LIMITING NUTRIENT DETERMINATION IN LOTIC ECOSYSTEMS USING A QUANTITATIVE NUTRIENT ENRICHMENT PERIPHYTOMETER1

ABSTRACT: The decline of water quality in United States’ lotic ecosystems (streams and rivers) has been linked to nonpoint source nutrient loading (U.S. EPA, 1990). Determining limiting nutrients in streams is difficult due to the variable nature of lotic ecosystems. We developed a quantitative passive diffusion periphyton nutrient enrichment system, called the Matlock Periphytometer, to measure the response of attached algae (periphyton) to nutrient enrichment. The system is simple to build and provides quantitative nutrient enrichment of a surface for periphytic growth. The periphyton grow on a glass fiber filter, which allows complete recovery of periphyton for chlorophyll a analysis. A 14-kilodalton dialysis membrane was used as a biofilter to prevent bacterial and algal contamination of the nutrient solution. We determined the rates of diffusion of nitrogen and phosphorus ions across the Matlock Periphytometer's dialysis membrane and glass fiber filter over a 21-day period (42 and 22 μg/cm²/hr, respectively). We used the Matlock Periphytometer to determine the limiting nutrient in a woodland stream. Six replicates each of a control, nitrogen, and phosphorus treatment were placed in the stream for 14 days. The results indicated that phosphorus was the limiting nutrient in the stream for the period and location sampled.     

Recovery of lotic communities and ecosystems from disturbance—A narrative review of case studies

We present a narrative account of case studies of the recovery of flowing water systems from disturbance, focusing on the investigators' conclusions about recovery time and the factors contributing to recovery. We restrict our attention to case studies in which the recovery of some biological property of the system has been examined, excluding those that deal only with physical or chemical properties. Although natural processes and rates of recovery are emphasized, studies of reclamation or restoration of damaged ecosystems are included where they contribute to an understanding of recovery processes. For the majority of studies examined, the systems recovered quite rapidly. The most commonly cited reasons for short recovery times were: (1) life history characteristics that allowed rapid recolonization and repopulation of the affected areas, (2) the availability and accessibility of unaffected up-stream and downstream areas and internal refugia to serve as sources of organisms for repopulation, (3) the high flushing rates of lotic systems that allowed them to quickly dilute or replace polluted waters, and (4) the fact that lotic systems are naturally subjected to a variety of disturbances and the biota have evolved life history characteristics that favor flexibility or adaptability. In general, longer recovery times were observed in disturbances, such as channelization, that resulted in alterations to physical conditions. This review also indicates that much of our knowledge of recovery in lotic ecosystems is fragmented and uncoordinated. In addition to establishing the bounds of recovery time, our review identifies some research gaps that need to be filled.     

Biogeochemical cycling constraints on stream ecosystem recovery

In systems where production is limited by the availability of a nutrient, nutrient input to and recycling within the system is related to the resilience, or speed of recovery, of a system to its steady state following a disturbance. In particular, it is shown that the return timeT _s of the system to steady state, or the inverse of the resilience, is approximately equal to the mean turnover time of the limiting nutrient in the system. From this relationship, it is possible to understand and predict how various properties of food webs and their environments affect resilience. These properties include nutrient input rate, loss rate, size of the detritus compartment, and trophic structure. The effects of these properties on resilience are described by using simple mathematical models. To test model predictions, experimental studies of the response of periphyton-dominated stream ecosystems to disturbance are being conducted on a set of laboratory streams in which nutrient inputs and grazing intensity are regulated at different levels. In streams without snail grazers (low-grazed streams), 90% recirculation of stream water to reduce nutrient inputs resulted in longer turnover times (T _r ) of phosphorus within the stream compared with once-through flow. However, in streams with snail grazers (high-grazed streams), there were no differences in phosphorus turnover time between once-through and partially recirculated treatments. Results on the rate of recovery of periphyton from a flood/scour disturbance to each stream partially support the model prediction of a positive relationship between ecosystem return time (T _s ) and nutrient turnover time (T _r ) within the streams.     

Recovery of lotic macroinvertebrate communities from disturbance

Ecosystem disturbances produce changes in macrobenthic community structure (abundances, biomass, and production) that persist for a few weeks to many decades. Examples of disturbances with extremely long-term effects on benthic communities include contamination by persistent toxic agents, physical changes in habitats, and altered energy inputs. Stream size, retention, and local geomorphology may ameliorate the influence of disturbances on invertebrates. Disturbances can alter food webs and may select for favorable genotypes (e.g., insecticidal resistance). Introductions of pesticides into lotic ecosystems, which do not result in major physical changes within habitats, illustrate several factors that influence invertebrate recovery time from disturbance. These include: (1) magnitude of original contamination, toxicity, and extent of continued use; (2) spatial scale of the disturbance; (3) persistence of the pesticide; (4) timing of the contamination in relation to the life history stages of the organisms; (5) vagility of populations influenced by pesticides; and (6) position within the drainage network. The ability of macroinvertebrates to recolonize denuded stream habitats may vary greatly depending on regional life histories, dispersal abilities, and position within the stream network (e.g., headwaters vs larger rivers). Although downstream drift is the most frequently cited mechanism of invertebrate recolonization following disturbance in middle- and larger-order streams, evidence is presented that shows aerial recolonization to be potentially important in headwater streams. There is an apparent stochastic element operating for aerial recolonization, depending on the timing of disturbance and flight periods of various taxa. Available evidence indicates that recolonization of invertebrate taxa without an aerial adult stage requires longer periods of time than for those that possess winged, terrestrial adult stages (i.e., most insects). Innovative, manipulative experiments are needed in order to address recolonization mechanisms of animals inhabiting streams that differ in size, latitude, disturbance frequency and magnitude, as well as the potential influence of early colonists on successional sequences of species following disturbance.     

Do Nutrients Limit Algal Periphyton in Small Blackwater Coastal Plain Streams?1

Abstract: We examine the potential for nutrient limitation of algal periphyton biomass in blackwater streams draining the Georgia coastal plain. Previous studies have investigated nutrient limitation of planktonic algae in large blackwater rivers, but virtually no scientific information exists regarding how algal periphyton respond to nutrients under different light conditions in smaller, low‐flow streams. We used a modification of the Matlock periphytometer (nutrient‐diffusing substrata) to determine if algal growth was nutrient limited and/or light limited at nine sites spanning a range of human impacts from relatively undisturbed forested basins to highly disturbed agricultural sites. We employed four treatments in both shaded and sunny conditions at each site: (1) control, (2) N (NO₃‐N), (3) P (PO₄‐P), and (4) N + P (NO₃‐N + PO₄‐P). Chlorophyll a response was measured on 10 replicate substrates per treatment, after 15 days of in situ exposure. Chlorophyll a values did not approach what have been defined as nuisance levels (i.e., 100‐200 mg/m²), even in response to nutrient enrichment in sunny conditions. For Georgia coastal plain streams, algal periphyton growth appears to be primarily light limited and can be secondarily nutrient limited (most commonly by P or N + P combined) in light gaps and/or open areas receiving sunlight.     

Application of ecological theory to determining recovery potential of disturbed lotic ecosystems: Research needs and priorities

This article summarizes the views of aquatic scientists who gathered to assess the ability of stream ecosystem theory to predict recovery from disturbance. Two views of disturbance were evident: a discrete removal of organisms vs an unusual deviation from normal. These were perceived as applying to different scales and/or objectives. Long-term information is required from both points of view to define recovery. Recovery also may be defined in different ways, but it is clear that recovery has both spatial and temporal components, and includes both physical and biological processes. Consensus was very strong that a major role (and challenge) for theory lies in the understanding of spatial aspects, temporal scales, coupling of physics and biology, and the interaction of these features in recovery processes. Some progress is evident in the articles of this volume, but among the topics identified as critical for further theoretical contributions were: homogeneous vs heterogeneous distribution of disturbance, local extent of disturbance relative to a regional context, critical vs noncritical patches (size and location) of disturbance at different spatial scales and temporal frequencies, delineation of reversible and nonreversible processes, and physical and biological constraints on the time frame for recovery. Such concepts need attention across different types of lotic ecosystems. Thus, there was strong consensus that a national monitoring system of representative lotic ecosystems within ecological regions be established. The purpose of this monitoring system would be to acquire long-term data on natural variability, to establish viable indicators of spatial and temporal aspects of recovery, and to develop and test emerging theoretical developments.     

Recovery of temperate-stream fish communities from disturbance: A review of case studies and synthesis of theory

To evaluate the relative effect of autecologic factors, site-specific factors, disturbance characteristics, and community structure on the recovery of temperate-stream fish communities, we reviewed case histories for 49 sites and recorded data on 411 recovery end points. Most data were derived from studies of low-gradient third- or fourth-order temperate streams located in forested or agricultural watersheds. Species composition, species richness, and total density all recovered within one year for over 70% of systems studied. Lotic fish communities were not resilient to press disturbances (e.g., mining, logging, channelization) in the absence of mitigation efforts (recovery time >5 to >52 yr) and in these cases recovery was limited by habitat quality. Following pulse disturbances, autecological factors, site-specific factors, and disturbance-specific factors all affected rates of recovery. Centrarchids and minnows were most resilient to disturbance, while salmonid populations were least resilient of all families considered. Species within rock-substrate/nest-spawning guilds required significantly longer time periods to either recolonize or reestablish predisturbance population densities than did species within other reproductive guilds. Recovery was enhanced by the presence of refugia but was delayed by barriers to migration, especially when source populations for recolonization were relatively distant. Median population recovery times for systems in which disturbances occurred during or immediately prior to spawning were significantly less than median recovery times for systems in which disturbances occurred immediately after spawning. There was little evidence for the influence of biotic interactions on recovery rates.     

Self-purification processes along a medium-sized stream

Water quality levels and loads of nutrients transported by the Agüera stream (northern Spain) were studied for a year to assess the self-purification capacity of this system. The main villages produce an increase of nutrient concentrations and a degradation of water quality. Nevertheless, the high retention capacity, especially for phosphate, allows the stream to recover its previous levels of quality after a short reach. The retention of nutrients depends on a complex combination of the flow level, hydrologic stability and the development of periphytic communities.     

The Role of Periphyton in Mediating the Effects of Pollution in a Stream Ecosystem

The effects of pollutants on primary producers ramify through ecosystems because primary producers provide food and structure for higher trophic levels and they mediate the biogeochemical cycling of nutrients and contaminants. Periphyton (attached algae) were studied as part of a long-term biological monitoring program designed to guide remediation efforts by the Department of Energy’s Y-12 National Security Complex on East Fork Poplar Creek (EFPC) in Oak Ridge, Tennessee. High concentrations of nutrients entering EFPC were responsible for elevated periphyton production and placed the stream in a state of eutrophy. High rates of primary production at upstream locations in EFPC were associated with alterations in both invertebrate and fish communities. Grazers represented >50% of the biomass of invertebrates and fish near the Y-12 Complex but <10% at downstream and reference sites. An index of epilithic periphyton production accounted for 95% of the site-to-site variation in biomass of grazing fish. Analyses of heavy metals in EFPC periphyton showed that concentrations of zinc, cadmium, copper and nickel in periphyton decreased exponentially with distance downstream from Y-12. Zinc uptake by periphyton was estimated to reduce the concentration of this metal in stream water ~60% over a 5-km reach of EFPC. Management options for mitigating eutrophy in EFPC include additional reductions in nutrient inputs and/or allowing streamside trees to grow and shade the stream. However, reducing periphyton growth may lead to greater downstream transport of contaminants while simultaneously causing higher concentrations of mercury and PCBs in fish at upstream sites.     

Relationships among nutrients, chlorophyll-a, and dissolved oxygen in agricultural streams in Illinois

A better understanding of the controls on algae and dissolved O2 in agricultural streams of Illinois is needed to aid in development of nutrient standards. We investigated the relationships between dissolved nutrients, algal abundance, and dissolved O2 in five streams in east-central Illinois from March through November 2004. The streams drained watersheds from 25 to 777 km2 that were dominated by row crop agriculture. Three sites had open canopies and two were bordered by a narrow forest of deciduous trees. Algal abundance was measured as chlorophyll-a (chl-a) concentration in the water column (sestonic) and on the streambed (periphytic). Mean NO3-N concentrations ranged from 5.5 to 8.8 mg N L(-1) and did not relate to algal abundance. Sestonic chl-a values ranged from nearly zero to >15 mg m(-3) with no differences between open and shaded streams and only a weak correlation with dissolved reactive P (mean concentrations were 44-479 microg L(-1)). The results suggest that sestonic chl-a is a poor criterion for assessing nutrient-related problems in these streams. Greatest periphytic chl-a occurred during low flow from August through October, but periphyton occurred consistently in only two of the five streams. The abundance of filamentous algae explained 64% of the variation in diel O2 saturation, but was not correlated with nutrients. Currently it appears that hydrology and light, rather than nutrients, control algal abundance in these streams, and in the agricultural landscape of east-central Illinois, it may not be possible to reduce nutrient concentrations sufficiently to limit filamentous algal blooms.     

Periphyton Nutrient Limitation and Maximum Potential Productivity in the Beaver Lake Basin,United States1

Ludwig, Andrea, Marty Matlock, Brian Haggard, and Indrajeet Chaubey, 2012. Periphyton Nutrient Limitation and Maximum Potential Productivity in the Beaver Lake Basin, United States. Journal of the American Water Resources Association (JAWRA) 48(5): 896‐908. DOI: 10.1111/j.1752‐1688.2012.00657.x Abstract: The objectives of this study were to measure periphytic growth responses to enrichment with nitrogen (N), phosphorus (P), and simultaneous N and P using in situ bioassays in streams draining Beaver Reservoir Basin, Northwest Arkansas; compare periphytic growth responses measured with in situ bioassays with a range of land use and point sources; and test the lotic ecosystem trophic status index (LETSI) as a simplifying metric to compare effects of nonpoint‐source pollutant‐limiting variables of N, P, and sediment across the basin. P limitation was observed at sites across a transect of stream orders throughout the basin; however, at the two sites with highest ambient nitrogen concentrations, limitation was often coupled with nitrogen limitation. Nutrients were at nonlimiting levels at both of two sites below wastewater treatment plants in all seasonal deployments. A Michaelis‐Menten growth equation described LETSI as a function of ambient PO₄‐P concentrations (p < 0.05); the midpoint (LETSI of 0.50) corresponded with a PO₄‐P concentration of approximately 3 μg/l. Change‐point analysis indicated a threshold point at LETSI of 0.80 and 15 μg/l PO₄‐P. These low values show that the periphytic community has a high affinity for available P, and that the watershed as a whole is sensitive to available nutrient inputs.     

Tree seedling establishment in dry tropics: an urgent need of interaction studies

The current anthropogenic activities and climate change are increasingly becoming a growing global concern for dry tropical forests. Worldwide, these ecologically important forests have degraded considerably since the past few decades due to such factors. These factors have harmful consequences on the vegetation structure and diversity especially tree seedlings, which may further aggravate climate change. Generally, the vegetation recovery is very slow and unpredictable in the dry tropics due to complex interaction among tree seedling, site (particularly, soil) and climatic conditions. We inculcated that a better understanding of the behavior of individuals of different tree species at seedling stage in dry forests is of immense importance. It is increasingly being recognized for explaining and managing the future composition of plant communities under changing environmental conditions. In this regard, the multi-factorial interaction studies under various resource–disturbance combinations are needed in dry tropical ecosystems to understand the: (1) impact of relative variability in resources and disturbances on the responses of tree seedlings of native species and (2) how the later relates to distinct functional and life history traits of the individual tree species. Most importantly, such studies would improve our limited understanding of how variation (natural and man-made) in nutrient availability, under the influence of other local environmental factors (such as water, light, grass competition, herbivory, fire, allelopathy and enhanced CO₂ conditions), would affect the dynamics of dry tropical forest community. It may help in the proper management of these forests. Moreover, it may prove helpful in the current climate change scenario, as change in forest community dynamics may have consequences on soil C sequestration and CO₂ efflux at global scale.     

Ecosystem Modeling Applied to Nutrient Criteria Development in Rivers

Threshold concentrations for biological impairment by nutrients are difficult to quantify in lotic systems, yet States and Tribes in the United States are charged with developing water quality criteria to protect these ecosystems from excessive enrichment. The analysis described in this article explores the use of the ecosystem model AQUATOX to investigate impairment thresholds keyed to biological indexes that can be simulated. The indexes selected for this exercise include percentage cyanobacterial biomass of sestonic algae, and benthic chlorophyll a. The calibrated model was used to analyze responses of these indexes to concurrent reductions in phosphorus, nitrogen, and suspended sediment in an enriched upper Midwestern river. Results suggest that the indexes would respond strongly to changes in phosphorus and suspended sediment, and less strongly to changes in nitrogen concentration. Using simulated concurrent reductions in all three water quality constituents, a total phosphorus concentration of 0.1 mg/l was identified as a threshold concentration, and therefore a hypothetical water quality criterion, for prevention of both excessive periphyton growth and sestonic cyanobacterial blooms. This kind of analysis is suggested as a way to evaluate multiple contrasting impacts of hypothetical nutrient and sediment reductions and to define nutrient criteria or target concentrations that balance multiple management objectives concurrently. Any opinions, findings, conclusions, or recommendations expressed in this article are those of the authors alone, and do not necessarily reflect the views of the U.S. Environmental Protection Agency or of the U.S. Government.     

Overview of case studies on recovery of aquatic systems from disturbance

An extensive review of the published literature identified more than 150 case studies in which some aspect of resilience in freshwater systems was reported. Approximately 79% of systems studied were lotic and the remainder lentic. Most of the stressor types were chemical with DDT (N=29) and rotenone (N=15) the most common. The most common nonchemical stressors were logging activity (N=16), flooding (N=8), dredging (N=3), and drought (N=7). The variety of endpoints to which recovery could be measured ranged from sparse data for phytoplankton (N=13), periphyton (N=6), and macrophytes (N=8) to relatively more data for fish (N=412) and macroinvertebrates (N=698). Unfortunately the same characteristics were rarely measured consistently among sites. For example, with respect to fish, more than 30 different species were studied and recovery was measured in many ways, most commonly on the basis of: (1) first reappearance of the species, (2) return time of predisturbance densities, and (3) return time of predisturbance average individual size. Based on these criteria, all systems in these studies seem to be resilient to most disturbances with most recovery times being less than three years. Exceptions included when (1) the disturbance resulted in physical alteration of the existing habitat, (2) residual pollutants remained in the system, or (3) the system was isolated and recolonization was suppressed.     

Linking Excess Nutrients,Light, and Fine Bedded Sediments to Impacts on Faunal Assemblages in Headwater Agricultural Streams1

Griffith, Michael B., F. Bernard Daniel, Matthew A. Morrison, Michael E. Troyer, James M. Lazorchak, and Joseph P. Schubauer‐Berigan, 2009. Linking Excess Nutrients, Light, and Fine Bedded Sediments to Impacts on Faunal Assemblages in Headwater Agricultural Streams. Journal of the American Water Resources Association (JAWRA) 45(6):1475‐1492. Abstract: Biological impairments in streams are typically defined by regulatory agencies in terms of altered invertebrate or fish assemblages. While nutrients, canopy cover, and sediment fines contribute to these impairments, these stressors are often defined, at least in part, by their impacts on periphyton. Path analysis can extend these assessments to impacts on invertebrates and fish by characterizing the direct and indirect relationships among variables along defined model pathways. With data from headwater tributaries in the Little Miami River, Ohio, we tested models of the impacts of nutrients [total nitrogen (TN), total phosphorus (TP), and the nitrogen to phosphorus (N/P) ratio], the percentage of (%) open canopy, and the % sand and fines on three periphyton metrics [periphytic ash‐free dry mass (AFDM), the percent abundance of cyanobacteria (% cyanobacteria), and the percent abundance of Chlorophyta (% Chlorophyta)] and, in turn, on selected invertebrate or fish metrics. Our objective was to develop and evaluate a statistical model that assesses the direct and indirect impacts of excess nutrients on macroinvertebrate and fish in these streams and demonstrate how this approach might be applicable elsewhere. The results suggest indirect pathways for the influences of nutrients, canopy cover, and fine bedded sediments on invertebrates or fish that are mediated by their influences on periphyton. This is in addition to any direct impacts of these stressors on the invertebrate and fish metrics. In most models, all three periphyton metrics increased with % open canopy. Periphytic AFDM increased with TN, while % cyanobacteria decreased. The % cyanobacteria also decreased with % sand and fines, but % Chlorophyta increased. The metrics, percent abundance of (%) three most dominant (macroinvertebrate) taxa, % Trichoptera, and % herbivorous fish all increased with periphytic AFDM, while % climbers, % swimmers, and %Lepomis cyanellus Rafinesque decreased. Lepomis cyanellus is an indicator species, because it is generally common in these streams and relatively tolerant to various common environmental stressors. The % three most dominant macroinvertebrate taxa increased while % Hydropsychidae (Trichoptera) and %L. cyanellus decreased with % cyanobacteria. The % Trichoptera and %L. cyanellus increased with % Chlorophyta. Some macroinvertebrate metrics, such as the % burrowers and number of burrower taxa, did not have any statistically significant relationships with the periphyton metrics but did exhibit a direct pathway with % sand and fines. These analyses illustrate how path analysis can be used to estimate the relationships among the variables in a conceptual model, modify the model, assess the relative importance of different paths, and explore responses resulting from stressors with interacting and indirect impacts.     

Minimizing Impacts of Maintenance Dredged Material Disposal in the Coastal Environment: A Habitat Approach

At present, coastal disposal of maintenance dredged material constitutes one of the most important problems in coastal zone management and in some coastal areas represents the major anthropogenic disturbance to the benthos. In this review we first propose, based on the classic literature, that macrofaunal communities typical of environmentally stressed habitats are more resilient than those of more environmentally stable habitats, and we outline the macrofaunal successional changes following a disturbance. Second, from a review and analysis of the published and unpublished literature on macrofaunal recovery following maintenance dredged material deposition in the coastal environment, we compare the successional sequences and recovery rates in euhaline and polyhaline systems. The review reveals that invertebrate recovery following dredged material disposal in relatively unstressed marine environments generally takes between 1 and 4 years, while in more naturally stressed areas, recovery is generally achieved within 9 months, although deeper polyhaline habitats can take up to 2 years to recover. Differences in recovery times are attributed to the number of successional stages required to regain the original community composition and that species typical of naturally unstressed assemblages do not possess life-history traits to allow rapid recolonization of disturbances. In the last section of this review, the management implications of these findings are discussed in terms of minimizing dredged material disposal impacts on fisheries resources. Since the natural disturbance regime appears to be very important in determining the response of a benthic community following dredged material disposal, it is recommended that when predicting the potential environmental impact of an operation, the nature of the physical environment in combination with the status (and role) of associated marine benthic communities should be considered.     

Lack of theoretical basis for predicting rate and pathways of recovery

An inadequate basis for precisely predicting the outcome of lotic ecosystem recovery, whether due to unaided natural processes or management techniques or both, exists because: (1) the field of ecology has not yet matured as a rigorous predictive science; (2) the precise sequence of events, including climatic occurrences, affecting the recovery process may be unique events and thus rarely or never repeated; and (3) even when attempts are made to control the recolonization process through introduction of species, etc., the interaction of these species may not follow deterministic models. Although this symposium focuses on lotic ecosystems, such systems are influenced strongly by exports from the surrounding land mass and, under certain circumstances, this may be the overriding influence on the recovery process; therefore, unless the boundary conditions are determined realistically, the recovery process may not follow desirable pathways. Despite the lack of a robust theoretical support base for lotic ecosystem recovery, some remarkable and rapid recoveries have occurred to either a close approximation of the original condition or to a condition ecologically superior to the damaged condition. In some cases, the recovery was due entirely to natural processes and, in others, often followed relatively straightforward management practices. There is evidence indicating that lotic ecosystem restoration is both cost effective and likely to produce satisfying results relatively rapidly. It is both fortunate that this is the case, since society is likely to support such efforts when the results have been extraordinarily successful, and unfortunate since restoration ecology needs a predictive capability.     

Benthic Algal (Periphyton) Growth Rates in Response to Nitrogen and Phosphorus: Parameter Estimation for Water Quality Models

Nitrogen (N) and phosphorus (P) are significant pollutants that can stimulate nuisance blooms of algae. Water quality models (e.g., Water Quality Simulation Program, CE‐QUAL‐R1, CE‐QUAL‐ICM, QUAL2k) are valuable and widely used management tools for algal accrual due to excess nutrients in the presence of other limiting factors. These models utilize the Monod and Droop equations to associate algal growth rate with dissolved nutrient concentration and intracellular nutrient content. Having accurate parameter values is essential to model performance; however, published values for model parameterization are limited, particularly for benthic (periphyton) algae. We conducted a 10‐day mesocosm experiment and measured diatom‐dominated periphyton biomass accrual through time as chlorophyll a (chl a) and ash‐free dry mass (AFDM) in response to additions of N (range 5–11,995 µg nitrate as nitrogen [NO₃‐N]/L) and P (range 0.89–59.51 µg soluble reactive phosphorus/L). Resulting half‐saturation coefficients and growth rates are similar to other published values, but minimum nutrient quotas are higher than those previously reported. Saturation concentration for N ranged from 150 to 2,450 µg NO₃‐N/L based on chl a and from 8.5 to 60 µg NO₃‐N/L when based on AFDM. Similarly, the saturation concentration for P ranged from 12 to 29 µg‐P/L based on chl a, and from 2.5 to 6.1 µg‐P/L based on AFDM. These saturation concentrations provide an upper limit for streams where diatom growth can be expected to respond to nutrient levels and a benchmark for reducing nutrient concentrations to a point where benthic algal growth will be limited.     

Island biogeographical theory: Can it be used to predict lotic recovery rates?

Classic island biogeographic theory predicts that equilibrium will be reached when immigration and extinction rates are equal. These rates are modified by number of species in source area, number of intermediate islands, distance to recipient island, and size of intermediate islands. This general model has been variously modified and proposed to be a stochastic process with minimal competitive interaction or heavily deterministic. Predictive models of recovery (regardless of the end point chosen) have been based on the appropriateness of the MacArthur-Wilson models. Because disturbance frequency, severity, and intensity vary in their effect on community dynamics, we propose that disturbance levels should first be defined before evaluating the applicability of island biogeographical theory. Thus, we suggest a classification system of four disturbance levels based on recovery patterns by primary and secondary succession and faunal organization by primary (invasion of vacant areas) and secondary (remnant of previous community remains) processes. Level 1A disturbances completely destroy communities with no upstream or downstream sources of colonizers, while some component of near surface interstitial or hyporheic flora and fauna survive level 1B disturbances. Recovery has been reported to take from five years to longer than 25 years, when most invading colonists do not have an aerial form. Level 2 disturbances destroy the communities but leave upstream and downstream colonization sources (level 2A) and, sometimes, a hyporheic pool of colonizers (level 2B). Recovery studies have indicated primary succession and faunal structuring patterns (2A) with recovery times of 90–400 days or secondary succession and faunal structuring patterns (2B) with recovery times of 40–250 days. Level 3 disturbances result in reduction in species abundance and diversity along a stream reach; level 4 disturbances result in reduction of abundance and diversity in discrete patches. Both disturbance types lead to secondary succession and secondary faunal organization. Recovery rates can be quite rapid, varying from less than 10 days to 100 or more days. We suggest that island biogeographical models seem appropriate to recovery by secondary processes after level 3 and 4 disturbances, where competition may be an important organizing factor, while models of numerical abundance and resource tracking are probably of better use where community development is by primary succession (levels 1 and 2). Development of predictive recovery models requires research that addresses a number of fundamental questions. These include the role of hydrologic patterns on colonization dynamics, the role of nonaerial colonizers in recovery from level 1 disturbances, and assessment of the impact of changes in the order of invasion by colonizers of varying energetic efficiencies. Finally, we must be able to assemble these data and determine whether information that guides community organization at one level of disturbance can provide insights into colonization dynamics at other levels.