Ecological relationships between phytoplankton communities and different spatial scales in European reservoirs: implications at catchment level monitoring programmes

Phytoplankton communities are structured by factors acting over temporal and spatial scales. Identifying which factors are driving spatial patterns in aquatic communities is the central aim of ecology. In this study, data sets of phytoplankton communities and environmental data of two Portuguese reservoirs types (lowland “riverine reservoirs” and higher altitude “artificial lake reservoirs”) were used to determine the importance of environmental variables at different spatial (geographical, regional and local) and time scales (seasons, years) on the community structure. In all the data sets, the multivariate ordination technique Canonical Correspondence Analysis (CCA) showed that regional and local scales explained the majority (9–18% and 13–19%, respectively) of the taxa variance. However, for “riverine reservoirs”, time variables were more important, explaining 27% of the variability in phytoplankton assemblages. Variance partitioning was used to assess the individual importance of the three spatial scales and time for the community structure of the two reservoir types. The majority of among-site variability (5.9–21.4%) was accounted for by time variables, with local, regional, and geographical scale variables accounting for 3.3–5.6%, 3.7–4.5% and 2.6–2.9%, respectively. The effects of different spatial scales on phytoplankton communities were clearly interrelated; thus, implying that phytoplankton assemblages are capable of detecting stress from catchment to site scales. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users. Handling editor: J. Padisak     

Littoral macroinvertebrate communities: spatial scale and ecological relationships

SUMMARY 1. Spatial correlations between ecological patterns and processes are thought to be scale-dependent, yet surprisingly few studies have evaluated the correspondence between different levels of spatial scale and ecosystem structure and function.
2. We evaluated the strength of relationships between the benthic macroinvertebrate communities of stony littoral habitats and levels of ecological scale and geographical position, using partial constrained ordination. Our hypothesis was that correlation strength would be inversely related to ecological scale, i.e. habitat > ecosystem > riparian > catchment > ecoregion.
3. The effect of habitat was greater than that of other levels of spatial scale: 23% of the variance in taxonomic composition and 11% of that in functional composition was explained by habitat variables alone. However, greater spatial scales were also important. For example, the combined influence of riparian, catchment and ecoregion classification accounted for 24% (taxonomic) and 11% (functional) of the explained variance.
4. Relationships between organisms and scale variables were, however, non-linear and a substantial amount of the functional variance was hidden in joint effects. These findings were not unexpected, and presumably indicate a close interdependence between local and regional-scale variables.     

Variability in stream macroinvertebrates at multiple spatial scales

1. We intensively sampled 16 western Oregon streams to characterize: (1) the variability in macroinvertebrate assemblages at seven spatial scales; and (2) the change in taxon richness with increasing sampling effort. An analysis of variance (ANOVA) model calculated spatial variance components for taxon richness, total density, percent individuals of Ephemeroptera, Plecoptera and Trichoptera (EPT), percent dominance and Shannon diversity.
2. At the landscape level, ecoregion and among‐streams components dominated variance for most metrics, accounting for 43–72% of total variance. However, ecoregion accounted for very little variance in total density and 36% of the variance was attributable to differences between streams. For other metrics, variance components were more evenly divided between stream and ecoregion effects.
3. Within streams, approximately 70% of variance was associated with unstructured local spatial variation and not associated with habitat type or transect position. The remaining variance was typically split about evenly between habitat and transect. Sample position within a transect (left, centre or right) accounted for virtually none of the variance for any metric.
4. New taxa per stream increased rapidly with sampling effort with the first four to eight Surber samples (500–1000 individuals counted), then increased more gradually. After counting more than 50 samples, new taxa continued to be added in stream reaches that were 80 times as long as their mean wetted width. Thus taxon richness was highly dependent on sampling effort, and comparisons between sites or streams must be normalized for sampling effort.
5. Characterization of spatial variance structure is fundamental to designing sampling programmes where spatial comparisons range from local to regional scales. Differences in metric responses across spatial scales demonstrate the importance of designing sampling strategies and analyses capable of discerning differences at the scale of interest.     

Temporal dynamics of stream fish assemblages and the role of spatial scale in quantifying change

1. Spatial grain of studies of communities is often based on arbitrary convention. Few studies have examined how spatial scaling of grain size affects estimates of compositional change over time, despite its broad implications.
2. Fish assemblage structure was compared between 1974 and 2014 at 33 sampling locations in the Muddy Boggy River drainage, USA. The two main objectives for this comparison were to quantify change in assemblage structure and to test for a relationship between compositional change and spatial scale. Spatial scale was manipulated by pooling assemblage data into a continuous range of groups, which increased in size from K = 33 pairs (i.e., local scale) to K = 1 pair (i.e., global scale), via clustering algorithm based on pair‐wise fluvial distance.
3. Local assemblages (stream reaches) varied in the degree of assemblage change over time (range = 0.10–0.99 dissimilarity; mean = 0.66). The global assemblage (drainage), however, remained relatively similar. A discontinuity in the relationship between compositional change and spatial scale occurred at K = 15 (mean dissimilarity = 0.56; p = .062), and this grouping is roughly the size of the headwater/tributary drainages (i.e., stream order ≤ 3) in the study system.
4. Spatial scale can impact estimates of biodiversity change over time. These results suggest assemblages are more dynamic at individual stream reaches than at the scale of the entire drainage. The decline in assemblage change at the spatial scale of K = 15 deserves further attention given the marginal significance, despite a small sample size (n = 15). This pattern could suggest regional and meta‐community processes become more important in shaping assemblage dynamics at the scale of headwater drainages, whereas the factors responsible for driving individual stream reach dynamics (e.g., stochasticity) become less important. Defining assemblages at a larger scale will result in different estimates of species persistence. Biodiversity monitoring efforts must take the effect of spatial scaling into consideration.

     

Relationships between land use, spatial scale and stream macroinvertebrate communities

1. The structure of lotic macroinvertebrate communities may be strongly influenced by land‐use practices within catchments. However, the relative magnitude of influence on the benthos may depend upon the spatial arrangement of different land uses in the catchment. 2. We examined the influence of land‐cover patterns on in‐stream physico‐chemical features and macroinvertebrate assemblages in nine southern Appalachian headwater basins characterized by a mixture of land‐use practices. Using a geographical information system (GIS)/remote sensing approach, we quantified land‐cover at five spatial scales; the entire catchment, the riparian corridor, and three riparian ‘sub‐corridors’ extending 200, 1000 and 2000 m upstream of sampling reaches. 3. Stream water chemistry was generally related to features at the catchment scale. Conversely, stream temperature and substratum characteristics were strongly influenced by land‐cover patterns at the riparian corridor and sub‐corridor scales. 4. Macroinvertebrate assemblage structure was quantified using the slope of rank‐abundance plots, and further described using diversity and evenness indices. Taxon richness ranged from 24 to 54 among sites, and the analysis of rank‐abundance curves defined three distinct groups with high, medium and low diversity. In general, other macroinvertebrate indices were in accord with rank‐abundance groups, with richness and evenness decreasing among sites with maximum stream temperature. 5. Macroinvertebrate indices were most closely related to land‐cover patterns evaluated at the 200 m sub‐corridor scale, suggesting that local, streamside development effectively alters assemblage structure. 6. Results suggest that differences in macroinvertebrate assemblage structure can be explained by land‐cover patterns when appropriate spatial scales are employed. In addition, the influence of riparian forest patches on in‐stream habitat features (e.g. the thermal regime) may be critical to the distribution of many taxa in headwater streams draining catchments with mixed land‐use practices.     

Beta diversity of stream fish assemblages: partitioning variation between spatial and environmental factors

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Hierarchy,spatial configuration,and nutrient cycling in a desert stream

Abstract Recent studies of nutrient cycling in Sycamore Creek in Arizona, USA, suggest that a thorough understanding requires a spatially explicit, hierarchical approach. Physical configuration determines the path that water follows as it moves downstream. Water follows flowpaths through surface stream components, the hyporheic zone beneath the surface stream, and the parafluvial (sand bar) zone. Characteristic biogeochemical processes in these subsystems alter nitrogen (N) species in transport, in part as a function of available concentrations of N species. At several hierarchical levels, substrate materials are an important determinant of nitrogen dynamics in desert streams. Sand is present in bars of variable size and shape, each of which can be considered a unit, interacting with the surface stream. Groups of these stream-sandbar units form a higher level, the reach. At the next higher scale, sandy reaches (runs) alternate with riffles. Where flowpaths converge, rates of N transformation are high and, as a result, change in concentration is a non-linear function of flowpath length. Disturbance by flash floods alters sandbar configuration. Between floods, the interaction of subsurface and surface flowpaths shapes configuration in each, thus a self-organizing element of spatial structure exists. Sandy runs are dominated by subsurface processes and are likely to be net nitrifiers while riffles are dominated by surface flow and are nitrogen fixers. Whether a stream ecosystem retains nitrogen, or transports it to downstream recipient systems, or is a net emitter of gaseous forms of N, depends upon the dynamics of a spatial mosaic of interacting elements. An understanding of the net effect of this mosaic requires a spatially explicit, hierarchical, multi-scale approach.     

Relationships between distribution and abundance vary with spatial scale and ecological group in stream bryophytes

1. We examined whether the local abundance of stream bryophytes in a boreal drainage basin (Koutajoki system in northeastern Finland) correlated with their: (i) regional occupancy; (ii) provincial distribution in northwestern Europe; and (iii) global range size. We specifically tested whether aquatic and semi‐aquatic species differ in their distribution–abundance relationships. We also analysed the frequency distributions of occupancy at two spatial scales: within the focal drainage system and across provinces of northwestern Europe. 2. Regional occupancy and mean local abundance of stream bryophytes were positively correlated, and the relationship was rather strong in aquatic species but very weak in semi‐aquatic species. Local abundance was related neither to provincial distribution nor global distribution. 3. Species frequency distributions differed between regional occupancy and provincial distribution. While most species were rare with regard to their regional occupancy within the focal drainage system, most of the same set of species were common and occurred in most provinces in northwestern Europe. 4. The results indicate the presence of dominants (core species) and transients/subordinates (satellite species) among stream bryophytes, highlighting marked differentiation in life‐history strategies and growth form. The observed abundance–occupancy relationships suggest that dispersal limitation and metapopulation processes may govern the dynamics of obligatory aquatic stream bryophytes. In semi‐aquatic species, however, habitat availability may be more important in contributing to regional occupancy.     

Taxonomic and biological trait differences of stream macroinvertebrate communities between mediterranean and temperate regions: implications for future climatic scenarios

Streams in mediterranean regions have highly seasonal discharge patterns, with predictable torrential floods and severe droughts. In contrast, discharge is less variable in temperate regions and intermittent flow conditions are uncommon. Hydroclimatic models predict that climate change would increase frequency and severity of floods and droughts across Europe, thus increasing the proportion of streams with mediterranean characteristics in actually temperate areas. Correspondingly, understanding actual ecological differences between mediterranean and temperate streams may help to anticipate large‐scale ecological impacts of climate change. Given that large‐scale factors determine local community composition, we hypothesized that climatic differences between mediterranean and temperate regions should affect the taxonomic and biological trait composition in streams. We assembled the abundance of stream macroinvertebrate genera of 265 sites each from the Mediterranean Basin and from temperate Europe and linked these abundances to published information on 61 categories of 11 biological traits reflecting the potential of resilience from and resistance to disturbances. Although regional taxonomic richness was higher in the mediterranean than in the temperate region, local taxonomic richness and diversity did not significantly differ between regions. Local trait richness and diversity were significantly higher in the mediterranean region. Both local taxonomic and trait‐community composition differed between regions, but the former varied much more than the latter, highlighting that climate change could produce large changes in the taxonomic but rather weak changes in the trait composition. The mediterranean region was characterized by macroinvertebrates with higher dispersion and colonization capabilities, suggesting that species loss in the temperate region, by extinction or northward emigration of taxa, would be compensated for by immigration of southern mediterranean taxa. Thus, climate change would likely have stronger implications for the local conservation of taxa than for the trait composition of stream macroinvertebrate communities.     

Response of three lotic assemblages to riparian and catchment‐scale land use: implications for designing catchment monitoring programmes

1. Although many studies have focussed on the effects of catchment land use on lotic systems, the importance of broad (catchment) and fine (segment/reach) scale effects on stream assemblages remain poorly understood. 2. Nine biological metrics for macrophytes (498 sites), benthic macroinvertebrates (491) and fish (478) of lowland and mountain streams in four ecoregions of France and Germany were related to catchment and riparian buffer land use using partial Redundancy Analysis and Boosted Regression Trees (BRTs). 3. Lotic fauna was better correlated (mean max., r = 0.450) than flora (r = 0.277) to both scales of land use: the strongest correlations were noted for mountain streams. BRTs revealed strong non‐linear relationships between mountain assemblage metrics and land use. Correlations increased with increasing buffer lengths, suggesting the importance of near‐stream land use on biotic assemblages. 4. Several metrics changed markedly between 10–20% (mountain ecoregions) and 40–45% (lowland) of arable land use, irrespective of the buffer size. At mountain sites with >10% catchment arable land use, metric values differed between sites with <30% and sites with >30% forest in the near‐stream riparian area. 5. These findings support the role of riparian land use in catchment management; however, differences between mountain and lowland ecoregions support the need for ecoregion‐specific management.     

The relationships between local and regional species richness and spatial turnover

Aim To determine the empirical relationships between species richness and spatial turnover in species composition across spatial scales. These have remained little explored despite the fact that such relationships are fundamental to understanding spatial diversity patterns. Location South‐east Scotland. Methods Defining local species richness simply as the total number of species at a finer resolution than regional species richness and spatial turnover as turnover in species identity between any two or more areas, we determined the empirical relationships between all three, and the influence of spatial scale upon them, using data on breeding bird distributions. We estimated spatial turnover using a measure independent of species richness gradients, a fundamental feature which has been neglected in theoretical studies. Results Local species richness and spatial turnover exhibited a negative relationship, which became stronger as larger neighbourhood sizes were considered in estimating the latter. Spatial turnover and regional species richness did not show any significant relationship, suggesting that spatial species replacement occurs independently of the size of the regional species pool. Local and regional species richness only showed the expected positive relationship when the size of the local scale was relatively large in relation to the regional scale. Conclusions Explanations for the relationships between spatial turnover and local and regional species richness can be found in the spatial patterns of species commonality, gain and loss between areas.     

The role of trout in stream food webs: integrating evidence from field surveys and experiments

1. We evaluated the effects of brown trout on boreal stream food webs using field surveys and enclosure/exclosure experiments. Experimental results were related to prey preference of uncaged trout in the same stream, as well as to a survey of macroinvertebrate densities in streams with vs. without trout. Finally, we assessed the generality of our findings by examining salmonid predation on three groups of macroinvertebrate prey (chironomid midges, epibenthic grazers, invertebrate predators) in a meta-analysis. 2. In a preliminary experiment, invertebrate predators showed a strong negative response to trout, whereas chironomids benefited from trout presence. In the main experiment, trout impact increased with prey size. Trout had the strongest effect on invertebrate predators and cased caddis larvae, whereas Baetis mayfly and chironomid larvae were unaffected. Trout impact on the largest prey seemed mainly consumptive, because prey emigration rates were low and independent of fish presence. Despite strong effects on macroinvertebrates, trout did not induce a trophic cascade on periphyton. Uncaged trout showed a strong preference for the largest prey items (predatory invertebrates and aerial prey), whereas Baetis mayflies and chironomids were avoided by trout. 3. Densities of invertebrate predators were significantly higher in troutless streams. Baetis mayflies also were less abundant in trout streams, whereas densities of chironomids were positively, although non-significantly, related to trout presence. Meta-analysis showed a strong negative impact of trout on invertebrate predators, a negative but variable impact on mobile grazers (mainly mayfly larvae) and a slightly positive impact on chironomid larvae. 4. Being size-selective predators, salmonid fishes have a strong impact on the largest prey types available, and this effect spans several domains of scale. Discrepancies between our experimental findings and those from the field survey and meta-analysis show, however, that for most lotic prey, small-scale experiments do not reflect fish impact reliably at stream-wide scales. 5. Our findings suggest that small-scale experiments will be useful only if the experimental results are evaluated carefully against natural history information about the experimental system and interacting species across a wide array of spatial scales.     

Persistence and stability of macroinvertebrate communities in streams of Denali National Park, Alaska: implications for biological monitoring

1. Macroinvertebrate communities were studied from 1994 to 2001/2002 (except 1997) in six streams in Denali National Park, interior Alaska. All six streams were potential reference streams with no known impairment. 2. Abundance of individual taxa varied markedly from year to year. Overall, abundance decreased over the study period, particularly with respect to mayflies. Stonefly taxa showed lower persistence and were sometimes absent from a stream in any particular year. 3. Mean community persistence for the six streams, as measured by Jaccard's similarity coefficients between years, varied from 0.48 in the year pair 1999–2000 to 0.78 in 1998–99. Tattler Creek (a small stable stream) supported the most persistent macroinvertebrate community and Highway Pass Creek (a small, unstable creek) the least. Mean community persistence showed a significant relationship with mean winter snowfall (November to March) for the six streams. 4. The highest community compositional stability was found in Tattler Creek and the lowest in Highway Pass Creek, but stability varied markedly over time for the six streams, peaking in 1994–95 and reaching a minimum in 2000–01. Compositional stability was significantly related to the Pfankuch Index of channel stability. 5. The composition metrics % Chironomidae, % dominant taxa, %EPT, % Ephemeroptera and % Plecoptera, employed as part of the Alaska Stream Condition Index, varied over almost their entire range in these pristine streams across the 9 years of the study. 6. This study demonstrates the wide range of natural variation that occurs in benthic macroinvertebrate communities in these pristine central Alaskan streams, potentially limiting the applicability of composition metrics for the biological monitoring of water quality in these systems.     

Land-use influence on stream water quality and diatom communities in Victoria, Australia: a response to secondary salinization

Diatom communities were analyzed in 39 streams located in drainages with varied land-use practices throughout Victoria, Australia. Thirteen water quality parameters were also measured in each stream. Most streams had low HCO₃ ^1- concentrations (low buffering capacity) with >90% of the waters dominated by Na¹⁺ and Cl^1-. Phosphate concentrations ranged from 0.003 to 2.0 mg/L. Diatom communities (245 taxa) were strongly correlated with land-use practices, i.e. historic clear cutting, and secondary salinization. Streams influenced by heavy irrigation practices and dryland farming had reduced species diversity and richness compared to systems with low to moderate land use. A nonmetric multidimensional ordination of diatom communities in the 39 streams was conducted. An ANOSIM on the ordination showed that diatom communities in upland watersheds with native forest canopies and low salinization, lowland streams in watersheds with cleared forest canopies, moderate agricultural utilization and salinization, and lowland streams in areas with high irrigation and salinization were all significantly different (p<0.001) from one another. Community ordination techniques showed that both specific conductance (salinity) and phosphorus interacted to determine stream diatom community structure in drainages with high secondary salinization. Drainages with low to moderate agricultural activity and low nutrients, but with a wide range of salinities showed strong associations with the diatom taxa Amphora coffeaeformis, Cymbella pusilla and Entomoneis paludosa, whereas, streams in regions with heavy agricultural practices and high phosphorus had Bacillaria paradoxa, Nitzschia hungarica, N. frustulum and Aulacoseira granulata as numerically important diatoms. In contrast, Rhizosolenia eriensis, Frustulia rhomboides, Eunotia pectinatus and Tabellaria flocculosa were strongly associated with upland streams with fast current, relatively low O-PO₄ ^3- concentrations, low pH, low salinity, and low temperature. In general, the diatom communities in saline streams (3 mS) were similar to those previously reported in saline lakes in Victoria.     

Identifying the scales of variability in stream macroinvertebrate abundance, functional composition and assemblage structure

1. Many natural ecosystems are heterogeneous at scales ranging from microhabitats to landscapes. Running waters are no exception in this regard, and their environmental heterogeneity is reflected in the distribution and abundance of stream organisms across multiple spatial scales. 2. We studied patchiness in benthic macroinvertebrate abundance and functional feeding group (FFG) composition at three spatial scales in a boreal river system. Our sampling design incorporated a set of fully nested scales, with three tributaries, two stream sections (orders) within each tributary, three riffles within each section and ten benthic samples in each riffle. 3. According to nested anova s, most of the variation in total macroinvertebrate abundance, abundances of FFGs, and number of taxa was accounted for by the among‐riffle and among‐sample scales. Such small‐scale variability reflected similar patterns of variation in in‐stream variables (moss cover, particle size, current velocity and depth). Scraper abundance, however, varied most at the scale of stream sections, probably mirroring variation in canopy cover. 4. Tributaries and stream sections within tributaries differed significantly in the structure and FFG composition of the macroinvertebrate assemblages. Furthermore, riffles in headwater (second order) sections were more variable than those in higher order (third order) sections. 5. Stream biomonitoring programs should consider this kind of scale‐dependent variability in assemblage characteristics because: (i) small‐scale variability in abundance suggests that a few replicate samples are not enough to capture macroinvertebrate assemblage variability present at a site, and (ii) riffles from the same stream may support widely differing benthic assemblages.     

Gradients in stream fish assemblages across a Mediterranean landscape: contributions of environmental factors and spatial structure

1. Despite wide recognition that fish assemblages are influenced by factors operating over a range of spatial scales, little effort has been devoted to quantifying large‐scale variation and the multiscale dependencies of assemblage patterns and processes. This is particularly true for Mediterranean streams, where seasonally predictable drying‐up may lead to a strong association between assemblage attributes and large‐scale factors affecting the distribution of population sources and extinction likelihood. 2. The contribution of large‐scale factors to stream fish assemblage variation was quantified across a Mediterranean landscape, in south‐west Portugal. Fish abundance and species composition were estimated at 166 sites across third‐ to sixth‐order streams, in March–July 1998. Variance partitioning by redundancy analyses was used to analyse assemblage variation against three sets of predictor variables: environmental (catchment position, and geomorphic and hydrological factors), large‐scale spatial trends and neighbourhood effects. 3. Environmental variables and spatial trends accounted for 34.6% of the assemblage variation across the entire region, and for 36.6 and 57.8% within the two largest catchments (Mira and Seixe). Neighbourhood effects were analysed at the catchment scale, increasing the explained variation to 56.1% (Mira) and 70.7% (Seixe). 4. A prevailing environmental gradient was reflected in an increase in the abundance of all species and size‐classes in relation to catchment position, with more fish present in larger streams and in downstream reaches. Variables describing geomorphic and hydrological settings were less important in explaining assemblage variation. 5. Spatial trends always accounted for the smallest fraction of assemblage variation, and they were probably associated with historical barriers to fish dispersal. The strong neighbourhood effects may be related to spatially autocorrelated habitat conditions, but they are also a likely consequence of fish emigration/extinction and colonisation processes. 6. These results emphasise that a substantial proportion of fish assemblage variation in Mediterranean streams may be explained by large‐scale factors, irrespective of microhabitats and local biotic interactions. It is suggested that this pattern results to a large extent from the seasonal drying‐up, with the summer shortage of surface water limiting fish occurrence in headwaters, and consequently the key core areas for fish concentrating in larger streams and tributaries adjacent to large streams because of neighbourhood effects.     

The effect of spatial scale on interactions between two weevils and their parasitoid

1. The effect of spatial scale on the interactions between three hymenopteran parasitoids and their weevil hosts was investigated. The parasitoid Mesopolobus incultus (Walker) parasitised Gymnetron pascuorum Gyll.; the parasitoids Entodon sparetus (Walker) and Bracon sp. parasitised Mecinus pyraster Herbst. Both of these weevils develop inside the seedhead of Plantago lanceolata L. but occupy different niches. Seedheads were sampled annually from 162 plants at each of two experimental sites consisting of a series of habitat patches of two distinct sizes. Data were analysed from three site‐years. 2. Parasitoid densities at each site‐year were closely related to the abundance of their respective weevil hosts. The overall proportion of hosts parasitised was more variable for M. incultus than for E. sparetus and Bracon sp. 3. Changes in spatial scale affected the variability of parasitoid densities. For M. incultus, there was generally a greater degree of additional heterogeneity for all increases of scale; for E. sparetus, this was true only at the largest scales; for Bracon sp., all components of variance were negative. 4. The rate of parasitism was related to host density in different ways at different spatial scales. Mesopolobus incultus exhibited inverse density dependence at the finest (seedhead) scale, direct density dependence at the intermediate (plant) scale, and density independence at the large (habitat area 729 m²) scale. Entodon sparetus showed no response to variation in host density at any spatial scale. Bracon sp. showed direct density dependence only at the intermediate and largest scales. 5. Parasitoids E. sparetus and Bracon sp. seemed able to detect more than one M. pyraster individual in seedheads with multiple host occupancy; a greater incidence of conspecific parasitoids than expected emerged from such seedheads.