Spatial and temporal variation in species composition of dung beetle assemblages in southern Ireland

Abstract. 1. This study attempts to identify the main community characteristics that contribute to variability in dung beetle assemblage composition and structure across a range of spatial and temporal scales.
2. Dung beetle assemblages ( Aphodius , Sphaeridium, and Geotrupes species) were monitored by dung-baited pitfall trapping at 10-day intervals during the seasonally active period at eleven sites in southern Ireland. Three of the sites were monitored over at least 2 years between 1991 and 1996.
3. Although the species composition of the above taxonomic groups was comparable among sites and years, relative abundances of component species varied considerably. Detrended correspondence analysis ordinations indicated a similar level of variability in dung beetle assemblage structure among years, and among sites ≈1–180 km apart.
4. Processes that may contribute to spatio-temporal variability in dung beetle assemblages are discussed, and strategies for future research are suggested.     

Local and ecoregion effects on fish assemblage structure in tributaries of the Rio Paraíba do Sul, Brazil

1.  We examined the effects of physical and chemical habitat variables and ecoregions on species occurrence and fish assemblage structure in streams of the Paraíba do Sul basin, in southeast Brazil.
2.  Fish and environmental data were collected from 42 sites on 26 first to fourth order streams (1 : 50 000 map scale) in three ecoregions. The sites occurred in one valley and two plateau ecoregions at altitudes of 40–1080 m and distances of 0.1–188 km from the main channel of the Rio Paraíba do Sul. Physical habitat (substratum, riparian cover, habitat types) and water quality (dissolved oxygen, pH, temperature and conductivity) variables were measured at each sampling site.
3.  A total of 2684 individuals in 16 families and 59 species were recorded.
4.  Ecoregion was a better predictor of the fish assemblage than the other environmental variables, according to the differences between the mean within-class and mean between-class similarities in assemblage data.
5.  Differing landscape characteristics were associated with differing local variables and thereby with differing fish assemblage structures. Riffles, shrub, grass, dissolved oxygen, conductivity and temperature were closely related to fish assemblage structure.
6.  Fish assemblages in sites far from the main river and at higher altitudes also differed from those near the Paraíba do Sul main channel, presumably as a result of differences in connectivity, covarying environmental factors and anthropogenic influence.
7.  These results reinforce the importance of understanding how stream communities are influenced by processes and patterns operating at local and regional scales, which will aid water resource managers to target those factors in their management and rehabilitation efforts.     

A Strategic Interpretation of Beetle (Coleoptera) Assemblages, Biotopes, Habitats and Distribution, and the Conservation Implications

The identification of beetle, and other invertebrate, biotopes based on the recording of species assemblages has become a regular occurrence as a result of increased survey work, with the approach expanded into classifications of grid square pooled species lists at the national scale. A reassessment of beetle biotopes and distribution has been attempted by interpreting the classifications in terms of the major environmental factors of productivity and disturbance, identified as being important drivers in work on habitat templates and strategic triangles. For grassland ground beetle biotopes, productivity was generally related to soil quality whilst disturbance was associated with land management or cover. Productivity of exposed riverine sediment ground beetle biotopes was dependent on deposited organic matter and disturbance on the effects of water flow on site structure. With both ground beetle biotopes, the distribution of assemblages was also affected by substrate water, another abiotic driver. Productivity in aquatic beetle biotopes was a function of base-status, generally reflected by pH, whilst disturbance was mainly due to water flow and wave action. However, disturbance in ditches was also the result of site vegetation management whilst another factor affecting assemblage distribution was water permanence, with temporary water having specific assemblages. For large-scale British, grid-based, classifications productivity and disturbance were interpreted from satellite-derived land cover data. At the national scale, the other important factor influencing distribution was temperature. There is a requirement for better quantification of a number of the factors influencing biotope and assemblage definition. Species strategies employed to cope with the various environmental variables were reflected in traits in such factors as morphology and life-cycles. The effects of environmental perturbations including climate change, pollution and land use are discussed in relation to the environmental variables and to the various species strategies. Biotope classifications, using standardised and reproducible survey methods, allied to a better understanding of the underlying environmental pressures, could produce a unified approach for the conservation of beetle and other invertebrate species.     

Macroinvertebrate distributions and the conservation value of aquatic Coleoptera, Mollusca and Odonata in the ditches of traditionally managed and grazing fen at Wicken Fen, UK

1. Water-filled ditches are an important ecological feature of lowland Britain. Originally dug to facilitate wetland drainage, ditches often provide refuges for aquatic flora and fauna of high conservation value.
2. The ditches, ponds and major waterways of a traditionally managed undrained fen and the ditches of a previously drained cattle-grazed fen meadow were sampled at Wicken Fen National Nature Reserve for Coleoptera, Mollusca and Odonata, and for physical, chemical and biological variables.
3. Multivariate analysis showed a clear division between ditches on the two fens, and between larger and smaller waterbodies.
4. Individual ditches possessed distinctive faunas. Ditch age and bank profile were important factors influencing faunal species composition.
5. Invertebrate distributions were shown to be significantly correlated with macrophyte distribution.
6. Waterbodies were ranked in terms of their species quality, using a point-scoring system; there was no correlation between rankings for the three invertebrate groups; only ditch age and detritus levels were correlated with species quality score (for Coleoptera).
7. Ditch management recommendations are discussed in the light of these results.     

Grouse moor management: habitat and conservation implications for invertebrates in southern Scotland

The distribution of ground beetle, rove beetle, spider and plant bug species assemblages, and the occurrence of other beetle species, was investigated in a five-year survey of a grouse moor in the Scottish borders, some of which was managed by burning, cutting and herbicide application. Multivariate analyses were used to assess the relationship between assemblage distribution and a number of environmental variables. The management of dry Calluna moor had a positive effect on the habitat diversity for ground beetles and plant bugs but had little effect on rove beetles and spiders. A total of 39 nationally rare and scarce species (38 beetle, 1 plant bug) was recorded in the survey. The most important habitat present was on streamside sites, especially sediment. A number of these rarer species were restricted to sites managed by burning and cutting but other species were only found on unmanaged wet Calluna moor. Molinia-dominated moor was generally of poorer quality than Calluna sites, with fewer rare and scarce species and lower site rarity values based on the ground beetle species recorded. The highest median site rarity scores were for dry, open, managed Calluna sites. In order to maximise both habitat diversity and the incidence of rare and scarce species on grouse moor, a mosaic of both managed and undisturbed patches differing in soil characteristics, plant composition and vegetation structure appears to be required.     

The Rove Beetles (Coleoptera, Staphylinidae) of Exposed Riverine Sediments in Scotland and Northern England: Habitat Classification and Conservation Aspects

Exposed riverine sediments (ERS) by four rivers in Scotland and northern England were sampled for beetles in 1996 and 1997. One hundred and sixty rove beetle (Staphylinidae) species lists were analysed using ordination and classification techniques in order to identify habitat groups within and between catchments and to assess which factors were affecting species assemblage distribution. There were major differences between the species assemblages of ERS by rivers of highland and lowland catchments. Within catchments, assemblage distribution was mainly influenced by the position of sites within the catchment; vegetation cover and sediment composition had less influence. The number of rove beetle habitats was not the same as those for ground and phytophagous beetle groups, indicating that conservation considerations should take into account variations in ERS habitat diversity. A considerable number of records of nationally rare and scarce rove beetle species were recorded, most on ERS by rivers and tributaries unaffected by river management or engineering.     

Comparing the effects of farming practices on ground beetle (Coleoptera: Carabidae) and spider (Araneae) assemblages of Scottish farmland

Multivariate techniques were used to compare and contrast the effects of land cover and farming practice on ground beetle and spider assemblages of Scottish farmland. For both ground beetles and spiders, the ordination and fuzzy clustering of sites were related to land cover rather than geographical location or year of sampling. The same four types of land cover group were identified: that is, heather moorland, semi-natural grassland, intensive grassland and arable land. The robustness of these land cover groups was tested using previously unsampled sites and it was found that 79 and 86% of sites, for ground beetle and spider assemblages respectively, were allocated to the land cover group predicted from their actual land cover. Furthermore, procrustes rotational analysis found a strong relationship between ground beetle and spider assemblages in intensively managed sites, suggesting that the assemblage structure of one group could be used to predict that of the other. The observed relationship between spider and ground beetle assemblages does not necessarily indicate that both groups were responding to agricultural practices in the same way. Indeed, the highest number of beetle species occurred in intensively managed grassland and arable sites while the highest number of spider species occurred in semi-natural grassland and heather sites. When conducting ecological assessments, one might wish to collect information on a wide range of ecologically different taxa; however, financial constraints make this unfeasible. From the results it could be concluded that spiders should be chosen in preference to ground beetles when seeking to make predictions on how farming practices influence invertebrates. However, such a conclusion would be premature since not only were spiders more numerous in the traps, but they were also more time consuming to process. In addition, the strong relationship found between the spider and ground beetle assemblages further justifies carabids as a target group when monitoring the influence of farming practices on biodiversity.     

Assignment and evaluation of ground beetle (Coleoptera: Carabidae) assemblages to sites on different scales in a grassland landscape

Carabid beetles and environmental parameters were investigated in 52 grassland sites with three replicate pitfall traps in each site and in the valley of the River Eider in Schleswig–Holstein (northern Germany) with 61 pitfall traps. Environmental parameters included water content of soil, sand content, organic matter content and pH. Ground beetle assemblages were derived by detrended correspondence analyses (DCA) and characterised by the specific environmental conditions as means for each assemblage. On the regional scale including all investigated sites of Schleswig–Holstein, five assemblages were differentiated. On the local scale including the investigated sites in the valley of the River Eider, three assemblages were found corresponding well with those found on the regional scale. Environmental conditions at the sites of the five assemblages were correlated with land use data, soil types, and water level stages provided by three maps of a geographic information system (GIS). The GIS maps were combined to develop smaller areas with land use, soil type and water level stage information. The characteristic environmental conditions were assigned to each area to derive the spatial distribution of the five ground beetle assemblages. Spatial prediction was correct for 65% of investigated sites. The potential area of each assemblage was estimated for the valley. The different grassland areas were evaluated as potential habitats for ground beetle species comparing total species richness with the regional species richness of each assemblage. The comparison shows that species richness in the evaluated assemblages is relatively low compared to the regional potential.     

Asymmetrical competition between Neotropical dung beetles and its consequences for assemblage structure

Abstract.  1. This study combines the results of laboratory experiments using representative assemblage components and pitfall trapping over a large geographical area to examine the hypothesis that ongoing interspecific competition structures Neotropical dung beetle assemblages.
2. From Guatemala to Panama assemblages of large to medium-sized, fast-tunnelling dung beetles include a single large, nocturnal dichotomiine species, Dichotomius annae (Kohlmann & Solís, 1997). In competition experiments, this species out-competed the medium-sized coprine species, Copris lugubris Boheman and Phanaeus demon Laporte-Castelnau, for dung and nesting space, in spite of earlier colonisation by the diurnal species, P. demon .
3. Differences in the abundance of D. annae at Central American sites did not affect total fast-tunnelling dung beetle assemblage richness over the rainy season. However, D. annae rank order was directly related to the probability of interspecific encounters (Hurlbert's Δ ₁) among species. These trends were also observed when species lists from published and unpublished studies of other large allopatric dichotomiine species, with a more northerly distribution, were included in the analyses.
4. The results obtained suggest that where large dichotomiine species are abundant, their efficient pre-emption of a considerable proportion of available resources drives all, or most, other fast-tunnelling species to a lower population density, thereby decreasing assemblage diversity.     

Changes in structure and composition of an assemblage of Hydroporus species (Coleoptera: Dytiscidae) along a pH gradient

SUMMARY. 1. Patterns of total abundance and species relative abundances in an assemblage of predatory dytiscid beetles of the genus Hydroporus were examined along a pH gradient within a ditch in north-cast England.
2. During spring, total abundance of adult Hydroporus was greatest at the less acidic (pH 5.6–6.2) upstream sites, whereas total abundance of larval Hydroporus was greatest at the more acidic (pH 4.5) downstream sites. Both adults and larvae were more abundant in samples from the edge than from the middle of the ditch.
3. Species number and evenness did not differ along the pH gradient.
4. Species relative abundance patterns varied along the pH gradient, with particular species dominating at different sites. However, relative abundances of species changed discontinuously along the pH gradient.
5. Comparison of species abundance patterns along this pH gradient with published patterns of presence/absence among bodies of water with different pH revealed that species relationships with pH are often not consistent. Possible sources of this inconsistency are: geographic variation in species' pH preferences; complex responses of individual species to many factors in addition to pH; or random variation among sites due to colonization and extinction history.
6. The overall results suggest that pH is not the only, or even the most important, factor influencing Hydroporus populations and assemblage organization.     

Application of an index of biological integrity (IBI) to fish assemblages of the rivers Khan and Kshipra (Madhya Pradesh), India

1. A multimetric index of fish assemblage integrity was developed and similarity analyses were conducted on fish species in two central Indian rivers and the effects of distance from municipal and industrial effluents on those indices then evaluated.
2. Five metrics from Karr et al . (1986 , Illinois Natural History Survey Special Publication 5 , Urbana, IL) were adopted: intolerant species richness, % omnivorous individuals, % top carnivore individuals, total number of individuals and % individuals with anomalies. Seven new metrics (native species richness, native family richness, benthic species richness, water column species richness, % non-native individuals, % tolerant individuals and % herbivorous individuals) were added.
3. Non-native individuals represented 1–55% of the assemblages at sampled sites which held fish.
4. Fish were present at eleven sites and not collected at two sites, despite heavy metal concentrations exceeding U.S. Environmental Protection Agency acute criteria at all sites.
5. Two types of metric scoring were examined. The traditional 5–3–1 method showed the same pattern as continuous scoring from 0 to 10, but produced a higher integrity class at one site.
6. Scores on our modified index of fish assemblage integrity increased with distance downstream from a major effluent source in each river. Jaccard similarity scores between the least disturbed downstream site and all other sites decreased with increasing distance and disturbance.
7. It was concluded that Karr's original index and its theoretical foundations are easily adaptable, even to an ichthyofauna containing no species, and only two families (Cyprinidae, Poeciliidae), in common with the midwestern United States.     

The diversity of beetle assemblages in different habitat types in Sabah, Malaysia

The diversity of beetle assemblages in different habitat types (primary forest, logged forest, acacia plantation and oil palm plantation) in Sabah, Malaysia was investigated using three different methods based on habitat levels (Winkler sampling, flight-interception-trapping and mist-blowing). The overall diversity was extremely high, with 1711 species recorded from only 8028 individuals and 81 families (115 family and subfamily groups). Different degrees of environmental changes had varying effects on the beetle species richness and abundance, with oil palm plantation assemblage being most severely affected, followed by acacia plantation and then logged forest. A few species became numerically dominant in the oil palm plantation. In terms of beetle species composition, the acacia fauna showed much similarity with the logged forest fauna, and the oil palm fauna was very different from the rest. The effects of environmental variables (number of plant species, sapling and tree densities, amount of leaf litter, ground cover, canopy cover, soil pH and compaction) on the beetle assemblage were also investigated. Leaf litter correlated with species richness, abundance and composition of subterranean beetles. Plant species richness, tree and sapling densities correlated with species richness, abundance and composition of understorey beetles while ground cover correlated only with the species richness and abundance of these beetles. Canopy cover correlated only with arboreal beetles. In trophic structure, predators represented more than 40% of the species and individuals. Environmental changes affected the trophic structure with proportionally more herbivores (abundance) but fewer predators (species richness and abundance) in the oil palm plantation. Biodiversity, conservation and practical aspects of pest management were also highlighted in this study.     

Beetle assemblages in ponds: effects of habitat and site age

• 1 Water beetle assemblages were sampled in each of 18 freshwater ponds, including 11 recently constructed sites designed to provide mitigation for wetlands destruction elsewhere, and seven older reference sites. There were three objectives: (a) to relate taxon richness and biomass of the beetles to the same properties of the wider aquatic invertebrate community, (b) to evaluate changes in beetle assemblage structure over time, and (c) to determine habitat effects on taxonomic composition, mean body size and trophic guild structure of the beetle assemblage.
• 2 Forty‐seven beetle genera were identified, representing 77 species. The beetles represented an average of 21.5% of total generic richness, but only 3.7% of total wet biomass of the wider invertebrate community.
• 3 Of all variables evaluated using canonical correspondence analysis (CCA), site age had the greatest influence on the beetle assemblage. Predatory dytiscids were early colonists at younger sites, while herbivorous curculionids and chrysomelids associated with particular types of vegetation typically occurred in older ponds. Mitigation ponds and reference ponds supported similar numbers of species. Reference sites, however, harboured substantially more unique species found at only a single site within the study area.
• 4 The presence of fish was also strongly related to beetle assemblage structure. Ponds with few or no fish contained about 3‐fold higher biomass and 3‐fold greater mean wet weight per individual compared to ponds with substantial fish assemblages.
• 5 Beetle assemblage composition varied among sites and sampling years, but beetle biomass, richness and species composition may be useful tools in evaluating the success of wetland mitigation efforts.

     

Detecting the effects of environmental change above the species level with beetles in a fragmented tropical rainforest landscape

Abstract.  1. Invertebrate data identified to coarse groupings according to taxonomy or ecology are easier to obtain than species-level data. However, it is unclear if such data are suitable for detecting environmental change.
2. We compared the performance of four types of data (species, family, trophic group, and body size) to quantify the responses of beetle assemblages with three types of environmental change on the Atherton Tableland in north-eastern Queensland, Australia. Clearing of rainforest creates two levels of environmental change: first the extreme, forest loss, and second, the more subtle change associated with forest fragmentation. A strong rainfall gradient across the study landscape also influences the biota, imposing a third type of environmental change, independent of rainforest modification.
3. Almost 20 000 beetles were sampled from six replicate sites in each of pasture, small rainforest remnants, and both the edges and interiors of large rainforest remnants.
4. All four types of data showed significant multivariate differences in assemblage composition between pasture and rainforest sites. Species-level data in multivariate analyses showed an effect of fragmentation on small remnants and also showed variation which corresponded with the spatial aridity gradient across sites. Both of these patterns were only weakly evident or were non-existent at the level of family, trophic group, or body size class. No level of taxonomic resolution gave consistently stronger results when univariate tests were applied to individual component taxa.
5. We conclude that data above the species level may not have the sensitivity required to detect more subtle forms of environmental change.     

Effects of fen management and habitat parameters on staphylinid beetle (Coleoptera: Staphylinidae) assemblages in north-eastern Germany

Because fens have undergone dramatic declines in recent decades, an important question is which management regimes and habitat parameters are most effective in preserving fen biodiversity. The aim of the present study was to assess the effects of five different management regimes (intensive grassland, moist meadows, summer harvested sites, winter harvested sites, fallows) on staphylinid beetle assemblages. During the study period 5,989 individuals from 92 staphylinid beetle species were recorded. Species richness and abundance were highest on intensive grassland and fallows, and water level and vegetation height had significant impacts on the abundance of staphylinids. On winter-harvested sites species richness and abundance were lowest. In general, staphylinid beetles did not show pronounced variation among management regimes, while the environmental factors water level, vegetation height and top soil mineralisation seemed to have a larger structuring impact. The number of threatened species and the conservation index were highest on summer-harvested sites and fallows, representing fairly well-preserved fens. We conclude that summer harvest of reed beds or no management at all appears to be most beneficial for the conservation of staphylinid beetles.     

Invasion and retreat: shifting assemblages of dung beetles amidst changing agricultural landscapes in central Peru

This study examines the effects of deforestation, and the habitat value of coffee and regenerated forest for tropical dung beetles, a functionally significant insect group. Pitfall trapping was conducted at 22 sites in a montane region of central Peru during April and November/December of 2002. Sites included primary and secondary forest, shade-coffee, regenerated forest and open farms (mainly with banana, yuca, and corn). Ordination techniques indicated that beetle assemblages in forests, regenerated forest and coffee were relatively similar. However, assemblage compositions in forested areas differed even at similar altitudes under the influence of biogeographical factors, and the assemblages at disturbed sites (farms/coffee) were influenced by beetle dispersal from adjacent forests. During dry months, when beetle activity is low, communities at all habitat types tended to converge because fewer unique species were recorded in forests at that time and habitat/season generalists were dominant. Preliminary results also indicate that beetles in shady crops such as bananas responded to plant growth: as the banana canopy closed-in, producing more shade, open-habitat specialists retreated and forest/shade specialists invaded the sites. Chronosequence data at two of the sites demonstrate the rapid and dramatic changes in species richness and assemblage composition caused by deforestation. As forests become increasingly fragmented, and open farms continue to expand, dung beetles will become more restricted to the remaining fragments and reserves. In the mosaic landscape studied here, shade crops, like coffee, act as habitat and corridors for many dung beetle species. Small farm size and the consequent magnitude of edge effects, likely contributed to beetle movement between habitat types and determined the apparent generalist nature of many of the dung beetle species in this study.     

Livestock Trampling Reduces the Conservation Value of Beetle Communities on High Quality Exposed Riverine Sediments

Exposed riverine sediments (ERS) are habitat for a large number of rare and threatened specialist species of invertebrates and are of considerable conservation importance. Livestock trampling is believed to be one of the most widespread causes of damage to ERS habitats in the UK, and as such, its effects were the focus of this investigation. Beetle density was measured at two points within 25 distinct patches of habitat along ∼ ∼47 km of the Afon Tywi special site of scientific interest in South Wales, which is known to support an extremely good quality ERS beetle fauna in a UK context. Partial canonical correspondence analyses were used to explore the relationship between beetle assemblage and a range of environmental variables. The percentage of fine (<8 mm) sediments, median sediment size, distance downstream, cattle stocking levels, and counts of sheep faeces were found to best relate to beetle abundance and assemblage structure. Species richness was positively associated with stocking levels, probably because of the addition of species associated with resultant elevated levels of silt and organic matter. The ERS quality score, which is a measure of conservation value based on the rarity of specialist ERS beetles, was negatively associated with measures of trampling damage. It was therefore concluded that livestock trampling reduces the conservation value of beetle communities on high quality ERS and management should restrict trampling in sites of high conservation importance.     

Survey of the bryophytes of arable land in Britain and Ireland 1: a classification of arable field assemblages

Abstract

Knowledge of the distribution of arable bryophytes lags behind that of bryophytes of many other habitats, and we have almost no information on their occurrence in relation to crop type and agricultural management regime. A survey of arable fields in Britain and Ireland carried out by members of the British Bryological Society in 2001–05 was designed to provide such baseline data. We surveyed a stratified random sample of 200 fields in the main areas of arable agriculture and a further 620 fields in these areas and in areas where arable fields are less frequent. The species present in each field were listed and their frequency within the field assessed. Data on the crop, the cover of vascular plants, bryophytes, trash and bare soil, and the pH and texture of the soil were recorded for each field. Six species assemblages are described on the basis of a classification of the species recorded in each field and their frequency values. The Tortula truncata–Anthoceros and Dicranella staphylina–Riccia glauca assemblages are the most species-rich and are concentrated on acidic soils in northern and western Britain and in Ireland. By contrast, the Barbula unguiculata–Bryum klinggraeffii and Phascum cuspidatum–Microbryum davallianum assemblages are mainly found on calcareous soils in southern and eastern England. The Bryum dichotomum–Marchantia polymorpha assemblage is characterised by widespread, weedy generalist species and is concentrated in East Anglia. The final assemblage, Brachythecium rutabulum–Fissidens taxifolius, characteristically occurs in late-successional fields and was scattered throughout the area sampled. A comparison of the results from Kent with those of a survey of Kentish fields (using a different methodology) by A.G. Side in 1973–74 suggests that the Bryum dichotomum–Marchantia polymorpha assemblage may possibly have replaced the Dicranella staphylina–Riccia glauca assemblage on acidic soils in the last 30 years. The considerable geographical variation within the arable flora of Britain and Ireland revealed by the survey should be taken into account when assessing the meagre evidence for historic changes in the arable bryophyte flora.     

Seasonal fluctuation, phenology and turnover of chafer assemblages – insight to the structural plasticity of insect communities in tropical farmlands

1 Studies on chafer assemblages were conducted on two farmland sites in the Terai lowland of Nepal (200 m above sea level) using light traps. During the course of a 2-year field monitoring program, a total of 4503 specimens was captured and an unexpectedly high number of syntopically co-occurring species was found: 52 from Gunganagar (GN) and 36 from Gaindakot (GK), respectively. Highest species abundances and species numbers were found during April and May.
2 Species occurrence was strongly correlated with air temperature and the maximum soil temperature, at least during the pre-monsoon season. However, assemblage structure from the two sites showed significant qualitative and quantitative changes seasonally, as well as from 1 year to the next. Turnover rates between adjacent months were in the range 26–62% (GN) and 37–70% (GK), whereas the turnover from 2004 to 2005 was 25.8% (GN) and 21.4% (GK) respectively.
3 When only dominant and subdominant taxa are considered, the seasonal change in species composition was even more striking.
4 Strong fluctuation in chafer assemblage over time suggests: (i) a possible influence of patchy habitat types and soil working on seasonal assemblage structure and (ii) colonization of suitable habitats (fields) in great part by chance.