Scales of spatiotemporal variability in macroinvertebrate abundance and diversity in monsoonal streams: detecting environmental change

1. We quantified spatial and temporal variability in benthic macroinvertebrate species richness, diversity and abundance in six unpolluted streams in monsoonal Hong Kong at different scales using a nested sampling design. The spatial scales were regions, stream sites and stream sections within sites; temporal scales were years (1997–99), seasons (dry versus wet seasons) and days within seasons. 2. Spatiotemporal variability in total abundance and species richness was greater during the wet season, especially at small scales, and tended to obscure site‐ and region‐scale differences, which were more conspicuous during the dry season. Total abundance and richness were greater in the dry season, reflecting the effects of spate‐induced disturbance during the wet season. Species diversity showed little variation at the seasonal scale, but variability at the site scale was apparent during both seasons. 3. Despite marked variations in monsoonal rainfall, inter‐year differences in macroinvertebrate richness and abundance at the site scale during the wet season were minor. Inter‐year differences were only evident during the dry season when streams were at base flow and biotic interactions may structure assemblages. 4. Small‐scale patchiness within riffles was the dominant spatial scale of variation in macroinvertebrate richness, total abundance and densities of common species, although site or region was important for some species. The proportion of total variance contributed by small‐scale spatial variability increased during the dry season, whereas temporal variability associated with days was greater during the wet season. 5. The observed patterns of spatiotemporal variation have implications for detection of environmental change or biomonitoring using macroinvertebrate indicators in streams in monsoonal regions. Sampling should be confined to the dry season or, in cases where more resources are available, make use of data from both dry and wet seasons. Sampling in more than one dry season is required to avoid the potentially confounding effects of inter‐year variation, although variability at that scale was relatively small.     

Understanding landscape patterns of temporal variability in avian populations to improve environmental impact assessments

It is recognized that wildlife populations exhibit spatial and temporal variability in patterns of species richness across heterogeneous landscapes. This phenomenon can prove problematic for environmental practitioners when attempting to complete comprehensive environmental assessments (EAs) with limited field surveys. A better understanding of regional spatio-temporal patterns in population dynamics should enhance site-level decision-making. In this study, the variability of seasonal data across the Credit River Watershed, southern Ontario, is assessed for a hierarchy of conservation measures including species richness, and two conservation wildlife response guilds based on primary habitat and area sensitivity. Bird populations were monitored at 24 forest monitoring plots across the watershed by the authors twice a season from 2003 to 2010 following the protocol of Environment Canada's Forest Bird Monitoring Program. The monitoring plots are located within four land management zones identified as 1) urban, 2) transitional, 3) escarpment and 4) rural. Data from the monitoring program are used to compare species richness among plots across the watershed and among land management zones. In addition, the variability of records from each plot over the 8 year period was determined by means of the Coefficient of Variation (CV) statistic. The mean variability of these records at each site within each land management zone was determined in order to assess whether the temporal variability of bird records might affect the integrity of short term assessments. Finally, a one-way ANOVA was applied to learn whether the result of short-term assessments may be further compounded by differences in the response of selected bird guilds to landscape heterogeneity. The results show that there is a significant difference in mean richness of forest birds among the four management zones. The ANOVAs indicate that significant difference is due to the temporal variability of a) breeding forest interior birds rather than edge birds or generalist species and b) breeding area sensitive species rather than area non-sensitive species. Recommendations are made that environmental assessments targeting forest interior bird populations need to plan sampling strategies that recognize this variability, especially for sites within the transitional and urban zones. Planning in the transition or urbanizing landscape should incorporate landscape ecology principles to sustain current levels of richness in forest species.     

Improving species occupancy estimation when sampling violates the closure assumption

Site occupancy models that account for imperfect detection of species are increasingly utilized in ecological research and wildlife monitoring. Occupancy models require replicate surveys to estimate detection probability over a time period where the occupancy status at sampled sites is assumed closed. Unlike mark–recapture models, few studies have examined how violations of closure can bias occupancy estimates. Our study design allowed us to differentiate among two processes that violate the closure assumption during a sampling season: 1) repeated destructive sampling events that result in either short‐ or long‐term site avoidance by the target species and 2) sampling occurring over a time period during which non‐random movements of the target species result in variable occupancy status. We used dynamic occupancy models to quantify the potential bias in occupancy estimation associated with these processes for a terrestrial salamander system. Our results provide strong evidence of a systematic decrease in salamander occupancy within a field season. Chronic disturbance due to repeated searches of natural cover objects accelerated natural declines in species occurrence on the forest surface as summer progressed. We also observed a strong but temporary disturbance effect on salamander detection probability associated with repeated sampling within a 24‐h. period. We generalized our findings by conducting a simulation to evaluate how violations of closure can bias occupancy estimates when local extinction occurs within a sampling season. Our simulation study revealed general sensitivity of estimates from single‐season occupancy models to violations of closure, with the strength and direction of bias varying between scenarios. Bias was minimal when extinction proba bility or the number of sample occasions was relatively low. Our research highlights the importance of addressing closure in occupancy studies and we provide multiple solutions, using both design‐ and model‐based frameworks, for minimizing bias associated with non‐random changes in occupancy and repeated sampling disturbances.     

Seasonal Change of Species Diversity Patterns of Non‐volant Small Mammals along Three Subtropical Elevational Gradients

Our understanding of geographic patterns of species diversity and the underlying mechanisms is increasing rapidly, whereas the temporal variation in these patterns remains poorly understood. We examined the seasonal species richness and species turnover patterns of non‐volant small mammals along three subtropical elevational gradients in southwest China. Small mammal diversity was surveyed in two seasons (early wet season and late wet season) using a standardized sampling protocol. The comparison of species richness patterns between two seasons indicated a temporal component in magnitude and shape, with species richness at high elevations clearly increased during the late wet season. Species richness demonstrated weak correlations with modelled temperature and precipitation. The elevational pattern of species turnover measured by Chao‐Sørenson similarity index also changed seasonally, even though the temporal pattern varied with scale. Species turnover between neighboring elevations at high elevations was slower in the late wet season. Meanwhile, there was an acceleration of species turnover along the whole range of the gradient. The seasonal change in species diversity patterns may be due to population‐level increases in abundance and elevational migration, whereas seasonal variation in factors other than temperature and precipitation may play a greater role in driving seasonal diversity patterns. Our study strongly supports the seasonality in elevational patterns of small mammal diversity in subtropical montane forests. Thus it is recommended that subsequent field surveys consider temporal sampling replicate for elevational diversity studies.     

Monitoring temporal trends in spatially structured populations: how should sampling effort be allocated between space and time?

Estimating temporal trends in spatially structured populations has a critical role to play in understanding regional changes in biological populations and developing management strategies. Designing effective monitoring programmes to estimate these trends requires important decisions to be made about how to allocate sampling effort among spatial replicates (i.e. number of sites) and temporal replicates (i.e. how often to survey) to minimise uncertainty in trend estimates. In particular, the optimal mix of spatial and temporal replicates is likely to depend upon the spatial and temporal correlations in population dynamics. Although there has been considerable interest in the ecological literature on understanding spatial and temporal correlations in species’ population dynamics, little attention has been paid to its consequences for monitoring design. We address this issue using model‐based survey design to identify the optimal allocation of sampling effort among spatial and temporal replicates for estimating population trends under different levels of spatial and temporal correlation. Based on linear trends, we show that how we should allocate sampling effort among spatial and temporal replicates depends crucially on the spatial and temporal correlations in population dynamics, environmental variation, observation error and the spatial variation in temporal trends. When spatial correlation is low and temporal correlation is high, the best option is likely to be to sample many sites infrequently, particularly when observation error and/or spatial variation in temporal trends are high. When spatial correlation is high and temporal correlation is low, the best option is likely to be to sample few sites frequently, particularly when observation error and/or spatial variation in temporal trends are low. When abundances are spatially independent, it is always preferable to maximise spatial replication. This provides important insights into how spatio‐temporal monitoring programmes should be designed to estimate temporal trends in spatially structured populations.     

The inherent multidimensionality of temporal variability: how common and rare species shape stability patterns

Empirical knowledge of diversity–stability relationships is mostly based on the analysis of temporal variability. Variability, however, often depends on external factors that act as disturbances, which makes comparisons across systems difficult to interpret. Here, we show how variability can reveal inherent stability properties of ecological communities. This requires that we abandon one‐dimensional representations, in which a single variability measurement is taken as a proxy for how stable a system is, and instead consider the whole set of variability values generated by all possible stochastic perturbations. Despite this complexity, in species‐rich systems, a generic pattern emerges from community assembly, relating variability to the abundance of perturbed species. Strikingly, the contrasting contributions of different species abundance classes to variability, driven by different types of perturbations, can lead to opposite diversity–stability patterns. We conclude that a multidimensional perspective on variability helps reveal the dynamical richness of ecological systems and the underlying meaning of their stability patterns.     

Random placement models predict species–area relationships in duck communities despite species aggregation

Species–area relationships are the product of many ecological processes and their interactions. Explanations for the species–area relationship (SAR) have focused on separating putative niche‐based mechanisms that correlate with area from sampling effects caused by patches with more individuals containing more species than patches with fewer individuals. We tested the hypothesis that SARs in breeding waterfowl communities are caused by sampling effects (i.e. random placement from the regional species pool). First, we described observed SARs and patterns of species associations for fourteen species of ducks on ponds in prairie Canada. Second, we used null models, which randomly allocated ducks to ponds, to test if observed SARs and patterns of species associations differed from those expected by chance. Consistent with the sampling effects hypothesis, observed SARs were accurately predicted by null models in three different years and for diving and dabbling duck guilds. This is the first demonstration that null models can predict SARs in waterbirds or any other aquatic organisms. Observed patterns of species association, however, were not well predicted by null models as in all years there was less observed segregation among species (i.e. more aggregation) than under the random expectation, suggesting that intraspecific competition could play a role in structuring duck communities. Taken together, our results indicate that when emergent properties of ecological communities such as the SAR appear to be caused by random processes, analyses of species associations can be critical in revealing the importance of niche‐based processes (e.g. competition) in structuring ecological communities.     

Moderate patchiness optimizes heterogeneity,stability, and beta diversity in mesic grassland

Heterogeneous disturbance patterns are fundamental to rangeland conservation and management because heterogeneity creates patchy vegetation, broadens niche availability, increases compositional dissimilarity, and enhances temporal stability of aboveground biomass production. Pyrodiversity is a popular concept for how variability in fire as an ecological disturbance can enhance heterogeneity, but mechanistic understanding of factors that drive heterogeneity is lacking. Mesic grasslands are examples of ecosystems in which pyrodiversity is linked strongly to broad ecological processes such as trophic interactions because grazers are attracted to recently burned areas, creating a unique ecological disturbance referred to as the fire–grazing interaction, or pyric herbivory. But several questions about the application of pyric herbivory remain: What proportion of a grazed landscape must burn, or how many patches are required, to create sufficient spatial heterogeneity and reduce temporal variability? How frequently should patches burn? Does season of fire matter? To bring theory into applied practice, we studied a gradient of grazed tallgrass prairie landscapes created by different sizes, seasons, and frequencies of fire, and used analyses sensitive to nonlinear trends. The greatest spatial heterogeneity and lowest temporal variability in aboveground plant biomass, and greatest plant functional group beta diversity, occurred in landscapes with three to four patches (25%–33% of area burned) and three‐ to four‐year fire return intervals. Beta diversity had a positive association with spatial heterogeneity and negative relationship with temporal variability. Rather than prescribing that these results constitute best management practices, we emphasize the flexibility offered by interactions between patch number and fire frequency for matching rangeland productivity and offtake to specific management goals. As we observed no differences across season of fire, we recommend future research focus on fire frequency within a moderate proportion of the landscape burned, and consider a wider seasonal burn window.     

A comparison of estimates of relative abundance from a weakly structured mass‐participation bird atlas survey and a robustly designed monitoring scheme

Estimates of population size are frequently used in conservation. Volunteer‐conducted surveys are often the only source of information available, but their reliability is unclear. We compare data from a weakly structured national bird atlas collected by volunteer surveyors free to choose where and when to visit with data from an independent suite of monitoring surveys that used a stratified sampling design. We focus on the Mount Lofty Ranges, South Australia, a region that has lost most of its native vegetation. Both datasets comprise several thousand 20‐min 2‐ha searches carried out between 1999 and 2007. The atlas dataset reported more species, and covered habitats more comprehensively, but showed greater variability in the temporal and spatial distribution of survey effort. However, after we restricted the atlas dataset to native eucalypt woodlands, reporting rates from the two schemes were very strongly correlated. The structured surveys tended to record more species that are normally detected by call and the unstructured surveys recorded more species using edges and open habitats. Minimum population estimates from the two datasets agreed very well. The strength of concordance depended on whether overflying birds were included, highlighting the importance of distinguishing such records in future surveys. We conclude that appropriate calibration using selected regional surveys, including surveys to estimate absolute densities, can enable volunteer‐collected and weakly structured atlas data to be used to generate robust occupancy and minimum population estimates for many species at a regional scale.     

Autumn bird migration phenology: A potpourri of wind,precipitation and temperature effects

Climate change has caused a clear and univocal trend towards advancement in spring phenology. Changes in autumn phenology are much more diverse, with advancement, delays, and ‘no change' all occurring frequently. For migratory birds, patterns in autumn migration phenology trends have been identified based on ecological and life‐history traits. Explaining interspecific variation has nevertheless been challenging, and the underlying mechanisms have remained elusive. Radar studies on non‐species‐specific autumn migration intensity have repeatedly suggested that there are strong links with weather. In long‐term species‐specific studies, the variance in autumn migration phenology explained by weather has, nevertheless, been rather low, or a relationship was even lacking entirely. We performed a spatially explicit time window analysis of weather effects on mean autumn passage of four trans‐Saharan and six intra‐European passerines to gain insights into this apparent contradiction. We analysed data from standardized daily captures at the Heligoland island constant‐effort site (Germany), in combination with gridded daily temperature, precipitation and wind data over a 55‐year period (1960–2014), across northern Europe. Weather variables at the breeding and stopover grounds explained up to 80% of the species‐specific interannual variability in autumn passage. Overall, wind conditions were most important. For intra‐European migrants, wind was even twice as important as either temperature or precipitation, and the pattern also held in terms of relative contributions of each climate variable to the temporal trends in autumn phenology. For the trans‐Saharan migrants, however, the pattern of relative trend contributions was completely reversed. Temperature and precipitation had strong trend contributions, while wind conditions had only a minor impact because they did not show any strong temporal trends. As such, understanding species‐specific effects of climate on autumn phenology not only provides unique insights into each species' ecology but also how these effects shape the observed interspecific heterogeneity in autumn phenological trends.     

Quantification of climatic niches in birds: adding the temporal dimension

Quantification of the climatic niche from geographic occurrences is an increasingly important tool for studying species’ relationships to their environment, for example to predict responses to climate change. However, as the geographic distributions of birds are seasonally dynamic, they pose a challenge to carrying out comparable and appropriate quantification of climatic niches. In this review, we first assess how relevant seasonal dynamics are across birds as a whole by compiling a database of migratory behaviour for 10 443 bird species. Second, we examine how studies have quantified climatic niches of birds. Finally, using Australia as a case study, we investigate how well existing distribution datasets represent temporal dynamics by comparing seasonal patterns of species richness obtained from point‐occurrence data with those from range maps and assess the consequences for niche quantification. We provide a consistent classification of migratory behaviour across all birds, and find that a huge variety exists between and within species that should be considered when quantifying climatic niches. Despite this, our review of the literature revealed that seasonal dynamics have often not been accounted for. For future studies, we provide a framework for selecting appropriate occurrence data depending on migratory behaviour and data availability. Our comparison of seasonal species richness patterns obtained from extent‐of‐occurrence range maps and point‐occurrence data suggests that range maps are less able to detect temporal dynamics of bird distributions than point‐occurrence data. We conclude that seasonally explicit range maps combined with climatic data for the corresponding time period can be used to adequately quantify climatic niches for resident birds, but are not adequate to quantify the climatic niches of migratory and nomadic species. Therefore, consistent quantification of climatic niches across all birds requires temporally explicit occurrence points. As such, increasing the availability of occurrence data and methods correcting biases should be a priority.     

Proportionate spatial sampling and equal‐time sampling of mobile animals: A dilemma for inferring areal dependence

Abstract Patch or island area is one of the most frequently used variables for inference in conservation biology and biogeography, and is often used in ecological applications. Given that all of these disciplines deal with large spatial scales, exhaustive censusing is not often possible, especially when there are large numbers of patches (e.g. for replication and control purposes). Therefore, data for patches or islands are usually collected by sampling. We argue that if area is to be used as an inferential factor, then the objects under study (i.e. the patches) must be characterized on an areal basis. This necessarily means that fixed‐area sampling is inadequate (e.g. a single standard quadrat or transect set within patches irrespective of the patch area) and that some form of area‐proportionate sampling is needed (e.g. a fixed areal proportion of each patch is surveyed by random allocation of standard quadrats across each patch). However, use of area‐proportionate sampling is not usually dissociated from the increased temporal intensity of sampling that arises from using this approach. The dilemma we see is deciding how much of the area‐specificity of variables such as species richness, rare‐species indices or probabilities of occurrence of individual species is related to the area‐proportionate survey protocol and how much is due to the temporal intensity of surveys. We undertook a study in which we balanced temporal and spatial effects by increasing the time spent surveying smaller patches of vegetation to account for the area‐ratio difference. The estimated species richness of birds of the box–ironbark system of central Victoria, Australia, was found to depend strongly upon area when area‐proportionate sampling alone was performed. When time‐balancing was imposed upon area‐proportionate sampling, the differences between smaller (10‐ha) and larger (40‐ha) areas were much reduced or effectively disappeared. We show that species found in the additional surveys used to conduct the time‐balancing were significantly less abundant than species recorded in area‐proportionate sampling. This effect is probably most severe for mobile animals, but may emerge in other forms of sampling.     

Species–area and species–sampling effort relationships: disentangling the effects

Species numbers tend to increase with both the area surveyed (species–area relationship, SAR) and the number of samples taken (species–sampling effort relationship, SSER). These two relationships differ in their nature and underlying mechanisms but are not clearly distinguished in field studies. To discriminate the effects of area (spatial extent) and sampling effort (SE) on species richness, several models explicitly involving both variables were proposed and tested against 13 datasets from marine micro‐, meio‐ and macrobenthos. A combination of power SSER and piecewise power SAR terms was found to have the best fit. The effects of area and SE were both significant, but the former one was noticeably weaker. The SSERs were roughly linear in log‐log space, whereas the SARs demonstrated scale‐dependent behavior with a noticeable threshold (slope breakpoint). Species richness was almost area‐independent below this threshold (the “small area effect”, SAE) but followed typical power‐law SAR beyond the threshold. This effect was similar to the “small island effect” but occurred for arbitrarily delineated areas within continuous habitats. Parameters of the SAR curves depended on organism size. The upper limit of the SAE increased from microorganisms to meiofauna to macrofauna. Also, SAR curves for unicellular groups had significantly lower slopes. SAE is supposed to indicate a spatial range of statistical homogeneity in species composition. Its upper limit corresponds to the characteristic size of a local community (a single habitat occupied by a common species pool). Interpretations of SAR and SSER parameters in terms of α‐ and β‐diversity are proposed. Both SAR and SSER slopes obtained from univariate regressions are overestimated. This upward bias depends on sampling design, decreasing for SAR but increasing for SSER with more unequally spaced samples. Both spatial extent and sampling effort should be taken into account to disentangle properly their effects on diversity.     

The ‘standardized search’: An improved way to conduct bird surveys

Abstract Bird surveys are among the most widely used biodiversity inventories and serve as the basis for an increasing proportion of pure and applied ecological research. It is rarely possible to conduct exhaustive censuses of all individuals present at a particular site, so stopping rules are routinely used to determine when sampling should finish. Most bird survey methods use (implicit) effort‐based stopping rules, either fixed times, fixed sampling areas (quadrats) or both, to standardize samples of different sites. If between‐site variation is high, however, a fixed sampling effort will generate samples of variable completeness with samples from smaller, less complex sites being more representative and complete than samples from larger, more complex sites. More importantly, quadrat‐based methods shift the scope of the overall study from bird occurrence in sites to bird occurrence in quadrats within sites, diminishing the impact of the research given that results cannot be extrapolated to relevant biological and management scales. Here I advocate an alternative means of conducting bird surveys, whereby the entire site is sampled and a results‐based stopping rule is used to ensure sample completeness is uniform across all sites. For example, a researcher may decide to continue sampling each site until two or fewer previously unencountered species are recorded in a 40‐min period. Samples of different sites will vary in both area and duration but will all be equivalently accurate estimates of species richness. This approach allows the avifauna of entire sites (whether territories, woodland remnants or catchments) to be sampled and compared directly, generating results and implications at the appropriate scale. In addition to yielding reliable measures of species richness, data collected this way can be used to calculate estimates of sample completeness and species incidence, two valuable metrics for ecological studies. This paper includes detailed worked examples of how to conduct a ‘standardized search’ and calculate sample completeness and species incidence estimates. I encourage further research on bird survey methods, and suggest that most current methods are insufficient, inconsistent and unreliable.     

Assemblage structure and dynamics of terrestrial birds in the southwest Amazon: a camera‐trap case study

Habitat spatial distribution, seasonal variation, and activity patterns influence changes in vertebrate assemblages over time. Terrestrial birds play major roles in the dynamics of tropical forests, but there are few effective methods to study these species due to their cryptic coloration and elusive behavior. We used camera‐trap data collected during 16 mo (February 2017–June 2018) to describe the terrestrial avifauna in southeastern Peru, assess to what extent the composition of terrestrial avifauna changes among seasons and across two major habitats (terra firme and floodplain forests), and determine daily activity patterns of terrestrial birds. We used overlap analyses to examine temporal co‐occurrence between ecologically similar and sympatric species. Camera traps recorded 16 species, including eight species in the family Tinamidae. Capture rates were highest for Pale‐winged Trumpeters (Psophia leucoptera; Psophiidae) and Gray‐fronted Doves (Leptolila rufaxilla; Columbidae). Species composition did not differ between habitats or seasons, and capture rates between habitats only differed for White‐throated Tinamous (Tinamus guttatus). Overlaps of activity patterns were high between ecologically similar species and species found in terra firme habitats (White‐throated Tinamous and Cinereous Tinamous, Crypturellus cinereus) and in both habitat types (Pale‐winged Trumpeters and Gray‐fronted Doves). Low numbers of captures of possibly locally rare or less abundant species hindered a complete analysis of spatial and seasonal patterns of terrestrial bird assemblages. We suggest a greater sampling effort and greater spatial replication to better understand the spatial and seasonal dynamics of the terrestrial avifauna. Further studies that assess the mechanisms that allow the coexistence of sympatric tinamous would be valuable, both in our study area and elsewhere. The use of camera traps in long‐term monitoring projects proved to be an effective tool for monitoring terrestrial birds, identifying cryptic and often rare animals to species level, and providing valuable ecological information at species and community levels.     

Dispersal evolution in temporally variable environments: implications for plant range dynamics

Dispersal—the movement of an individual from the site of birth to a different site for reproduction—is an ecological and evolutionary driver of species ranges that shapes patterns of colonization, connectivity, gene flow, and adaptation. In plants, the traits that influence dispersal often vary within and among species, are heritable, and evolve in response to the fitness consequences of moving through heterogeneous landscapes. Spatial and temporal variation in the quality and quantity of habitat are important sources of selection on dispersal strategies across species ranges. While recent reviews have evaluated the interactions between spatial variation in habitat and dispersal dynamics, the extent to which geographic variation in temporal variability can also shape range-wide patterns in dispersal traits has not been synthesized. In this paper, we summarize key predictions from metapopulation models that evaluate how dispersal evolves in response to spatial and temporal habitat variability. Next, we compile empirical data that quantify temporal variability in plant demography and patterns of dispersal trait variation across species ranges to evaluate the hypothesis that higher temporal variability favors increased dispersal at plant range limits. We found some suggestive evidence supporting this hypothesis while more generally identifying a major gap in empirical work evaluating plant metapopulation dynamics across species ranges and geographic variation in dispersal traits. To address this gap, we propose several future research directions that would advance our understanding of the interplay between spatiotemporal variability and dispersal trait variation in shaping the dynamics of current and future species ranges.     

Seasonal variation in diet breadth of folivorous Lepidoptera in the Brazilian cerrado

Documented patterns of specialization and species interactions often omit plasticity in resource use across space and time, yet such variation is an important part of species interactions. To examine temporal variation in resource use, we compared species‐ and community‐level patterns of host plant use by folivorous caterpillars between the dry and rainy seasons in four preserved areas of cerrado vegetation in the Distrito Federal, Brazil. We sampled plants and caterpillars in 10 m circular plots monthly from March 2010 to March 2011. At the community level, we found a significant increase in the mean diet breadth of dry season caterpillars relative to those collected in the rainy season. Families and species of moths varied in diet breadth, but most exhibited a seasonal expansion. In particular, intraspecific comparisons showed a 30% increase in the number of host plants used in the dry season compared to the rainy season. These results provide a clear example of how temporal variation in resource use is heterogeneous, and more generalized patterns of resource use can emerge from studies at large temporal scales. Thus, seasonal or annual heterogeneity may obscure ecologically relevant specialized interactions that occur at smaller scales.     

Estimating bird species richness: How should repeat surveys be organized in time?

Abstract Estimates of species richness for a given area require that repeat surveys be taken, so that the statistical robustness of the estimate can be assessed. But how should these repeat surveys be organized in time? Here we present a case study of Australian woodland birds, surveyed using the ‘active timed area search’ method, which has become the standard unit for the Australian Bird Atlas, a continental‐scale bird survey. To date, there has been no assessment of how estimates of species richness derived from this method are affected by the temporal organization of the repeat surveys. For instance, can conducting the repeat surveys in sequence on the same day effectively capture richness, or will additional species be obtained by repeating the surveys on different days within a season? If so, does the spacing of the repeat visits throughout the season have an effect? To answer these questions, we surveyed woodland birds in the Mount Lofty Ranges, South Australia, during late spring–summer 1999–2000, and compared the performance of two different temporal configurations of repeat visits to sites: (i) six repeat surveys performed on the same day; and (ii) three repeat surveys on different days. For both, we calculated the average number of species actually sighted and also estimated total species richness. The data supported our hypothesis that the same‐day surveys would yield fewer species and underestimate total species richness. The different‐day repeats captured significantly more species per unit of survey effort, and yielded a higher richness estimate. However, the timespan over which different‐day surveys were conducted within a season did not have a significant influence on species richness estimates, evincing a qualitative advantage to surveying on different days, regardless of the spacing of repeat visits. These results may be of assistance to conservation managers when planning cost‐efficient monitoring regimes.     

Patterns of spatial and temporal variation of macrofauna under boulders in a sheltered boulder field

Abstract Spatial and temporal patterns of abundance of animals and plants must be quantified before models to explain distributions can be developed. These patterns also provide essential data for measuring potential effects of environmental disturbances. Studies in many different habitats have shown that most organisms, particularly invertebrates, have highly variable and interactive patterns of abundance, with much variability at the smallest temporal and spatial scales. Intertidal boulder fields in New South Wales, Australia, support a diverse fauna, many species of which are relatively rare. These habitats are commonly found near rock‐platforms and in sheltered estuaries and are subjected to many human disturbances. Although there have been a few studies on the fauna in boulder fields, none has documented variability of the assemblage using multivariate and univariate techniques and most studies have not incorporated different spatial and temporal scales. This study quantifies spatial variation at three scales (metres, tens of metres alongshore and tens of metres upshore) and temporal variation at two scales (3 months and 2 years) of the assemblage of molluscs and echinoderms in a sheltered boulder field subjected to little natural or human disturbance. Multivariate analyses revealed that each site contained a distinct assemblage, mainly due to the relative abundances of a few species. Most species, those generally only found under boulders and common, widespread species, had considerable spatial variability in abundances, with more than 90% measured at the smallest scale, that is metre to metre within a site. Changes in abundances over 3 months or 2 years varied among species and sites in unpredictable ways. These data show that sampling designs to measure impacts on these fauna will need to be complex and must incorporate a number of spatial and temporal scales if they are to be able to detect impact against such a variable background.     

More individuals drive the species energy–area relationship in an experimental zooplankton community

The shape of the species–area relationship (SAR) often varies with the amount of available energy; SARs from high‐energy habitats typically have higher intercepts and steeper slopes than SARs from low‐energy habitats. Such patterns are often assumed to result from a shift in the mechanisms of coexistence between high and low energy habitats. However, a plausible but unexplored alternative mechanism emerges from proportional sampling, if there are simply more individuals in larger or more productive habitats, without the need to invoke differing coexistence mechanisms. Here, we examined proportional versus disproportional responses of a diverse assemblage of freshwater zooplankton to manipulations of experimental pond size and energy inputs. We found that higher energy treatments had higher species richness in large, but not small, ponds, leading to a steeper SAR with higher energy input. The total abundances of individuals also increased with energy in large, but not small ponds. By using a sample‐independent rarefaction technique (probability of interspecific encounter), we found that SAR patterns resulted from changes in the total, but not relative, abundance of individuals, and thus proportional, rather than disproportional, responses of species. Overall, our results emphasize the need to consider how both the total and relative abundances of species respond to ecological drivers such as energy and area before inferring the underlying mechanisms that lead to biodiversity patterns. Further, our results may implicate a proportionally smaller influence of energy on patterns of biodiversity when habitats are destroyed.