Alexander D. Huryn

Length-Mass Relationships for Freshwater Macroinvertebrates in North America with Particular Reference to the Southeastern United States

Estimation of invertebrate biomass is a critical step in addressing many ecological questions in aquatic environments. Length-dry mass regressions are the most widely used approach for estimating benthic invertebrate biomass because they are faster and more precise than other methods. A compilation and analysis of length-mass regressions using the power model, M (mass) = a L (length)b, are presented from 30 y of data collected by the authors, primarily from the southeastern USA, along with published regressions from the rest of North America. A total of 442 new and published regressions are presented, mostly for genus or species, based on total body length or other linear measurements. The regressions include 64 families of aquatic insects and 12 families of other invertebrate groups (mostly molluscs and crustaceans). Regressions were obtained for 134 insect genera (155 species) and 153 total invertebrate genera (184 species). Regressions are provided for both body length and head width for some taxa. In some cases, regressions are provided from multiple localities for single taxa. When using body length in the equations, there were no significant differences in the mean value of the exponent b among 8 insect orders or Amphipoda. The mean value of b for insects was 2.79, ranging from only 2.69 to 2.91 among orders. The mean value of b for Decapoda (3.63), however, was significantly higher than all insects orders and amphipods. Mean values of a were not significantly different among the 8 insect orders and Amphipoda, reflecting considerable variability within orders. Reasons for potential differences in b among taxa are explained with hypothetical examples showing how b responds to changes in linear dimensions and specific gravity. When using head width as the linear dimension in the power model, the mean value of b was higher (3.11) than for body length and more variable among orders (2.8-3.3). Values of b for Ephemeroptera (3.3) were significantly higher than those for Odonata, Megaloptera, and Diptera. For those equations in which ash-free dry mass was used, % ash varied considerably among functional feeding groups (3.3-12.4%). Percent ash varied from 4.0% to 8.5% among major insect orders, but was 18.9% for snails (without shells). Family-level regressions also are presented so that they can be used when generic equations are unavailable or when organisms are only identified to the family level. It is our intention that these regressions be used by others in estimating mass from linear dimensions, but potential errors must be recognized.

The effects of amphibian population declines on the structure and function of Neotropical stream ecosystems

Amphibians can be important consumers in both aquatic and terrestrial habitats and may represent an important energetic link between the two, particularly in the tropics, where amphibian species richness and abundance are high. In the past 20 years, amphibian populations have declined dramatically around the world; numbers have decreased catastrophically in protected upland sites throughout the neotropics, usually resulting in the disappearance of over 75% of amphibians at a given site, particularly those species that breed in streams. Most studies of amphibian declines have focused on identifying causes and documenting changes in adult abundance, rather than on their ecological consequences. Here, we review evidence for the potential ecological effects of catastrophic amphibian declines, focusing on neotropical highland streams, where impacts will likely be greatest. Evidence to date suggests that amphibian declines will have large-scale and lasting ecosystem-level effects, including changes in algal com...

Impervious Surface Area as a Predictor of the Effects of Urbanization on Stream Insect Communities in Maine, U.S.A.

The influence of urbanization on stream insect communities was determined by comparing physical, chemical, and biological characteristics of streams draining 20 catchments with varyinglevels of urban land-cover in Maine (U.S.A). Percent total impervious surface area (PTIA), which was used to quantify urbanland-use, ranged from ∼1–31% among the study catchments.Taxonomic richness of stream insect communities showed an abruptdecline as PTIA increased above 6%. Streams draining catchmentswith PTIA < 6% had the highest levels of both total insect and EPT (Ephemeroptera + Plecoptera + Trichoptera) taxonomic richness. These streams contained insect communities with a totalrichness averaging 33 taxa in fall and 31 taxa in spring; EPT richness ranged from an average of 15 taxa in fall and 13 taxa inspring. In contrast, none of the streams draining catchments with6–27% PTIA had a total richness > 18 taxa or an EPT richness> 6 taxa. Insect communities in streams with PTIA > 6% were characterized by the absence of pollution-intolerant taxa. The distribution of more pollution-tolerant taxa (e.g.Acerpenna (Ephemeroptera); Paracapnia, Allocapnia (Plecoptera); Optioservus, Stenelmis (Coleoptera); Hydropsyche, Cheumatopsyche (Trichoptera)), however, showed little relation to PTIA. In contrast to the apparent threshold relationship between PTIA and insect taxonomic richness, both habitat qualityand water quality tended to decline as linear functions of PTIA.Our results indicate that, in Maine, an abrupt change in stream insect community structure occurs at a PTIA above a threshold ofapproximately 6% of total catchment area. The measurement of PTIA may provide a valuable tool for predicting thresholds for adverse effects of urbanization on the health of headwater streams in Maine.

URBANIZATION AFFECTS STREAM ECOSYSTEM FUNCTION BY ALTERING HYDROLOGY, CHEMISTRY, AND BIOTIC RICHNESS

Catchment urbanization can alter physical, chemical, and biological attributes of stream ecosystems. In particular, changes in land use may affect the dynamics of organic matter decomposition, a measure of ecosystem function. We examined leaf-litter decomposition in 18 tributaries of the St. Johns River, Florida, USA. Land use in all 18 catchments ranged from 0% to 93% urban which translated to 0% to 66% total impervious area (TIA). Using a litter-bag technique, we measured mass loss, fungal biomass, and macroinvertebrate biomass for two leaf species (red maple [Acer rubrum] and sweetgum [Liquidambar styraciflua]). Rates of litter mass loss, which ranged from 0.01 to 0.05 per day for red maple and 0.006 to 0.018 per day for sweetgum, increased with impervious catchment area to levels of approximately 30-40% TIA and then decreased as impervious catchment area exceeded 40% TIA. Fungal biomass was also highest in streams draining catchments with intermediate levels of TIA. Macroinvertebrate biomass ranged from 17 to 354 mg/bag for red maple and from 15 to 399 mg/bag for sweetgum. Snail biomass and snail and total invertebrate richness were strongly related to breakdown rates among streams regardless of leaf species. Land-use and physical, chemical, and biological variables were highly intercorrelated. Principal-components analysis was therefore used to reduce the variables into several orthogonal axes. Using stepwise regression, we found that flow regime, snail biomass, snail and total invertebrate richness, and metal and nutrient content (which varied in a nonlinear manner with impervious surface area) were likely factors affecting litter breakdown rates in these streams.

Effect of riparian land use on contributions of terrestrial invertebrates to streams

Since terrestrial invertebrates are often consumed by stream fishes, land-use practices that influence the input of terrestrial invertebrates to streams are predicted to have consequences for fish production. We studied the effect of riparian land-use regime on terrestrial invertebrate inputs by estimating the biomass, abundance and taxonomic richness of terrestrial invertebrate drift from 15 streams draining catchments with three different riparian land-use regimes and vegetation types: intensive grazing — exotic pasture grasses (4 streams), extensive grazing — native tussock grasses (6 streams), reserve — native forest (5 streams). Terrestrial invertebrate drift was sampled from replicated stream reaches enclosed by two 1 mm mesh drift nets that spanned the entire channel. The mean biomass of terrestrial invertebrates that entered tussock grassland (12 mg ash-free dry mass m−2 d−1) and forest streams (6 mg AFDM m−2 d−1) was not significantly different (p > 0.05). However, biomass estimated for tussock grassland and forest streams was significantly higher than biomass that entered pasture streams (1 mg AFDM m−2 d−1). Mean abundance and richness of drifting terrestrial invertebrates was not significantly different among land-use types. Winged insects contributed more biomass than wingless invertebrates to both pasture and tussock grassland streams. Winged and wingless invertebrates contributed equally to biomass entering forest streams. Land use was a useful variable explaining landscape-level patterns of terrestrial invertebrate input for New Zealand streams. Evidence from this study suggests that riparian land-use regime will have important influences on the availability of terrestrial invertebrates to stream fishes.

Benthic invertebrate production—facilitating answers to ecological riddles in freshwater ecosystems

Abstract Invertebrate secondary production, or the formation of invertebrate biomass through time, has been estimated in many freshwater benthic habitats. It has been a major research theme for the North American Benthological Society (NABS), and many of its members have made significant contributions to the subject, both before and during the existence of J-NABS. Although some benthic production work occurred before 1960, the major methods were developed primarily during the 1960s and 1970s. Most of these methods also were applied in terrestrial and marine environments. The main focus of our paper is how secondary production has been used as an essential variable in facilitating answers to a wide variety of ecological questions. Benthic freshwater production studies before the inception of J-NABS were primarily related to life history, interpopulation comparisons, niche overlap/competition, predator–prey relationships, differences in production/biomass (P/B), energy flow, the trophic basis of production, habitat-specific microdistributions, effects of pollution and dams, and quantification of aquatic–terrestrial linkages. Since that time, new applications have been related to habitat-specific macrodistributions, quantitative food webs, experimental and tracer-based studies of trophic resources, chemical flows/stoichiometric relationships, diversity/function relationships, influence of nonnative species and landuse changes, implications of metabolic theory, and the importance of meiofauna vs macrofauna. J-NABS has been a major outlet for many of these applications, has probably included a higher fraction of papers incorporating secondary production analysis than any other journal, and probably will continue to be a leader in this area.

Interactions between temperature and nutrients across levels of ecological organization

Temperature and nutrient availability play key roles in controlling the pathways and rates at which energy and materials move through ecosystems. These factors have also changed dramatically on Earth over the past century as human activities have intensified. Although significant effort has been devoted to understanding the role of temperature and nutrients in isolation, less is known about how these two factors interact to influence ecological processes. Recent advances in ecological stoichiometry and metabolic ecology provide a useful framework for making progress in this area, but conceptual synthesis and review are needed to help catalyze additional research. Here, we examine known and potential interactions between temperature and nutrients from a variety of physiological, community, and ecosystem perspectives. We first review patterns at the level of the individual, focusing on four traits--growth, respiration, body size, and elemental content--that should theoretically govern how temperature and nutrients interact to influence higher levels of biological organization. We next explore the interactive effects of temperature and nutrients on populations, communities, and food webs by synthesizing information related to community size spectra, biomass distributions, and elemental composition. We use metabolic theory to make predictions about how population-level secondary production should respond to interactions between temperature and resource supply, setting up qualitative predictions about the flows of energy and materials through metazoan food webs. Last, we examine how temperature-nutrient interactions influence processes at the whole-ecosystem level, focusing on apparent vs. intrinsic activation energies of ecosystem processes, how to represent temperature-nutrient interactions in ecosystem models, and patterns with respect to nutrient uptake and organic matter decomposition. We conclude that a better understanding of interactions between temperature and nutrients will be critical for developing realistic predictions about ecological responses to multiple, simultaneous drivers of global change, including climate warming and elevated nutrient supply.

Direct and Indirect Effects of Geology on the Distribution, Biomass, and Production of the Freshwater Snail Elimia

We studied the relationship between geology and lotic secondary production by analyzing geographical patterns of the distribution, biomass, and production of the freshwater snail Elimia (Pleuroceridae) in Alabama. Nine streams were selected for study, three each in three physiographically distinct regions of uniform climate but contrasting lithology. Our objectives were to assess: 1) the production and biomass of Elimia among streams with contrasting alkalinities due to differences in regional geology--sandstone (5.7 mg/L as CaCo3) < phyllite (19.4) < carbonate (103.4); p < 0.05--and 2) geologically mediated variation in thermal regime as an alternative factor contributing to the widely observed correlation between alkalinity and productivity. Although conspicuous in phyllite and carbonate streams, Elimia was absent from sandstone streams. Biomass (B) and annual production (P) of Elimia were significantly lower (p ≤ 0.03) in streams draining phyllite (B = 1102 mg/m2, P = 1565 mg/m2) compared with carbonate catchments (B = 2990 mg/m2, P = 2501 mg/m2). However, whereas the correlation between biomass and alkalinity was consistent throughout the year, the correlation between production and alkalinity was not. Production did not differ significantly during the summer months (April-October, p = 0.41), and differences in annual production between regions were attributable to near cessation of production in phyllite streams during the winter months (October-April, p = 0.04). Although mean annual stream temperature was not significantly different between regions (p = 0.51), a simulation showed that the low winter temperatures of phyllite streams (⪡10°C) should cause the cessation of production and high net losses of snail biomass and result in insufficient post-winter biomass for rapid compounding of production during spring and summer. Consequently, biomass should be regulated at relatively lower levels in phyllite than in carbonate streams where production occurs year-round because of a more moderate thermal regime (e.g., winter minima ∼10°C).

Effect of a whole-catchment N addition on stream detritus processing

The Bear Brook Watershed in Maine (BBWM) is a paired catchment study investigating ecosystem effects of N and S deposition. Because of the decade long (NH4)2SO4 addition, the treatment catchment has higher stream NO3− and enriched foliar N concentrations compared to the reference catchment. We investigated how both stream N and foliar N affect stream detritus processing. Differences in litter processing were assessed by measuring mass loss, tissue softness, and shredder biomass. To examine both stream effects and leaf source, N-enriched and reference litter bags were prepared for 3 leaf species and placed in each stream. Red maple leaves were examined in 1997, 1998, and 1999. Sugar maple and American beech were examined in 1999. In all years, the only stream effects were increased mass loss for sugar maple and higher shredder biomass for red maple in 1998 in the treatment stream. Several leaf source effects were observed. N-enriched leaves of sugar maple, American beech, and red maple in 1998 had significantly higher microbial activity, as indicated by softer tissue, and had higher total mass loss. Further, shredder biomass tended to be highest in N-enriched litter bags. Although significant effects were detected, our results suggest that elevated dissolved N concentrations as a result of N deposition play a minimal role in regulating stream detritus processing at BBWM. Increased foliar N, however, did influence rates of stream detritus processing by increasing microbial activity, and possibly increasing shredder biomass. Our study shows how N deposited on entire catchments can affect litter processing in stream ecosystems.

CONTRASTING BEHAVIORAL RESPONSES BY DETRITIVOROUS AND PREDATORY MAYFLIES TO CHEMICALS RELEASED BY INJURED CONSPECIFICS AND THEIR PREDATORS

Larvae of the mayfly Siphlonisca are predators of the detritivorous mayfly Siphlonurus in floodplain wetlands in Maine (USA). Both mayflies are natural prey of brook trout (Salvelinus fontinalis). We exposed larvae of Siphlonurus and Siphlonisca to chemicals from injured conspecifics and their predators. Significant decreases in movement activity by Siphlonurus were elicited by chemicals released from Siphlonisca, chemicals released from brook trout fed conspecifics, and by chemicals released from injured conspecifics. A significant decrease in movement activity by Siphlonisca was elicited by chemicals released from brook trout fed either conspecifics or Siphlonurus. Movement activity by either Siphlonurus or Siphlonisca was not significantly affected by chemicals released from trout feeding on brine shrimp (Artemia). Both Siphlonurus and Siphlonisca were able to detect chemicals that provided information about past feeding behaviour by brook trout. However, their response to the chemicals used in this study was context-specific. A reduction in movement activity, a behavior that presumably reduces the probability of being consumed by visual predators, occurred only when mayflies were exposed to chemicals released by brook trout feeding on conspecific (Siphlonurus) or confamilial (Siphlonisca) prey.