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Module 4: Basic biology of wetland animals

An assortment of animals inhabit wetlands, with many invertebrates, amphibians and reptiles, fishes, birds, and some mammals being wetland specialists. Wetlands provide ample water, food, and cover that benefits resident animals, but periodic high floods, frequent drying, and harsh water qualities (low-oxygen, acidic, saline) can stress wetland animals. Animals that rely on wetlands have developed numerous adaptations to cope with these stresses. The animal biodiversity in wetlands is particularly valued by people, from deep historical and cultural connections, to modern ecotourists and bird watchers who observe it and hunters and anglers who appreciate the consumptive resources they provide. Animals such as alligators and beavers create or physically-modify wetland environments; this activity is called ecosystem engineering and makes these animals among the most important organisms to wetland ecology. Wetlands are high in biodiversity and support rich communities of wetland invertebrates, are primary breeding habitats for most amphibians, and nearly all migratory birds rely on wetlands during some part of their annual cycle. The interactions of animals with plants (herbivory, cutting) and each other (predation) strongly shape wetland environments and affect nutrient cycling. Human impacts on wetlands can negatively affect many animals through habitat loss and degradation, resulting in many wetland amphibians, reptiles, and birds being listed as threatened and endangered species. Thus, wetland conservation is vital to conserve many animal species.

Wetland Science & Practice↗

Natural disturbance shapes benthic intertidal macroinvertebrate communities of high latitude river deltas

Unlike lower latitude coastlines, the estuarine nearshore zones of the Alaskan Beaufort Sea are icebound and frozen up to 9 months annually. This annual freezing event represents a dramatic physical disturbance to fauna living within intertidal sediments. The main objectives of this study were to describe the benthic communities of Beaufort Sea deltas, including temporal changes and trophic structure. Understanding benthic invertebrate communities provided a baseline for concurrent research on shorebird foraging ecology at these sites. We found that despite continuous year-to-year episodes of annual freezing, these estuarine deltas are populated by a range of invertebrates that represent both marine and freshwater assemblages. Freshwater organisms like Diptera and Oligochaeta not only survive this extreme event, but a marine invasion of infaunal organisms such as Amphipoda and Polychaeta rapidly recolonizes the delta mudflats following ice ablation. These delta sediments of sand, silt, and clay are fine in structure compared to sediments of other Beaufort Sea coastal intertidal habitats. The relatively depauperate invertebrate community that ultimately develops is composed of marine and freshwater benthic invertebrates. The composition of the infauna also reflects two strategies that make life on Beaufort Sea deltas possible: a migration of marine organisms from deeper lagoons to the intertidal and freshwater biota that survive the 9-month ice-covered period in frozen sediments. Stable isotopic analyses reveal that both infaunal assemblages assimilate marine and terrestrial sources of organic carbon. These results provide some of the first quantitative information on the infaunal food resources of shallow arctic estuarine systems and the long-term persistence of these invertebrate assemblages. Our data help explain the presence of large numbers of shorebirds in these habitats during the brief summer open-water period and their trophic importance to migrating waterfowl and nearshore populations of estuarine fishes that are the basis of subsistence lifestyles by native inhabitants of the Beaufort Sea coast.

Estuaries and Coasts↗

Increased mercury and reduced insect diversity in linked stream-riparian food webs downstream of a historical mercury mine

Historical mining left a legacy of abandoned mines and waste rock in remote headwaters of major river systems in the western United States. Understanding the influence of these legacy mines on culturally and ecological important downstream ecosystems is not always straight-forward because of elevated natural levels of mineralization in mining-impacted watersheds. To test the ecological effects of historic mining in the headwaters of the upper Salmon River watershed (USA), we measured multiple community and chemical endpoints in downstream linked aquatic-terrestrial food webs. Mining inputs impacted downstream food webs through increased mercury accumulation and decreased insect biodiversity. Total mercury (THg) in seston, aquatic insect larvae, adult aquatic insects, riparian spiders, and fish at sites up to 7.6 km downstream of mining was in much higher concentrations (1.3 to 11.3-fold) and isotopically distinct compared with sites immediately upstream of mining inputs. Methylmercury (MeHg) concentrations in bull trout and riparian spiders were sufficiently high (732 – 918 and 347 – 1,140 ng MeHg g -1 dw) to affect humans, birds, and piscivorous fish. Furthermore, the alpha-diversity of benthic insects was locally depressed by 12-20% within 4.3 to 5.7 km downstream of from the mine. However, because total insect biomass was not affected by mine inputs, the mass of mercury in benthic insects at a site (i.e., ng Hg m -2 ) was extremely elevated downstream (10 – 1,778-fold) compared with directly upstream of mining inputs. Downstream adult aquatic insect-mediated fluxes of total mercury were also high (~16 ng THg m -2 d -1 ). Abandoned mines can have ecologically important effects on downstream communities, including reduced biodiversity and increased mercury flux to higher order consumers, including fish, birds, and humans.

Idaho↗

Aleuts, sea otters, and alternate stable state communities

Reexamination of stratified faunal components of a prehistoric Aleut midden excavated on Amchitka Island, Alaska, indicates that Aleut prey items changed dramatically during 2500 years of aboriginal occupation. Recent ecological studies in the Aleutian Islands have shown the concurrent existence of two alternate stable nearshore communities, one dominated by macroalgae, the other by epibenthic herbivores, which are respectively maintained by the presence or absence of dense sea otter populations. Thus, rather than cultural shifts in food preference, the changes in Aleut prey were probably the result of local overexploitation of sea otters by aboriginal Aleuts.

Alaska↗

Breakpoint analysis and assessment of selected stressor variables on benthic macroinvertebrate and fish communities in Indiana streams: Implications for developing nutrient criteria

Water chemistry, periphyton and seston chlorophyll a (CHL a ), and biological community data were collected from 321 sites from 2001 through 2005 to (1) determine statistically and ecologically significant relations among the stressor (total nitrogen, total phosphorus, periphyton and seston CHL a , and turbidity) variables and response (biological community) variables; and, (2) determine the breakpoint of biological community attributes and metrics in response to changes in stressor variables. Because of the typically weak relations among the stressor and response variables, methods were developed to reduce the effects of non-nutrient biological stressors that could mask the effect of nutrients. Stressor variable concentrations ranged from 0.30 to 11.0 milligrams per liter (mg/L) for total nitrogen, 0.025 to 1.33 mg/L for total phosphorus, 2.9 to 768 milligrams per square meter (mg/m 2 ) for periphyton CHL a , and 0.37 to 42 micrograms per liter (µg/L) for seston CHL a . Turbidity, another stressor variable, ranged from 0.8 to 65.4 Nephelometric turbidity units (NTUs). When the nutrient and CHL a data were compared to Dodds’ trophic classifications, 75.0 percent of the values for total nitrogen, 46.6 percent of the values for total phosphorus, 35.8 percent of the values for periphyton CHL a , and 3.5 percent of the values for seston CHL a , were eutrophic. The invertebrate communities were dominated by families considered highly nutrient tolerant, Chironimidae, (41.7 percent relative abundance), Hydropsychidae, (17.3 percent relative abundance), and Baetidae, (10.2 percent relative abundance). Fish communities were dominated by algivores and nutrient-tolerant species, specifically central stonerollers (13.3 percent relative abundance), creek chubs (9.9 percent relative abundance), and bluntnose minnows (9.3 percent relative abundance). Although not the dominant taxa, white sucker, spotted sucker, green sunfish, and bluegill species were correlated (p ‹0.05) with the stressor variables. The median breakpoints ranged from 2.4 to 3.3 mg/L for total nitrogen, from 0.042 to 0.129 mg/L for total phosphorus, from 54 to 68 mg/m 2 for periphyton CHL a , from 4.5 to 7.5 µg/L for seston CHL a , and from 14.1 to 16.1 NTU for turbidity. The breakpoints determined in this study, in addition to Dodds’ trophic classifications, were used as multiple lines of evidence to show changes in fish and invertebrate community and attributes based on annual exposure to nutrients.

Indiana↗

Effects of low pH on the coral reef cryptic invertebrate communities near CO2 vents in Papua New Guinea

Small cryptic invertebrates (the cryptofauna) are extremely abundant, ecologically important, and species rich on coral reefs. Ongoing ocean acidification is likely to have both direct effects on the biology of these organisms, as well as indirect effects through cascading impacts on their habitats and trophic relationships. Naturally acidified habitats have been important model systems for studying these complex interactions because entire communities that are adapted to these environmental conditions can be analyzed. However, few studies have examined the cryptofauna because they are difficult to census quantitatively in topographically complex habitats and are challenging to identify. We addressed these challenges by using Autonomous Reef Monitoring Structures (ARMS) for sampling reef-dwelling invertebrates >2 mm in size and by using DNA barcoding for taxonomic identifications. The study took place in Papua New Guinea at two reef localities, each with three sites at varying distances from carbon dioxide seeps, thereby sampling across a natural gradient in acidification. We observed sharp overall declines in both the abundance (34–56%) and diversity (42–45%) of organisms in ARMS under the lowest pH conditions sampled (7.64–7.75). However, the overall abundance of gastropods increased slightly in lower pH conditions, and crustacean and gastropod families exhibited varying patterns. There was also variability in response between the two localities, despite their close proximity, as one control pH site displayed unusually low diversity and abundances for all invertebrate groups. The data illustrate the complexity of responses of the reef fauna to pH conditions, and the role of additional factors that influence the diversity and abundance of cryptic reef invertebrates.

Milne Bay Province↗

Identifying nutrient reference sites in nutrient-enriched regions-Using algal, invertebrate, and fish-community measures to identify stressor-breakpoint thresholds in Indiana rivers and streams, 2005-9

Excess nutrients in aquatic ecosystems can lead to shifts in species composition, reduced dissolved oxygen concentrations, fish kills, and toxic algal blooms. In this study, nutrients, periphyton chlorophyll a (CHLa), and invertebrate- and fishcommunity data collected during 2005-9 were analyzed from 318 sites on Indiana rivers and streams. The objective of this study was to determine which invertebrate and fish-taxa attributes best reflect the conditions of streams in Indiana along a gradient of nutrient concentrations by (1) determining statistically and ecologically significant relations among the stressor (total nitrogen, total phosphorus, and periphyton CHLa) and response (invertebrate and fish community) variables; and (2) determining the levels at which invertebrate- and fish-community measures change in response to nutrients or periphyton CHL a . For water samples at the headwater sites, total nitrogen (TN) concentrations ranged from 0.343 to 21.6 milligrams per liter (mg/L) (median 2.12 mg/L), total phosphorus (TP) concentrations ranged from 0.050 to 1.44 mg/L (median 0.093 mg/L), and periphyton CHL a ranged from 0.947 to 629 mg/L (median 69.7 mg/L). At the wadable sites, TN concentrations ranged from 0.340 to 10.0 mg/L (median 2.31 mg/L), TP concentrations ranged from 0.050 to 1.24 mg/L (median 0.110 mg/L), and periphyton CHLa ranged from 0.383 to 719 mg/L (median 44.7 mg/L). Recursive partitioning identified statistically significant low and high breakpoint thresholds on invertebrate and fish measures, which demonstrated the ecological response in enriched conditions. The combined community (invertebrate and fish) mean low and high TN breakpoint thresholds were 1.03 and 2.61 mg/L, respectively. The mean low and high breakpoint thresholds for TP were 0.083 and 0.144 mg/L, respectively. The mean low and high breakpoint thresholds for periphyton CHL a were 20.9 and 98.6 milligrams per square meter (mg/m 2 ), respectively. Additive quantile regression analysis found similar thresholds (TN of 0.656 mg/L, mean TP of 0.118 mg/L, and periphyton CHLa of 27.2 mg/m 2 ) for some stressor variables as determined by the breakpoint analysis. The TN and TP concentrations in this study showed a nutrient gradient that spanned three orders of magnitude. Sites were divided into Low, Medium, and High nutrient groups based on the 10th and 75th percentiles. The invertebrate and fish communities were similar along the nutrient gradient, using an analysis of similarity, demonstrating there was not a species trophic gradient. Within all nutrient groups, invertebrate and fish communities were dominated by nutrient tolerant taxa (algivores, herbivores, and omnivores) that included invertebrates, such as Cheumatopsyche sp., Physella sp., and fish such as Stonerollers ( Campostoma spp.) and Bluntnose Minnow ( Pimephales notatus ). To determine if low nutrient concentrations at some sites were caused by algal uptake and not oligotrophic conditions, sites with low nutrient concentrations (less than 10th percentile for TN or TP) were examined based on the Low (less than or equal to the 10th percentile) and High (greater than the 75th percentile) periphyton CHL a concentrations. Within low nutrient sites, the invertebrate and fish communities were statistically different between Low and High periphyton CHL a categories. The majority of variance between the Low and High periphyton CHL a categories was caused by Cheumatopsyche sp. (caddisfly), Physella sp. (pulmonate snail), and Caenis latipennis (a mayfly) in the invertebrate community; and caused by Stonerollers, Western Blacknose Dace ( Rhinichthys atratulus meleagris ), and Creek Chub ( Semotilus atromaculatus ) in the fish community. The dominance of tolerant herbivore and omnivore taxa in the High periphyton CHL a group indicates that low nutrient concentrations are a result of nutrient uptake and increased algal growth. This study highlights the importance of assessing multiple lines of evidence when attempting to identify the trophic condition of a site.

Indiana↗

Quantifying and predicting fuels and the effects of reduction treatments along successional and invasion gradients in sagebrush habitats

Sagebrush shrubland ecosystems in the Great Basin are prime examples of how altered successional trajectories can create dynamic fuel conditions and, thus, increase uncertainty about fire risk and behavior. Although fire is a natural disturbance in sagebrush, post-fire environments are highly susceptible to conversion to an invasive grass-fire regime (often referred to as a “grass-fire cycle”). After fire, native shrub-steppe plants are often slow to regenerate, whereas nonnative annuals, especially cheatgrass (Bromus tectorum) and medusahead (Taeniatherum caput-medusae), can establish quickly and outcompete native species. Once fire-prone annuals become established, fire occurrences increase, further promoting dominance of nonnative species. The invasive grass-fire regime also alters nutrient and hydrologic cycles, pushing ecosystems beyond ecological thresholds toward steady-state, fire-prone, nonnative communities. These changes affect millions of hectares in the Great Basin and increase fire risk, decrease habitat quality and biodiversity, accelerate soil erosion, and degrade rangeland resources for livestock production. In many sagebrush landscapes, constantly changing plant communities and fuel conditions hinder attempts by land managers to predict and control fire behavior, restore native communities, and provide ecosystem services (e.g., forage production for livestock). We investigated successional and nonnative plant invasion states and associated fuel loads in degraded sagebrush habitat in a focal study area, the Morley Nelson Snake River Birds of Prey National Conservation Area (hereafter the NCA), in the Snake River Plain Ecoregion of southern Idaho. We expanded our inference by comparing our findings to similar data collected throughout seven major land resource areas (MLRAs) across the Great Basin (JFSP Project “Fire Rehabilitation Effectiveness: A Chronosequence Approach for the Great Basin” [09-S-02-1]). 4 We used a combination of field-sampling, experimental treatments, and remotely sensed data to address the following questions: (1) How do fuel loads change along gradients of succession and invasion in sagebrush ecological sites? (2) How do fuel reduction treatments influence fuels in invaded areas formerly dominated by sagebrush? (3) How do fuel loads vary across landscapes and which remote sensing techniques are effective for characterizing them?

Idaho↗

Hydrothermal vents and methane seeps: Rethinking the sphere of influence

Although initially viewed as oases within a barren deep ocean, hydrothermal vent and methane seep communities are now recognized to interact with surrounding ecosystems on the sea floor and in the water column, and to affect global geochemical cycles. The importance of understanding these interactions is growing as the potential rises for disturbance from oil and gas extraction, seabed mining and bottom trawling. Here we synthesize current knowledge of the nature, extent and time and space scales of vent and seep interactions with background systems. We document an expanded footprint beyond the site of local venting or seepage with respect to elemental cycling and energy flux, habitat use, trophic interactions, and connectivity. Heat and energy are released, global biogeochemical and elemental cycles are modified, and particulates are transported widely in plumes. Hard and biotic substrates produced at vents and seeps are used by “benthic background” fauna for attachment substrata, shelter, and access to food via grazing or through position in the current, while particulates and fluid fluxes modify planktonic microbial communities. Chemosynthetic production provides nutrition to a host of benthic and planktonic heterotrophic background species through multiple horizontal and vertical transfer pathways assisted by flow, gamete release, animal movements, and succession, but these pathways remain poorly known. Shared species, genera and families indicate that ecological and evolutionary connectivity exists among vents, seeps, organic falls and background communities in the deep sea; the genetic linkages with inactive vents and seeps and background assemblages however, are practically unstudied. The waning of venting or seepage activity generates major transitions in space and time that create links to surrounding ecosystems, often with identifiable ecotones or successional stages. The nature of all these interactions is dependent on water depth, as well as regional oceanography and biodiversity. Many ecosystem services are associated with the interactions and transitions between chemosynthetic and background ecosystems, for example carbon cycling and sequestration, fisheries production, and a host of non-market and cultural services. The quantification of the sphere of influence of vents and seeps could be beneficial to better management of deep-sea environments in the face of growing industrialization.

Frontiers in Marine Science↗

A Bayesian hierarchical modeling approach for species diversity in ecology

Species diversity is the foundation of many ecological disciplines. This metric is often approximated using species richness and evenness, even though actual richness likely exceeds observations due to imperfect sampling methods. Estimating the “true” species richness, which includes identifying the number of missing species, has intrigued ecologists for decades. We adopted a parametric model that appeared in Fisher et al. (1943), which models the numbers of individuals from different species as random samples from a negative binomial distribution, and developed a Bayesian computational approach to directly estimate the distribution model parameters. The model parameters represent species abundance and evenness, and can be used to derive species richness. We evaluated our parametric approach using (1) a simulation study and (2) three historical data sets. Furthermore, we illustrated the hierarchical modeling approach to combine data from multiple parallel studies using a biannual fishery survey data set. Our parametric model formulation is computationally efficient, and the hierarchical structure facilitates embedding diversity estimation into broader application, such as assessing spatial and temporal trends in species diversity associated with environmental stressors. Additionally, because the two parameters of the negative binomial distribution model represent species abundance and evenness of a community, this parametric approach facilitates a deeper understanding of the ecological systems under study. The negative binomial distribution model works with a wide range of species frequency distribution types. As a result, our emphasis on a parametric model can help us characterize the structure of an ecosystem and provide a greater depth of ecologically meaningful information.

Ecological Informatics↗

Avifaunal responses to fire in southwestern montane forests along a burn severity gradient

The effects of burn severity on avian communities are poorly understood, yet this information is crucial to fire management programs. To quantify avian response patterns along a burn severity gradient, we sampled 49 random plots (2001-2002) at the 17 351-ha Cerro Grande Fire (2000) in New Mexico, USA. Additionally, pre-fire avian surveys (1986-1988, 1990) created a unique opportunity to quantify avifaunal changes in 13 pre-fire transects (resampled in 2002) and to compare two designs for analyzing the effects of unplanned disturbances: after-only analysis and before-after comparisons. Distance analysis was used to calculate densities. We analyzed after-only densities for 21 species using gradient analysis, which detected a broad range of responses to increasing burn severity: (I) large significant declines, (II) weak, but significant declines, (III) no significant density changes, (IV) peak densities in low- or moderate-severity patches, (V) weak, but significant increases, and (VI) large significant increases. Overall, 71% of the species included in the after-only gradient analysis exhibited either positive or neutral density responses to fire effects across all or portions of the severity gradient (responses III-VI). We used pre/post pairs analysis to quantify density changes for 15 species using before-after comparisons; spatiotemporal variation in densities was large and confounded fire effects for most species. Only four species demonstrated significant effects of burn severity, and their densities were all higher in burned compared to unburned forests. Pre- and post-fire community similarity was high except in high-severity areas. Species richness was similar pre- and post-fire across all burn severities. Thus, ecosystem restoration programs based on the assumption that recent severe fires in Southwestern ponderosa pine forests have overriding negative ecological effects are not supported by our study of post-fire avian communities. This study illustrates the importance of quantifying burn severity and controlling confounding sources of spatiotemporal variation in studies of fire effects. After-only gradient analysis can be an efficient tool for quantifying fire effects. This analysis can also augment historical data sets that have small samples sizes coupled with high non-process variation, which limits the power of before-after comparisons. ?? 2007 by the Ecological Society of America.

Ecological Applications↗

Flow variations and macroinvertebrate community responses in a small groundwater-dominated stream in south east England

Changes in the macroinvertebrate community in response to flow variations in the Little Stour River, Kent, UK, were examined over a 6 year period (1992-1997). This period included the final year of the 1988-1992 drought, followed by some of the wettest conditions recorded this century and a second period of drought between 1996 and 1997. Each year, samples were collected from 15 sites during late-summer base-flow conditions. Correspondence analysis identified clear differences between samples from upstream and downstream sites, and between drought and non-drought years. Step-wise multiple regression was used to identify hydrological indicators of community variation. Several different indices were used to describe the macroinvertebrate community, including macroinvertebrate community abundance, number of families and species, and individual species. Site characteristics were fundamental in accounting for variation in the unstandardized macroinvertebrate community. However, when differences between sites were controlled, hydrological conditions were found to play a dominant role in explaining ecological variation. Indices of high discharge (or their absence), 4-7 months prior to sampling (i.e. winter-spring), were found to be the most important variables for describing the late-summer community The results are discussed in relation to the role of flow variability in shaping instream communities and management implications. Copyright ?? 2000 John Wiley & Sons, Ltd.Changes in the macroinvertebrate community in response to flow variations in the Little Stour River, Kent, UK, were examined over a 6 year period (1992-1997). This period included the final year of the 1988-1992 drought, followed by some of the wettest conditions recorded this century and a second period of drought between 1996 and 1997. Each year, samples were collected from 15 sites during late-summer base-flow conditions. Correspondence analysis identified clear differences between samples from upstream and downstream sites, and between drought and non-drought years. Step-wise multiple regression was used to identify hydrological indicators of community variation. Several different indices were used to describe the macroinvertebrate community, including macroinvertebrate community abundance, number of families and species, and individual species. Site characteristics were fundamental in accounting for variation in the unstandardized macroinvertebrate community. However, when differences between sites were controlled, hydrological conditions were found to play a dominant role in explaining ecological variation. Indices of high discharge (or their absence), 4-7 months prior to sampling (i.e. winter-spring), were found to be the most important variables for describing the late-summer community. The results are discussed in relation to the role of flow variability in shaping instream communities and management implications.

Hydrological Processes↗

The role of habitat heterogeneity and canyon processes in structuring sediment macrofaunal communities associated with hard substrate habitats in Norfolk Canyon, USA

Topographic and hydrodynamic complexity in submarine canyons promotes steep gradients in food availability and geophysical parameters which affect ecological assemblages and beta diversity. While habitat heterogeneity in submarine canyons is known to support diverse and abundant megafaunal communities, due to difficulty in sampling little is known about infaunal communities adjacent to hard substrate habitats, their contribution to canyon assemblages, and overall deep-sea diversity. Sediments were collected in three distinct habitat types: within Norfolk Canyon (western Atlantic) adjacent to hard substrate habitats including canyon walls and large boulders, along the main axis of the canyon, and on the adjacent continental slope to quantify macrofaunal (>300 μm) density, diversity and community composition, and sediment geochemical parameters including grain size, organic content, and stable isotope composition. While macrofaunal densities were similar among habitats sampled at comparable depths, diversity was higher near the hard substrate environments. Discrete communities were present in each habitat type, while annelid functional composition was similar between hard substrate adjacent and canyon axis habitats. Although diversity and abundance of hard substrate adjacent sediment communities did not change with depth, communities were driven by sediments with low organic matter content, whereas canyon and slope community assemblages were best explained by depth and higher organic content. Beta diversity among hard substrate adjacent sediments and canyon axis communities was high, with 27% of canyon taxa and 10% of regional taxa only occurring in hard substrate habitats. Given the thousands of submarine canyons worldwide, our results highlight the overall importance of substrate habitat heterogeneity within canyons in supporting deep-sea benthic diversity and suggest that both within-canyon and regional diversity are underestimated.

Norfolk Canyon↗

Multilevel Hierarchical Modeling of Benthic Macroinvertebrate Responses to Urbanization in Nine Metropolitan Regions across the Conterminous United States

Multilevel hierarchical modeling methodology has been developed for use in ecological data analysis. The effect of urbanization on stream macroinvertebrate communities was measured across a gradient of basins in each of nine metropolitan regions across the conterminous United States. The hierarchical nature of this dataset was harnessed in a multi-tiered model structure, predicting both invertebrate response at the basin scale and differences in invertebrate response at the region scale. Ordination site scores, total taxa richness, Ephemeroptera, Plecoptera, Trichoptera (EPT) taxa richness, and richness-weighted mean tolerance of organisms at a site were used to describe invertebrate responses. Percentage of urban land cover was used as a basin-level predictor variable. Regional mean precipitation, air temperature, and antecedent agriculture were used as region-level predictor variables. Multilevel hierarchical models were fit to both levels of data simultaneously, borrowing statistical strength from the complete dataset to reduce uncertainty in regional coefficient estimates. Additionally, whereas non-hierarchical regressions were only able to show differing relations between invertebrate responses and urban intensity separately for each region, the multilevel hierarchical regressions were able to explain and quantify those differences within a single model. In this way, this modeling approach directly establishes the importance of antecedent agricultural conditions in masking the response of invertebrates to urbanization in metropolitan regions such as Milwaukee-Green Bay, Wisconsin; Denver, Colorado; and Dallas-Fort Worth, Texas. Also, these models show that regions with high precipitation, such as Atlanta, Georgia; Birmingham, Alabama; and Portland, Oregon, start out with better regional background conditions of invertebrates prior to urbanization but experience faster negative rates of change with urbanization. Ultimately, this urbanization-invertebrate response example is used to detail the multilevel hierarchical construction methodology, showing how the result is a set of models that are both statistically more rigorous and ecologically more interpretable than simple linear regression models.

Scientific Investigations Report↗

Morphological identification and COI barcodes of adult flies help determine species identities of chironomid larvae (Diptera, Chironomidae)

Establishing reliable methods for the identification of benthic chironomid communities is important due to their significant contribution to biomass, ecology and the aquatic food web. Immature larval specimens are more difficult to identify to species level by traditional morphological methods than their fully developed adult counterparts, and few keys are available to identify the larval species. In order to develop molecular criteria to identify species of chironomid larvae, larval and adult chironomids from Western Lake Erie were subjected to both molecular and morphological taxonomic analysis. Mitochondrial cytochrome c oxidase I (COI) barcode sequences of 33 adults that were identified to species level by morphological methods were grouped with COI sequences of 189 larvae in a neighbor-joining taxon-ID tree. Most of these larvae could be identified only to genus level by morphological taxonomy (only 22 of the 189 sequenced larvae could be identified to species level). The taxon-ID tree of larval sequences had 45 operational taxonomic units (OTUs, defined as clusters with >97% identity or individual sequences differing from nearest neighbors by >3%; supported by analysis of all larval pairwise differences), of which seven could be identified to species or ‘species group’ level by larval morphology. Reference sequences from the GenBank and BOLD databases assigned six larval OTUs with presumptive species level identifications and confirmed one previously assigned species level identification. Sequences from morphologically identified adults in the present study grouped with and further classified the identity of 13 larval OTUs. The use of morphological identification and subsequent DNA barcoding of adult chironomids proved to be beneficial in revealing possible species level identifications of larval specimens. Sequence data from this study also contribute to currently inadequate public databases relevant to the Great Lakes region, while the neighbor-joining analysis reported here describes the application and confirmation of a useful tool that can accelerate identification and bioassesment of chironomid communities.

Bulletin of Entomological Research↗

New ecological knowledge and practices for society and sustainability

As attributes of the Earth's ecosystems shift in the face of human impact and sustainability of ecosystem services becomes less certain, one important tool at the disposal of the scientific community and other groups is a blueprint for understanding, evaluating, and communicating the value of ecological services. The blueprint presented here is based on (1) an examination of the ecological and societal trade-offs accompanying any given action, (2) revised methods of communication, and (3) coordination of actions at many different scales. The Mexican National Commission for the Knowledge and Use of Biodiversity (CONABIO) is a good example of a demand-driven “bridging institution” between academia, government, and civil society, and it works to collect and convert scientific information into information for policy, management, and conservation. Intergovernmental organizations like the United Nations Educational, Scientific and Cultural Organization (UNESCO) are well placed to facilitate such coordination at the international level, through their work with member states. Through collaboration with the constituencies of such organizations as the Ecological Society of America, the blueprint described below has the potential to become an important tool for assessing and managing threats to ecosystem services that are essential to life.

Frontiers in Ecology and the Environment↗

Effects of feral horse herds on plant communities across a precipitation gradient

Feral horses are widespread in the western United States, with the majority of feral horse herds found in the Great Basin. There is a federal mandate to manage these herds in order to maintain “ecological balance”; however, understanding of the specific effects of feral horse grazing on rangeland plant communities in this region is incomplete. To address this research gap, we utilized long-term grazing exclosures and fenceline contrasts to evaluate the impacts of feral horses on several plant community variables (diversity, richness, dominance, and biomass) and species composition. Because the effects of grazing can vary with site precipitation and productivity, we selected 5 sites from 4 different rangeland types (Great Basin Desert, Colorado Plateau, Rocky Mountain grassland, and mixed grass prairie) that spanned a mean annual precipitation gradient of 229 to 413 mm. Our results did not reveal a significant effect of feral horse grazing on plant community composition, species richness, diversity, evenness, or dominance. In contrast, total aboveground herbaceous biomass and grass biomass were significantly reduced with feral horse grazing, but these effects did not vary with mean annual precipitation. Our results suggest that, at least at the sites we studied, feral horses have affected the plant community by reducing herbaceous biomass but have not caused plant community shifts. Additional multisite studies, preferably with standardized exclosures and larger sample sizes, would increase our understanding of feral horse grazing effects and inform management of feral horse herds in the western United States.

Colorado, Montana, Nevada, North Dakota, Utah↗

Ecology of the macrophyte Podostemum ceratophyllum Michx. (Hornleaf riverweed), a widespread foundation species of eastern North American rivers

Podostemum ceratophyllum , commonly called Hornleaf Riverweed, occurs in mid-order montane and piedmont rivers of eastern North America, where the plant grows submerged and attached to rocks and stable substrates in swift, aerated water. Multiple studies, mostly conducted in the southern portions of the plant’s range, have shown that Podostemum can variously influence benthic communities in flowing waters. However, a synthetic review of the biology and ecology of the plant is needed to inform conservation, particularly because P. ceratophyllum is reported to be in decline in much of its range, for mostly unknown reasons. We have thus summarized the literature showing that Podostemum provides substantial habitat for invertebrates and fish, may be consumed by invertebrates, turtles, and other vertebrates, removes and sequesters dissolved elements (i.e., nitrogen, phosphorus, calcium, zinc, etc.) from the water column, and contributes organic matter to the detrital pool. Podostemum may be tolerant to some forms of pollution but appears vulnerable to sedimentation, epiphytic over-growth, and hydrologic changes that result in desiccation, and possibly increased herbivory pressure. Much remains unknown about Podostemum , including aspects of morphological variation, seed dispersal, and tolerance to changes in temperature and water chemistry. Nonetheless, Podostemum may be considered a foundation species, whose loss from eastern North American rivers is likely to affect higher trophic levels and ecosystem processes.

Aquatic Botany↗