Abundance and distribution within a guild of benthic stream fishes: local processes and regional patterns

1. Spatial patterns at regional and local scales were examined for evidence that species interactions can influence distribution and abundance within a guild of benthic fishes in upland streams of Oklahoma, U.S.A. Three groups of community patterns were examined: the species–area relationship, species–habitat associations, and interspecific associations.
2. The species–area relationship for riffle habitats was compared to a null species–area model based on random placement. The observed species–area curve was steeper resulting in less species per unit area in small streams than predicted by the null model.
3. Small, species-poor streams had summed fish densities at least as high as larger, species-rich streams, suggesting density compensation.
4. Several significant patterns of negative covariation were found among species at the regional scale, before and after statistically accounting for effects due to measured habitat variables.
5. For two of these negatively covarying taxa ( Cottus carolinae and Etheostoma spectabile ), the influence of each species on the distribution of the other was evaluated experimentally in field enclosures varying in depth and current velocity. The sculpin C . carolinae caused a shift in habitat use by the darter E . spectabile , but no reciprocal shift was found.
6. These results indicate an agreement between local and regional patterns of distribution for C . carolinae and E . spectabile and suggest that biotic interactions can influence regional patterns of distribution for species within this guild.     

Abundance and distribution within a guild of benthic stream fishes: local processes and regional patterns

1. Spatial patterns at regional and local scales were examined for evidence that species interactions can influence distribution and abundance within a guild of benthic fishes in upland streams of Oklahoma, U.S.A. Three groups of community patterns were examined: the species–area relationship, species–habitat associations, and interspecific associations.
2. The species–area relationship for riffle habitats was compared to a null species–area model based on random placement. The observed species–area curve was steeper resulting in less species per unit area in small streams than predicted by the null model.
3. Small, species-poor streams had summed fish densities at least as high as larger, species-rich streams, suggesting density compensation.
4. Several significant patterns of negative covariation were found among species at the regional scale, before and after statistically accounting for effects due to measured habitat variables.
5. For two of these negatively covarying taxa ( Cottus carolinae and Etheostoma spectabile ), the influence of each species on the distribution of the other was evaluated experimentally in field enclosures varying in depth and current velocity. The sculpin C . carolinae caused a shift in habitat use by the darter E . spectabile , but no reciprocal shift was found.
6. These results indicate an agreement between local and regional patterns of distribution for C . carolinae and E . spectabile and suggest that biotic interactions can influence regional patterns of distribution for species within this guild.     

Closing the larval loop: linking larval ecology to the population dynamics of marine benthic invertebrates

The majority of marine benthic invertebrates exhibit a complex life cycle that includes separate planktonic larval, and bottom-dwelling juvenile and adult phases. To understand and predict changes in the spatial and temporal distributions, abundances, population growth rate, and population structure of a species with such a complex life cycle, it is necessary to understand the relative importance of the physical, chemical and biological properties and processes that affect individuals within both the planktonic and benthic phases. To accomplish this goal, it is necessary to study both phases within a common, quantitative framework defined in terms of some common currency. This can be done efficiently through construction and evaluation of a population dynamics model that describes the complete life cycle.

Two forms that such a model might assume are reviewed: a stage-based, population matrix model, and a model that specifies discrete stages of the population, on the bottom and in the water column, in terms of simultaneous differential equations that may be solved in both space and time. Terms to be incorporated in each type of model can be formulated to describe the critical properties and processes that can affect populations within each stage of the life cycle. For both types of model it is shown how this might be accomplished using an idealized balanomorph barnacle as an example species. The critical properties and processes that affect the planktonic and benthic phases are reviewed. For larvae, these include benthic adult fecundity and fertilization success, growth and larval stage duration, mortality, larval behavior, dispersal by currents and turbulence, and larval settlement. It is possible to predict or estimate empirically all of the key terms that should be built into the larval and benthic components of the model. Thus, the challenge of formulating and evaluating a full life cycle model is achievable. Development and evaluation of such a model will be challenging because of the diverse processes which must be considered, and because of the disparities in the spatial and temporal scales appropriate to the benthic and planktonic larval phases. In evaluating model predictions it is critical that sampling schemes be matched to the spatial and temporal scales of model resolution.     

Field experiments on the mobility of benthic invertebrates in a southern English stream

1. In each of twenty-six week-long experiments, the colonization by macroinvertebrates of boxes of natural sediment in a stony stream was measured. The experiments took place between February and November 1992 and environmental conditions prevailing during the weeks (particularly discharge and temperature) differed widely.
2. Colonization rate also varied widely between the weekly experiments and was sensitive to discharge, temperature and background benthic density, depending on the taxa considered.
3. A 'mobility index' measured colonization rate independently of background benthic density. This index was most strongly and positively correlated with discharge for the abundant stonefly Leuctra nigra, the net-spinning caddis Plectrocnemia conspersa and the Chironomidae, perhaps indicating that their mobility is physically driven by the influence of flow on drift.
4. Mobility in a second stonefly ( Nemurella pictetii ) was greatest in summer, when flows were low but temperature was high. Peak mobility in this species occurred during an ochreous bloom on the stream bed. It is likely the mobility of N. pictetii is more active than the other taxa.
5. There were thus differences among taxa in their mobility at baseflow and in their susceptibility to flow fluctuations. The population consequences of differences in mobility among taxa are discussed.     

Effects of winter warm reservoir release on benthic stream invertebrates

The benthos in three small mountain rivers, Holmevassåna and Tverråna (regulated) and Naustdalsåna (natural) 950–1 050 m a.s.l., were investigated during the summer seasons of 1971 and 1972. During the winter Holmevassåna became free of ice and snow due to the increased flow and higher temperature of the released water. In Tverråna River, the amount of released water was too small to break up the ice and snow cover, as was the case with Naustdalsåna River.The recorded fauna in the rivers was similar regarding groups and species, but differences in abundance were noted. The most conspicuous difference, however, was the drifting pattern of chironomid larvae, which reflected changes in winter growth. In Holmevassåna River the drift of larvae was highest in the daytime during July and August. In Tverråna River, larval drift was highest at night in July, but changed to daytime in August and September.It is suggested that the earlier development of insects in Holmevassåna, caused by the winter warm conditions can put adult insects into terrestrial environments, to which they are not adapted. In such cases the imagos can be immobilized to a great extent on snow-covered fields before reproduction.     

Small-sized invertebrates in a gravel stream: community structure and variability of benthic rotifers

1. The Rotifera assemblage inhabiting the streambed surface and the hyporheic zone of a gravel stream was investigated between October 1991 and October 1992. Forty-two species of Monogononta and 27 of Bdelloidea were identified. Within these two classes, dominant species differed between the surface and the hyporheic zone. At the streambed surface, the abundance of monogonont rotifers showed a seasonal pattern with significantly higher densities in pools, whereas bdelloids showed no clear temporal trend and did not differ significantly among sites. In the hyporheic zone, the depth distribution differed among the two rotifer groups, bdelloids occurred in highest densities between 0 and 30 cm sediment depth, while monogononts were most abundant at greater depths.
2. Species composition differed greatly between successive sampling dates (min. 5 to max. 26 days) at both the streambed surface and the hyporheos. At the streambed surface and in the shallow hyporheos a significantly higher percentage of species was replaced in riffles than in pools.
3. Few measured hydrophysical variables were associated with the Rotifera assemblage structure. At the streambed surface, species richness was negatively correlated with water temperature and substratum heterogeneity, and Monogononta rotifer densities declined with water depth and substratum roughness.
4. Permutation tests carried out on temporal serial correlations showed that, at riffle sites at the streambed surface, bdelloid rotifer densities, rotifer species richness and diversity did not differ significantly from a temporal, near-random pattern. The hyporheic rotifer assemblage followed similar near-random patterns.     

Frequency of injury and the ecology of regeneration in marine benthic invertebrates

Many marine invertebrates are able to regenerate lost tissue following injury, but regeneration can come at a cost to individuals in terms of reproduction, behavior and physiological condition, and can have effects that reach beyond the individual to impact populations, communities, and ecosystems. For example, removal and subsequent regeneration of clams' siphons, polychaetes' segments, and brittlestars' arms can represent significant energetic input to higher trophic levels. In marine soft-sediment habitats, injury changes infaunal bioturbation rates and thus secondarily influences sediment-mediated competition, adult-larval interactions, and recruitment success. The importance of injury and regeneration as factors affecting the ecology of marine invertebrate communities depends on the frequency of injury, as well as on individual capacity for, and speed of, regeneration. A key question to answer is: "How frequently are marine benthic invertebrates injured?" Here, I review the sources and the frequencies of injury in a variety of marine invertebrates from different benthic habitats, discuss challenges, and approaches for accurately determining injury rates in the field, consider evidence for species-specific, temporal and geographic variation in injury rates, and present examples of indirect effects of injury on marine invertebrates to illustrate how injury and regeneration can modify larger-scale ecological patterns and processes.     

Patterns and processes in abundance-body size relationships for marine benthic invertebrates

1. The nature of abundance-body size relationships in animal communities, and especially the drivers behind the observed patterns, have been a focus of persistent debate in animal ecology. In a recent review, Allen et al. (2006) categorized five mechanistic explanations behind the commonly observed polymodality in these relationships: energetic constraints; phylogenetic constraints; biogeographical determinants; habitat structure; and community interactions. Progress in understanding of these patterns and the processes underlying them have been hindered by the use of a range of methods that differ in their validity and robustness. 2. Here, we used data on invertebrate body sizes from a variety of sandy beaches in the UK to test the hypothesis that these communities display modality in their abundance-body size relationships. We quantified modality in the relationships using kernel density estimation and smoothed bootstrap resampling and then evaluated the competing explanations for this modality based on the patterns identified in conjunction with measurements of the physical beach environment. 3. We found bimodal distributions in the body size spectrum for benthic invertebrates at nine of 16 sites. There was a consistent trough in the spectrum at around 0·5-1 mm diameter, which reflected the traditional split between meiofauna and macrofauna. Beaches with finer particle sizes and more heterogeneous macrofauna hosted communities with more than two modes. 4. Our results suggest that modality in sandy beach benthic communities is unlikely to be explained by any single hypothesis. There will be an interplay between physical and biological factors, with different explanations accounting for modality at different scales.     

The benthic invertebrates of the Salton Sea: distribution and seasonal dynamics

The Salton Sea, California's largest inland water body, is an athalassic saline lake with an invertebrate fauna dominated by marine species. The distribution and seasonal dynamics of the benthic macroinvertebrate populations of the Salton Sea were investigated during 1999 in the first survey of the benthos since 1956. Invertebrates were sampled from sediments at depths of 2–12 m, shallow water rocky substrates, and littoral barnacle shell substrates. The macroinvertebrates of the Salton Sea consist of a few invasive, euryhaline species, several of which thrive on different substrates. The principal infaunal organisms are the polychaetes Neanthes succinea Frey & Leuckart and Streblospio benedicti Webster, and the oligochaetes Thalassodrilides gurwitschi Cook, T. belli Hrabe, and an enchytraeid. All but Neanthes are new records for the Sea. Benthic crustacean species are the amphipods Gammarus mucronatus Say, Corophium louisianum Shoemaker, and the barnacle Balanus amphitrite Darwin. Neanthes succinea is the dominant infaunal species on the Sea bottom at depths of 2–12 m. Area-weighted estimates of N. succinea standing stock in September and November 1999 were two orders of magnitude lower than biomass estimated in the same months in 1956. During 1999, population density varied spatially and temporally. Abundance declined greatly in offshore sediments at depths >2 m during spring and summer due to decreasing oxygen levels at the sediment surface, eventually resulting in the absence of Neanthes from all offshore sites >2 m between July and November. In contrast, on shoreline rocky substrate, Neanthes persisted year round, and biomass density increased nearly one order of magnitude between January and November. The rocky shoreline had the highest numbers of invertebrates per unit area, exceeding those reported by other published sources for Neanthes, Gammarus mucronatus, Corophium louisianum, and Balanus amphitrite in marine coastal habitats. The rocky shoreline habitat is highly productive, and is an important refuge during periods of seasonal anoxia for Neanthes and for the other invertebrates that also serve as prey for fish and birds.     

Stream flow and predation effects on the spatial dynamics of benthic invertebrates

Field experiments were carried out to determine the influence of predation and prey movements on the accumulation of prey in enclosures. Experimental enclosures permitted exchange of prey with the benthos, but not of the large, predatory larvae of the caddisfly, Plectrocnemia conspersa (Curtis). Unseasonally heavy rainfalls during the experiment resulted in high flows and enabled us to examine the effects of a major, abiotic disturbance on invertebrate spatial dynamics. Prey colonization rates of cages without predators were determined in nine 24 h periods. Colonization rates increased exponentially with flow and were species-specific, depending on dispersal behaviour. Prey accumulation and predator impacts were measured in cages, with and without P. conspersa larvae, placed in the stream for 1, 2 or 3 weeks. Prey densities in cages increased with exposure time, but increases were not gradual and depended on flow regime. Flow was reduced within cages and they accumulated large numbers of invertebrates during high discharge. Analogous, naturally occurring refugia in the stream channel could be important for the recovery of lotic communities after major disturbances. Overall, prey densities were lowest in cages with predators. For fast colonizers, predation effects were detectable early in the experiment, but quickly obscured thereafter by continuous exchange of prey. For slow colonists, predation effects were detectable later, but persisted longer. Consumption rates for P. conspersa varied with prey density and flow regime. We suggest that the spatial dynamics of benthic invertebrates, especially as they are influenced by stochastic events, are important in understanding and detecting predation effects in stream communities.     

Long-term effects of local disturbance history on mobile stream invertebrates

The patchy distribution of benthic invertebrates in streams and rivers is an important and widely researched phenomenon. Previous studies on reasons for this patchiness have neglected the potential role of local disturbance history, probably because most lotic invertebrates are mobile and any effect of disturbance history was thought to be short-lived. Here we demonstrate for a New Zealand gravel-bed stream that local disturbance history can have long-term effects on the distribution of highly mobile stream invertebrates. Buried scour chains (100 at each of three 20-m sites within a 350-m reach) indicated that a spate with a return period of 5 months caused a mosaic of bed patches with different stabilities. More than 2 months after the spate, we took random, quantitative samples at each site from five patches that had experienced 4 cm or more of scour during the spate, from five patches with 4 cm or more of fill, and from five stable patches. Density of the dominant invertebrate taxon, the highly mobile mayfly Deleatidium spp., and densities of another three of the seven most common taxa differed significantly between patch stability categories. Larvae of Deleatidium, the black fly Austrosimulium spp. and the dipteran Eriopterini were most abundant in fill patches, whereas Isopoda were most abundant in scour patches. Total invertebrate densities and densities of six common taxa also differed between sites, although these were only 95–120 m apart. These results show that local disturbance history can have long-term effects on lotic invertebrates and be an important cause of invertebrate patchiness. The observed effects might have been even stronger had we sampled sooner after the spate or after a large flood. Disturbance history may influence invertebrates both directly (through dislodgement or mortality) and indirectly, through effects on the spatial distribution of their resources. Our results suggest that the role of disturbance in structuring animal communities dominated by mobile species may be more important than previously thought. Received: 25 January 2000 / Accepted: 14 April 2000     

Microdistribution of benthic invertebrates in a rocky mountain (U.S.A.) stream

A study of the benthic invertebrate community inhabiting a small, foothill trout stream in the Rocky Mountains of Idaho was conducted over a two-year period. Monthly Hess samples and short-term experiments using substratum-filled trays were used to describe the spatial dispersion of the benthos and to examine the response of invertebrate populations to substratum and current. A method was devised for measuring available surface area which involved coating individual stones with latex and measuring the area of the print resulting from inking the impression left on the latex mold.The dispersion of all populations was clumped throughout the year. Alteration of the cross-sectional pattern of current velocity and stream bed composition changed the pattern of distribution but not the extent of clumping. Collections made in areas of depositing and eroding substrata revealed a more diverse fauna in the latter. Most groups of organisms found in the riffle were scarcer in the pools or absent from them. The pool fauna contained no important additions over those found in the riffles.After a year's study of invertebrate populations in an otherwise undisturbed riffle, the substratum was altered and the flow made more uniform; an increase in the abundance of most of the benthic invertebrates followed. No single factor was responsible for the increase, but the change in substratum size and degree of compaction accounted for most of the change. Interpretation of the results was aided by findings from experiments using substratum-filled trays.Two series of stream experiments using the trays were conducted: one to test the relative importance of current and substratum and the other to test the effect of particle size on the distribution of the benthic fauna. In the first series, placement of trays of stones in a pool resulted in an increase in numbers of some but not all of the invertebrates over numbers usually occurring in the pool. Trays filled with stones and placed in a riffle supported fewer animals than found on the adjacent stream bed but more than in the pool. Variations are attributed to differences in current velocity and amounts of imported organic and inorganic debris. Three different relationships of population numbers to current velocity were found for different members of the community (direct, indirect, and parabolic) over the range of 10 to 60 cm/sec. The second series of experiments consisted of two sets of trays filled with stones of medium or large pebbles, respectively. Nine taxa, as well as all of the combined taxa, showed a preference for trays of small stones over the natural stream bed. A few taxa were noticeably more abundant on the small substratum than on the large but most of the fauna showed only slight increases in numbers or remained constant on the two substrata. Only three taxa showed a direct relation of numbers to total surface area presented by the stones.Number and kinds of organisms found in trays filled with a uniform size of substratum did not correspond to those taken in Hess samples from the natural stream bed. This has important implications in terms of currently recommended pollution monitoring techniques. However, it is suggested that if the substratum composition of the trays more nearly matched that of the stream, the correspondence would be much better. The results of the present study also throw considerable doubt on the adequacy of generalizations derived from earlier studies of responses to substratum size and suggest several reasons for reevaluating current ideas regarding the influence of substratum on invertebrate distribution.     

Overview of the chemical ecology of benthic marine invertebrates along the western Antarctic peninsula

Thirteen years ago in a review that appeared in the American Zoologist, we presented the first survey of the chemical and ecological bioactivity of Antarctic shallow-water marine invertebrates. In essence, we reported that despite theoretical predictions to the contrary the incidence of chemical defenses among sessile and sluggish Antarctic marine invertebrates was widespread. Since that time we and others have significantly expanded upon the base of knowledge of Antarctic marine invertebrates' chemical ecology, both from the perspective of examining marine invertebrates in new, distinct geographic provinces, as well as broadening the evaluation of the ecological significance of secondary metabolites. Importantly, many of these studies have been framed within established theoretical constructs, particularly the Optimal Defense Theory. In the present article, we review the current knowledge of chemical ecology of benthic marine invertebrates comprising communities along the Western Antarctic Peninsula (WAP), a region of Antarctica that is both physically and biologically distinct from the rest of the continent. Our overview indicates that, similar to other regions of Antarctica, anti-predator chemical defenses are widespread among species occurring along the WAP. In some groups, such as the sponges, the incidence of chemical defenses against predation is comparable to, or even slightly higher than, that found in tropical marine systems. While there is substantial knowledge of the chemical defenses of benthic marine invertebrates against predators, much less is known about chemical anti-foulants. The sole survey conducted to date suggests that secondary metabolites in benthic sponges are likely to be important in the prevention of fouling by benthic diatoms, yet generally lack activity against marine bacteria. Our understanding of the sensory ecology of Antarctic benthic marine invertebrates, despite its great potential, remains in its infancy. For example, along the WAP, community-level non-consumptive effects occur when amphipods chemically sense fish predators and respond by seeking refuge in chemically-defended macroalgae. Such interactions may be important in releasing amphipods from predation pressure and facilitating their unusually high abundances along the WAP. Moreover, recent studies on the sensory biology of the Antarctic keystone sea star Odontaster validus indicate that chemotactile-mediated interactions between conspecifics and other sympatric predatory sea stars may have significant ramifications in structuring community dynamics. Finally, from a global environmental perspective, understanding how chemical ecology structures marine benthic communities along the WAP must increasingly be viewed in the context of the dramatic impacts of rapid climatic change now occurring in this biogeographic region.     

Energetic constraints, size gradients, and size limits in benthic marine invertebrates

Populations of marine benthic organisms occupy habitats witha range of physical and biological characteristics. In the intertidalzone, energetic costs increase with temperature and aerial exposure,and prey intake increases with immersion time, generating sizegradients with small individuals often found at upper limitsof distribution. Wave action can have similar effects, limitingfeeding time or success, although certain species benefit fromwave dislodgment of their prey; this also results in gradientsof size and morphology. The difference between energy intakeand metabolic (and/or behavioral) costs can be used to determinean energetic optimal size for individuals in such populations.Comparisons of the energetic optimal size to the maximum predictedsize based on mechanical constraints, and the ensuing mortalityschedule, provides a mechanism to study and explain organismsize gradients in intertidal and subtidal habitats. For specieswhere the energetic optimal size is well below the maximum sizethat could persist under a certain set of wave/flow conditions,it is probable that energetic constraints dominate. When theopposite is true, populations of small individuals can dominatehabitats with strong dislodgment or damage probability. Whenthe maximum size of individuals is far below either energeticoptima or mechanical limits, other sources of mortality (e.g.,predation) may favor energy allocation to early reproductionrather than to continued growth. Predictions based on optimalsize models have been tested for a variety of intertidal andsubtidal invertebrates including sea anemones, corals, and octocorals.This paper provides a review of the optimal size concept, andemploys a combination of the optimal energetic size model andlife history modeling approach to explore energy allocationto growth or reproduction as the optimal size is approached.     

Screening for antibacterial and antifungal activities in marine benthic invertebrates from northern Norway

Benthic marine invertebrates collected from sub-Arctic regions of northern Norway, were found to be a promising source of novel bioactive compounds against human and fish pathogenic bacteria and fungi. Lyophilized material from seven species of ascidians, six sponges and one soft alcyonid coral were extracted with 60% acidified acetonitrile (ACN). After separation into an ACN-rich phase (ACN-extract) and an aqueous phase, and subsequent solid-phase extraction of the aqueous phase, fractions differing in polarity were obtained and screened for antibacterial and antifungal activities, along with the more lipophilic ACN-extracts. Antimicrobial activity was determined against two Gram-negative, two Gram-positive bacteria, and two strains of fungi. Notably, all the invertebrate species in the study showed activity against all four strains of bacteria and the two strains of fungi. In general, the aqueous fractions displayed highest antimicrobial activity, and the most potent extracts were obtained from the colonial ascidian Synoicum pulmonaria which displayed activity against bacteria and fungi at a concentration of 0.02 mg/ml; the lowest concentration tested.     

Size-mass relationships of stream invertebrates in a northern Spain stream

We show the regression equations that relate the head width of 10 freshwater insect species or the length of the first thoracic segment of 2 amphipods with their dry mass in 3 reaches along a small stream in northern Spain. Spatial differences in size–mass relationships were found. Organisms stored in alcohol showed lower dry mass than those specimens that were dried immediately after collection. The use of both size measures for dry mass estimation in field samples is discussed. It is recommended that the size–mass relationships be constructed using non-preserved organisms. Furthermore, great care is required when such allometric equations are taken from the literature.     

Reproductive strategies of marine benthic invertebrates revisited: facultative feeding by planktotrophic larvae

Fecundity-time models of reproductive strategies in marine invertebrates all predict that reproductive success is maximized only at the extreme levels of investment. Selection should drive egg sizes toward small eggs and planktotrophy or large eggs and lecithotrophy. The existence of two distinct larval types, feeding and nonfeeding, has been taken as confirmation of this prediction and has established the current paradigm for larval ecology. However, comparative and experimental evidence does not support the prediction that egg size is minimized in species with planktotrophic larvae. Recent discoveries have documented the existence of planktotrophs that have intermediate egg sizes, differing degrees of dependence on exogenous food, and differing capacities for facultative feeding. A fecundity-time model is presented that includes facultative larval feeding by dissociating the onset of feeding capability from the need for exogenous food. The facultative feeding model shows that reproductive success can be maximized at intermediate levels of investment per offspring between the minimum for development and the threshold for lecithotrophy, depending on the amount of food available to larvae and the intensity of planktonic mortality. A continuum of larval strategies is predicted.     

Invertebrate drift and benthic community dynamics in a lowland neotropical stream, Costa Rica

In this study we quantified invertebrate drift and related it to the structure of the benthic community, over a 6–8 month period, in a 4th-order tropical stream in Costa Rica. Relative to reports from similar-sized temperate and tropical streams, drift densities were high (2-fold greater: mean 11.2 m⁻³; range 2.5–25 m⁻³), and benthic insect densities were relatively low (>3-fold lower: mean 890 m⁻²; range 228–1504 m⁻²). Drift was dominated by larval shrimps that represented more than 70% of total drift on any given date; the remaining 30% was composed of 54 insect taxa. Among insects, Simuliidae and Chironomidae (Diptera) and Baetidae, Leptohyphes and Tricorythodes (Ephemeroptera) comprised 24% of total drift. Drift periodicity was strongly nocturnal, with peaks at 18:00 h (sunset) and 03:00 h. Our results, and those of previous experiments in the study stream, suggest that nighttime drift is driven by the presence of predatory diurnal drift-feeding fishes and nocturnal adult shrimps. There were no clear seasonal patterns over both ‘dry’ and wet seasons, suggesting that benthic communities are subject to similar stresses throughout the year, and that populations grow and reproduce continuously. This revised version was published online in July 2006 with corrections to the Cover Date.     

Combined ecological factors permit classification of developmental patterns in benthic marine invertebrates: a discussion note

Traditional classifications of developmental patterns of marine benthic invertebrates are based on combinations of embryological (direct or indirect development) and ecological (such as nutritional source or habitat) characteristics. Different schemes have been proposed for different reasons, relating to ecology, evolution and/or development. However, these classifications contain interconnected characters that do not efficiently discriminate between developmental patterns and, thus, do not fully apply to either ecological or embryological studies. An ecological multifactor classification based on three independent two-state characters (pelagic/benthic, free/protected, and feeding/non-feeding) is proposed. It discriminates between eight developmental patterns and can encompass any ecological pattern of development among marine benthic invertebrates.     

Microhabitat as a determinant of diversity: stream invertebrates colonizing leaf packs

1. Macroinvertebrate colonization of artificial leaf packs of differing palatability to detritivores was measured in a low-order stream. 2. The most palatable leaf types—alder and young beech—were colonized mainly by detritivores and consumed rapidly, so that species diversity on the substrate remained low. In the case of the less palatable old beech and paper ‘leaves’, however, colonization was slower but species diversity approached that of the surrounding benthos. 3. After 1 week, species diversity in paper ‘leaf packs was equivalent to that of the benthos, but species composition was dissimilar. 4. I argue that species diversity of a patch of stream bed may be inversely related to the abundance or palatability of a given food resource, and that this relationship is maintained by the instability of the stream bed habitat, precluding a long-term community response to increased food availability. It may be, however, that the dispersal abilities of many aquatic insects allow a rapid response to the creation of novel habitats.