Salinity as a driver of aquatic invertebrate colonisation behaviour and distribution in the wheatbelt of Western Australia

To understand how environmental change will modify community assembly and the distribution of organisms it is valuable to understand mechanisms that drive the occurrence of organisms across the landscape. Salinisation of agricultural land in southwest Western Australia, as a result of land clearing, is a widespread environmental change, which threatens numerous taxa, but provides an opportunity to elucidate such mechanisms. Although salinisation affects terrestrial fauna and flora, the greatest impacts are seen in wetlands and waterways. Many aquatic insect taxa colonise ephemeral water bodies directly as adults or by oviposition. Few empirical studies, however, evaluate the influence of abiotic factors, such as water body salinity, on the colonisation behaviour of aquatic fauna. We conducted a manipulative experiment using mesocosms to test whether colonising insect fauna select aquatic habitats based upon salinity. We found that halosensitive fauna selected less saline mesocosms for oviposition and colonisation, demonstrating that behaviour can influence the distribution of aquatic organisms. Additionally, we utilised field surveys of insects from ephemeral water bodies across a broad region of southwest Western Australia to determine if mesocosm results reflected field observation. The abundance of the same insect taxa and taxonomic groups in the field were highly variable and, with the exceptions of Culex australicus Dobrotworksy and Drummond and Anopheles annulipes Giles (Diptera: Culicidae), did not show similar patterns of distribution to those observed in the mesocosm experiment. Both mesocosm and field assemblages exhibited similar and significant trajectories associated with the salinity gradient, even though there were differences in assemblage structure between the two. Our findings give empirical support to the importance of behaviour in the spatial distribution and assembly of some aquatic insects. Handling editor: K. Martens     

Restoration of a subtropical eutrophic shallow lake in China: effects on nutrient concentrations and biological communities

Disturbances caused by rainfall are common in streams with the impact on stream inhabitants determined by the frequency, intensity, and predictability of the event. Here, we examine the response of winter-emerging Chironomidae (Diptera) to extreme flooding disturbance. In August of 2007, a severe flood impacted southeastern Minnesota, imparting stress on aquatic communities. Chironomid pupal exuviae collections were obtained biweekly from 18 southeast Minnesota streams during the following winter to assess resistance and resilience of winter-active chironomids to flooding. Streams examined were divided into moderate (2.5–10 cm), heavy (10–20 cm), or extreme (20+ cm) rainfall categories with rainfall amounts in each category representing total precipitation during the 3-day storm. Post-flood samples were compared to samples obtained from the same localities during prior winters. Our findings contradict studies of responses by Chironomidae to flooding during warmer-water conditions and show that winter-emerging Chironomidae are resistant to stress imposed by summer spates. Significantly more taxa emerged during winter after flooding as compared to historic collections, and the number of species emerging in winter was positively correlated with rainfall severity, indicating that 15 species responded opportunistically to disturbance. This indicates that winter-active Chironomidae may be resistant to increased severity of summer spates associated with climate change predictions.     

Influence of fishes on macroinvertebrate communities and insect emergence production in intermittent stream refuges

1. Drying intermittent stream networks often have permanent water refuges that are important for recolonisation. These habitats may be hotspots for interactions between fishes and invertebrates as they become isolated, but densities and diversity of fishes in these refuges can be highly variable across time and space.
2. Insect emergence from streams provides energy and nutrient subsidies to riparian habitats. The magnitude of such subsidies may be influenced by in-stream predators such as fishes.
3. We examined whether benthic macroinvertebrate communities, emerging adult insects, and algal biomass in permanent grassland stream pools differed among sites with naturally varying densities of fishes. We also manipulated fish densities in a mesocosm experiment to address how fishes might affect colonisation during recovery from hydrologic disturbance.
4. Fish biomass had a negative impact on invertebrate abundance, but not biomass or taxa richness, in natural pools. Total fish biomass was not correlated with total insect emergence in natural pools, but orangethroat darter (Etheostoma spectabile) biomass was inversely correlated with emerging Chironomidae biomass and individual midge body size. The interaction in our models between predatory fish biomass and date suggested that fishes may also delay insect emergence from natural pools, altering the timing of aquatic–terrestrial subsidies.
5. There was an increase over time in algal biomass (chlorophyll-a) in mesocosms, but this did not differ among fish density treatments. Regardless, fish presence in mesocosms reduced the abundance of colonising insects and total invertebrate biomass. Mesocosm invertebrate communities in treatments without fishes were characterised by more Chironomidae, Culicidae, and Corduliidae.
6. Results suggest that fishes influence invertebrates in habitats that represent important refuges during hydrologic disturbance, hot spots for subsidy exports to riparian food webs, and source areas for colonists during recovery from hydrologic disturbance. Fish effects in these systems include decreasing invertebrate abundance, shifting community structure, and altering patterns of invertebrate emergence and colonisation.

     

Climate change modifies the size structure of assemblages of emerging aquatic insects

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Enhanced interannual precipitation variability increases plant functional diversity that in turn ameliorates negative impact on productivity

Although precipitation interannual variability is projected to increase due to climate change, effects of changes in precipitation variance have received considerable less attention than effects of changes in the mean state of climate. Interannual precipitation variability effects on functional diversity and its consequences for ecosystem functioning are assessed here using a 6‐year rainfall manipulation experiment. Five precipitation treatments were switched annually resulting in increased levels of precipitation variability while maintaining average precipitation constant. Functional diversity showed a positive response to increased variability due to increased evenness. Dominant grasses decreased and rare plant functional types increased in abundance because grasses showed a hump‐shaped response to precipitation with a maximum around modal precipitation, whereas rare species peaked at high precipitation values. Increased functional diversity ameliorated negative effects of precipitation variability on primary production. Rare species buffered the effect of precipitation variability on the variability in total productivity because their variance decreases with increasing precipitation variance.     

Conceptualising the interactive effects of climate change and biological invasions on subarctic freshwater fish

Climate change and species invasions represent key threats to global biodiversity. Subarctic freshwaters are sentinels for understanding both stressors because the effects of climate change are disproportionately strong at high latitudes and invasion of temperate species is prevalent. Here, we summarize the environmental effects of climate change and illustrate the ecological responses of freshwater fishes to these effects, spanning individual, population, community and ecosystem levels. Climate change is modifying hydrological cycles across atmospheric, terrestrial and aquatic components of subarctic ecosystems, causing increases in ambient water temperature and nutrient availability. These changes affect the individual behavior, habitat use, growth and metabolism, alter population spawning and recruitment dynamics, leading to changes in species abundance and distribution, modify food web structure, trophic interactions and energy flow within communities and change the sources, quantity and quality of energy and nutrients in ecosystems. Increases in temperature and its variability in aquatic environments underpin many ecological responses; however, altered hydrological regimes, increasing nutrient inputs and shortened ice cover are also important drivers of climate change effects and likely contribute to context‐dependent responses. Species invasions are a complex aspect of the ecology of climate change because the phenomena of invasion are both an effect and a driver of the ecological consequences of climate change. Using subarctic freshwaters as an example, we illustrate how climate change can alter three distinct aspects of species invasions: (1) the vulnerability of ecosystems to be invaded, (2) the potential for species to spread and invade new habitats, and (3) the subsequent ecological effects of invaders. We identify three fundamental knowledge gaps focused on the need to determine (1) how environmental and landscape characteristics influence the ecological impact of climate change, (2) the separate and combined effects of climate and non‐native invading species and (3) the underlying ecological processes or mechanisms responsible for changes in patterns of biodiversity.     

Changes in rainfall level and litter stoichiometry affect aquatic community and ecosystem processes in bromeliad phytotelmata

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Having your water and drinking it too: resource limitation modifies density regulation

Determining the interaction between extrinsic and intrinsic drivers of variation in population abundance through time continues to challenge ecologists. Chamaillé-Jammes and colleagues (this issue) examined African elephant time series to explore how water availability alters the density feedback mechanisms restricting population growth. The relationship between population growth rate and density shifted from an upward convex to a more linear form after controlling for rainfall. Spatial variation in water availability also attenuated density dependence as elephants adjusted their distribution relative to current environmental conditions. This work has important climate change implications for the conservation management of African herbivores.     

Response of a Mediterranean semiarid community to changing patterns of water supply

Climate change may cause profound effects on terrestrial ecosystems. Changes in rainfall patterns may have large effects on a wide range of biological processes such as seed germination, seedling establishment, plant growth, community composition, and population and community dynamics. Climate change models for the Mediterranean region forecast reduced annual precipitation and more extreme rain events (i.e., fewer rainy days and longer drought periods between rainfall events), along with seasonal changes. We experimentally addressed the response of a semiarid Mediterranean community to higher aridity and changes in seasonal rainfall patterns in two glasshouse experiments in which we manipulated water supply. We simulated a delay in the onset of autumn rainfalls (i.e., a longer summer drought period), decreased watering amount and frequency as predicted by climate models, and manipulated the seasonality of water supply. We found that delayed watering led to decreases in plant community productivity and to delays in flowering time, in terms of both date and number of days of water supply. Decreased watering amount and frequency, and accentuated seasonality, also diminished biomass and individuals recruited, but did not change flower phenology. Species diversity was not affected by watering delays; however, it was reduced by changes in frequency, amount and seasonality. Overall, these data underline the need to consider rainfall patterns as an important element that might alter community dynamics and ecosystem structure and functioning. Therefore, the analysis of climate change consequences must not depend on climatic means-based scenarios but must take into account expected seasonal changes in rainfall quantity and frequency.     

Composition and Structure of the Chironomidae (Insecta: Diptera) Community Associated with Bryophytes in a First-Order Stream in the Atlantic Forest, Brazil

This study describes the structure of the Chironomidae community associated with bryophytes in a first-order stream located in a biological reserve of the Atlantic Forest, during two seasons. Samples of bryophytes adhered to rocks along a 100-m stretch of the stream were removed with a metal blade, and 200-mL pots were filled with the samples. The numerical density (individuals per gram of dry weight), Shannon’s diversity index, Pielou’s evenness index, the dominance index (DI), and estimated richness were calculated for each collection period (dry and rainy). Linear regression analysis was employed to test the existence of a correlation between rainfall and the individual’s density and richness. The high numerical density and richness of Chironomidae taxa observed are probably related to the peculiar conditions of the bryophyte habitat. The retention of larvae during periods of higher rainfall contributed to the high density and richness of Chironomidae larvae. The rarefaction analysis showed higher richness in the rainy season related to the greater retention of food particles. The data from this study show that bryophytes provide stable habitats for the colonization by and refuge of Chironomidae larvae, mainly under conductions of faster water flow and higher precipitation.     

Weakening density dependence from climate change and agricultural intensification triggers pest outbreaks: a 37‐year observation of cotton bollworms

Understanding drivers of population fluctuation, especially for agricultural pests, is central to the provision of agro‐ecosystem services. Here, we examine the role of endogenous density dependence and exogenous factors of climate and human activity in regulating the 37‐year population dynamics of an important agricultural insect pest, the cotton bollworm (Helicoverpa armigera), in North China from 1975 to 2011. Quantitative time‐series analysis provided strong evidence explaining long‐term population dynamics of the cotton bollworm and its driving factors. Rising temperature and declining rainfall exacerbated the effect of agricultural intensification on continuously weakening the negative density dependence in regulating the population dynamics of cotton bollworms. Consequently, ongoing climate change and agricultural intensification unleashed the tightly regulated pest population and triggered the regional outbreak of H. armigera in 1992. Although the negative density dependence can effectively regulate the population change rate to fluctuate around zero at stable equilibrium levels before and after outbreak in the 1992, the population equilibrium jumped to a higher density level with apparently larger amplitudes after the outbreak. The results highlight the possibility for exogenous factors to induce pest outbreaks and alter the population regulating mechanism of negative density dependence and, thus, the stable equilibrium of the pest population, often to a higher level, posing considerable risks to the provision of agro‐ecosystem services and regional food security. Efficient and timely measures of pest management in the era of Anthropocene should target the strengthening and revival of weakening density dependence caused by climate change and human activities.     

Impacts of climate change on national biodiversity population trends

Climate change has had well‐documented impacts on the distribution and phenology of species across many taxa, but impacts on species’ abundance, which relates closely to extinction risk and ecosystem function, have not been assessed across taxa. In the most comprehensive multi‐taxa comparison to date, we modelled variation in national population indices of 501 mammal, bird, aphid, butterfly and moth species as a function of annual variation in weather variables, which through time allowed us to identify a component of species’ population growth that can be associated with post‐1970s climate trends. We found evidence that these climate trends have significantly affected population trends of 15.8% of species, including eight with extreme (> 30% decline per decade) negative trends consistent with detrimental impacts of climate change. The modelled effect of climate change could explain 48% of the significant across‐species population decline in moths and 63% of the population increase in winged aphids. The other taxa did not have significant across‐species population trends or consistent climate change responses. Population declines in species of conservation concern were linked to both climatic and non‐climatic factors respectively accounting for 42 and 58% of the decline. Evident differential impacts of climate change between trophic levels may signal the potential for future ecosystem disruption. Climate change has therefore already driven large‐scale population changes of some species, had significant impacts on the overall abundance of some key invertebrate groups and may already have altered biological communities and ecosystems in Great Britain.     

Does Facilitation of Faunal Recruitment Benefit Ecosystem Restoration? An Experimental Study of Invertebrate Assemblages in Wetland Mesocosms

We used wetland mesocosms (1) to experimentally assess whether inoculating a restored wetland site with vegetation/sediment plugs from a natural wetland would alter the development of invertebrate communities relative to unaided controls and (2) to determine if stocking of a poor invertebrate colonizer could further modify community development beyond that due to simple inoculation. After filling mesocosms with soil from a drained and cultivated former wetland and restoring comparable hydrology, mesocosms were randomly assigned to one of three treatments: control (a reference for unaided community development), inoculated (received three vegetation/sediment cores from a natural wetland), and stocked + inoculated (received three cores and were stocked with a poorly dispersing invertebrate group—gastropods). All mesocosms were placed 100 m from a natural wetland and allowed to colonize for 82 days. Facilitation of invertebrate colonization led to communities in inoculated and stocked + inoculated treatments that contrasted strongly with those in the unaided control treatment. Control mesocosms had the highest taxa richness but the lowest diversity due to high densities and dominance of Tanytarsini (Diptera: Chironomidae). Community structure in inoculated and stocked + inoculated mesocosms was more similar to that of a nearby natural wetland, with abundance more evenly distributed among taxa, leading to diversity that was higher than in the control treatment. Inoculated and stocked + inoculated communities were dominated by non‐aerial invertebrates, whereas control mesocosms were dominated by aerial invertebrates. These results suggest that facilitation of invertebrate recruitment does indeed alter invertebrate community development and that facilitation may lead to a more natural community structure in less time under conditions simulating wetland restoration.     

Contrasting consequences of climate change for migratory geese: Predation,density dependence and carryover effects offset benefits of high‐arctic warming

Climate change is most rapid in the Arctic, posing both benefits and challenges for migratory herbivores. However, population‐dynamic responses to climate change are generally difficult to predict, due to concurrent changes in other trophic levels. Migratory species are also exposed to contrasting climate trends and density regimes over the annual cycle. Thus, determining how climate change impacts their population dynamics requires an understanding of how weather directly or indirectly (through trophic interactions and carryover effects) affects reproduction and survival across migratory stages, while accounting for density dependence. Here, we analyse the overall implications of climate change for a local non‐hunted population of high‐arctic Svalbard barnacle geese, Branta leucopsis, using 28 years of individual‐based data. By identifying the main drivers of reproductive stages (egg production, hatching and fledging) and age‐specific survival rates, we quantify their impact on population growth. Recent climate change in Svalbard enhanced egg production and hatching success through positive effects of advanced spring onset (snow melt) and warmer summers (i.e. earlier vegetation green‐up) respectively. Contrastingly, there was a strong temporal decline in fledging probability due to increased local abundance of the Arctic fox, the main predator. While weather during the non‐breeding season influenced geese through a positive effect of temperature (UK wintering grounds) on adult survival and a positive carryover effect of rainfall (spring stopover site in Norway) on egg production, these covariates showed no temporal trends. However, density‐dependent effects occurred throughout the annual cycle, and the steadily increasing total flyway population size caused negative trends in overwinter survival and carryover effects on egg production. The combination of density‐dependent processes and direct and indirect climate change effects across life history stages appeared to stabilize local population size. Our study emphasizes the need for holistic approaches when studying population‐dynamic responses to global change in migratory species.     

Rainfall induces time-lagged changes in the proportion of tropical aquatic hosts infected with metazoan parasites

Rainfall serves as a powerful driving force, shifting temporal abundance and prevalence patterns in parasites and free-living aquatic organisms in tropical environments. However, there is a lack of sound evidence showing the temporal scales at which rainfall influences infection parameters of parasites in the tropics either directly by affecting the parasite life cycle or indirectly by modifying host population abundance. In the present study, we demonstrate that changes in rainfall patterns lead to changes in the proportion of infected hosts with several parasite species, causing immediate or lagged favourable conditions for an increase in levels of infection. However, the temporal scale of the influence of rainfall varied depending on the ecological characteristics of aquatic ecosystems. Despite the environmental heterogeneity and stochastic events (storms and hurricanes) which affect the study sites, the proportion of infected hosts shows frequency cycles on a yearly scale, suggesting that environmental changes are within the range of variability that naturally occur at the study sites. We propose that the incorporation of stochastic events into long-term predictive models is crucial for understanding the potential effects of global climate change on infection parameters of tropical parasites.     

Frog population viability under present and future climate conditions: a Bayesian state-space approach

1.?World-wide extinctions of amphibians are at the forefront of the biodiversity crisis, with climate change figuring prominently as a potential driver of continued amphibian decline. As in other taxa, changes in both the mean and variability of climate conditions may affect amphibian populations in complex, unpredictable ways. In western North America, climate models predict a reduced duration and extent of mountain snowpack and increased variability in precipitation, which may have consequences for amphibians inhabiting montane ecosystems. 2.?We used Bayesian capture-recapture methods to estimate survival and transition probabilities in a high-elevation population of the Columbia spotted frog (Rana luteiventris) over 10?years and related these rates to interannual variation in peak snowpack. Then, we forecasted frog population growth and viability under a range of scenarios with varying levels of change in mean and variance in snowpack. 3.?Over a range of future scenarios, changes in mean snowpack had a greater effect on viability than changes in the variance of snowpack, with forecasts largely predicting an increase in population viability. Population models based on snowpack during our study period predicted a declining population. 4.?Although mean conditions were more important for viability than variance, for a given mean snowpack depth, increases in variability could change a population from increasing to decreasing. Therefore, the influence of changing climate variability on populations should be accounted for in predictive models. The Bayesian modelling framework allows for the explicit characterization of uncertainty in parameter estimates and ecological forecasts, and thus provides a natural approach for examining relative contributions of mean and variability in climatic variables to population dynamics. 5.?Longevity and heterogeneous habitat may contribute to the potential for this amphibian species to be resilient to increased climatic variation, and shorter-lived species inhabiting homogeneous ecosystems may be more susceptible to increased variability in climate conditions.     

Evolutionary refugia and ecological refuges: key concepts for conserving Australian arid zone freshwater biodiversity under climate change

Refugia have been suggested as priority sites for conservation under climate change because of their ability to facilitate survival of biota under adverse conditions. Here, we review the likely role of refugial habitats in conserving freshwater biota in arid Australian aquatic systems where the major long‐term climatic influence has been aridification. We introduce a conceptual model that characterizes evolutionary refugia and ecological refuges based on our review of the attributes of aquatic habitats and freshwater taxa (fishes and aquatic invertebrates) in arid Australia. We also identify methods of recognizing likely future refugia and approaches to assessing the vulnerability of arid‐adapted freshwater biota to a warming and drying climate. Evolutionary refugia in arid areas are characterized as permanent, groundwater‐dependent habitats (subterranean aquifers and springs) supporting vicariant relicts and short‐range endemics. Ecological refuges can vary across space and time, depending on the dispersal abilities of aquatic taxa and the geographical proximity and hydrological connectivity of aquatic habitats. The most important are the perennial waterbodies (both groundwater and surface water fed) that support obligate aquatic organisms. These species will persist where suitable habitats are available and dispersal pathways are maintained. For very mobile species (invertebrates with an aerial dispersal phase) evolutionary refugia may also act as ecological refuges. Evolutionary refugia are likely future refugia because their water source (groundwater) is decoupled from local precipitation. However, their biota is extremely vulnerable to changes in local conditions because population extinction risks cannot be abated by the dispersal of individuals from other sites. Conservation planning must incorporate a high level of protection for aquifers that support refugial sites. Ecological refuges are vulnerable to changes in regional climate because they have little thermal or hydrological buffering. Accordingly, conservation planning must focus on maintaining meta‐population processes, especially through dynamic connectivity between aquatic habitats at a landscape scale.     

Body size variation of the floodwater mosquito Aedes albifasciatus in Central Argentina

An inverse relationship between larval density and adult body size has been reported for several mosquito species, affecting their survival and vector competence, response to repellents and other factors. Larvae of the floodwater mosquito Aedes (Ochlerotatus) albifasciatus (Macquart) (Diptera: Culicidae) develop quickly in temporary pools, so intraspecific competition (for food or space) might regulate population abundance and affect the size of adult mosquitoes. We investigated the temporal variation of adult female wing-length (an index of body-size) in natural populations of Ae. albifasciatus, using adults collected during each phase of the rainy season. The relationships between adult mosquito abundance, female wing-length, rainfall and temperature were analysed through simple regressions. Skewness of the frequency distribution of wing-lengths showed a strong negative relationship with mean wing-length. The distribution of wing-lengths varied seasonally and was correlated with rainfall 7-15 days previously as the major consequence of breeding site volume. Thus temporal variation of body size in natural populations of Ae. albifasciatus reflected density-dependent changes in the aquatic habitat where immature stages develop, influenced more by rainfall than by temperature or other environmental variables.     

Integrated modeling predicts shifts in waterbird population dynamics under climate change

Climate change has been identified as one of the most important drivers of wildlife population dynamics. The in‐depth knowledge of the complex relationships between climate and population sizes through density dependent demographic processes is important for understanding and predicting population shifts under climate change, which requires integrated population models (IPMs) that unify the analyses of demography and abundance data. In this study we developed an IPM based on Gaussian approximation to dynamic N‐mixture models for large scale population data. We then analyzed four decades (1972–2013) of mallard Anas platyrhynchos breeding population survey, band‐recovery and climate data covering a large spatial extent from North American prairies through boreal habitat to Alaska. We aimed to test the hypothesis that climate change will cause shifts in population dynamics if climatic effects on demographic parameters that have substantial contribution to population growth vary spatially. More specifically, we examined the spatial variation of climatic effects on density dependent population demography, identified the key demographic parameters that are influential to population growth, and forecasted population responses to climate change. Our results revealed that recruitment, which explained more variance of population growth than survival, was sensitive to the temporal variation of precipitation in the southern portion of the study area but not in the north. Survival, by contrast, was insensitive to climatic variation. We then forecasted a decrease in mallard breeding population density in the south and an increase in the northwestern portion of the study area, indicating potential shifts in population dynamics under future climate change. Our results implied that different strategies need to be considered across regions to conserve waterfowl populations in the face of climate change. Our modelling approach can be adapted for other species and thus has wide application to understanding and predicting population dynamics in the presence of global change.     

Seasonal and diel periodicity in the drift of aquatic insects in a subtropical Florida stream

A study of insect drift was conducted in a small, subtropical Florida stream from December 1971 to December 1972 to describe the seasonal and diel periodicity and to determine factors influencing behavioural drift. Paired samples of 2 h duration beginning 15 min after sunset were taken biweekly, and hourly collections over a 24-h period were made quarterly. Benthic invertebrates were collected on each date from three habitats (riffle, pool and aquatic vegetation) and temperature, dissolved oxygen and current velocity were measured. Drift rates ranged from 100 to 2125 organisms/m². h (0·03 to 0·49 organisms/m³) and were greatest in winter and early spring; minimal rates occurred in the summer months. The following six taxa, in order of relative abundance, comprised 87% of the drift: Baetis intercalaris, Cheumatopsyche sp., Stenonema exiguum, Chironomidae, Stenelmis fuscata and Simulium sp. Total drift showed no significant correlation with temperature, dissolved oxygen or mean benthic abundance and only slight correlation with current velocity (r=0·34). Stepwise, multiple regression analyses indicated that riffle density and mean size of drifting organisms were important factors influencing the drift rates of B. intercalaris (R=0·67) and S. exiguum (R=0·82); mean size, riffle density and water temperature influenced the drift of Cheumatopsyche sp. (R=0·78). The other taxa of drifting insects showed no significant correlation with the variables measured. Diel (24 h) studies of the major taxa showed marked differences in the periodicity, both within and between taxa, indicating the need for long-term studies with frequent sample intervals in subtropical habitats. A new drift pattern for the family Chironomidae, alternans type, was observed for late instars of Polypedilum halterale.