient-poor streams.

Nitrogen fixation and phosphatase activity in periphyton growing on nutrient diffusing substrata: evidence for differential nutrient limitation in stream periphyton

Abstract We explored N2 fixation and alkaline phosphatase activity (APA) in periphyton from a N-limited stream ecosystem by coupling measurements of these processes with nutrient diffusion substrata (NDS) experiments. We measured periphyton biomass accumulation (as ash-free dry mass [AFDM] and chlorophyll a [CHLA]), N2 fixation, and APA to evaluate the relative importance of N2 fixation as an N source to the periphyton community and APA as an indicator of P deficiency in a seemingly N-limited system. We used fritted-glass-disc NDS and estimated AFDM, CHLA, N2 fixation, and APA on days 6, 18, and 29 after deployment. Periphyton AFDM steadily increased on NDS over time, but was not influenced by nutrients. CHLA was elevated in the N treatment on days 18 and 29, indicating autotrophic N limitation. Consistent with N limitation, N2 fixation was high but not different in the control and P treatments and was virtually undetectable on the N treatment. N2 fixation in control and P treatments was detectable in both light and dark incubations, and dark rates were 4 to 73% of the light rates on days 18 and 29. The average contribution of total N2 fixation to periphyton in control and P treatments was 0.93 mg N/m2 on day 18 and 1.0 mg N/m2 on day 29. APA was significantly elevated on the control and was highest in the N treatment despite no apparent P limitation of periphyton biomass accumulation. P enrichment always decreased APA. Measurable N2 fixation and the change in CHLA suggest that autotrophs were primarily N limited. However, APA observed in controls demonstrated that some portion of the periphyton community was experiencing P deficiency. This result suggests that periphyton metabolism was related to both N and P availability, but that biomass accumulation might have been limited primarily by N. One explanation for these findings is that different organisms, perhaps occupying different trophic positions within the community, might have been limited by different elements.