Occurrence patterns of butterflies (Rhopalocera) in semi-natural pastures in southeastern Sweden

Occurrence patterns of butterflies (Rhopalocera) were analysed in relation to locality characteristics, and nested patterns were evaluated, based on a survey of 100 semi-natural pastures in the county of Östergötland, southeastern Sweden. Species richness of butterflies was positively related with locality-size, but not with the density of pastures in the surroundings. Species richness was lower in heavily grazed pastures compared with localities with a low or intermediate grazing pressure. This suggests that a uniformed, heavy grazing pressure should be avoided if butterflies are to be protected, even though such a management is favourable for many vascular plants. Out of 45 analysed species, 73% exhibited a significantly nested pattern, and species regarded to be sedentary had generally a more nested pattern. Several butterflies (such as Erynnis tages, Mellicta athalia and Pyrgus malvae) are possible to use as indicators of a relatively intact butterfly community. In a short-term perspective, the great bulk of butterflies confined to semi-natural pastures would be saved if management were adapted to the requirements of butterflies only at the localities richest in species. However, for conservation strategies to be successful over a longer time, whole landscapes harbouring the majority of the species and with a high density of semi-natural pastures should be selected and be given priority for conservation.     

Evaluation of community indices in seasonal assemblages of butterflies (Lepidoptera) at different frequency of transect count

Community indices were evaluated at different frequency of transect counts of butterflies by randomization. Randomized samples were obtained by re-arranging 1, 3, 6, 12 and 18 counts from 18 counts at a study site (Hiraoka), and 1, 2, 3, 5 and 15 from 15 at another site (Narukawa), in 1994. Morisita's new estimate of species richness, QM gave a good estimate when the number of counts was five or more. QM estimated from the sum of both assemblages (58 species) was 70.7, while 73 species were recorded at least once from 1990 to 1994. The Octave method always resulted in an underestimation. Diversity indices, Shannon's H, Simpson's D and , Hurlbert's E(Sn), Itow's b, and Morisita's H gave good estimates when the number of counts was five or more. Temporariness of univoltine species in flight period made a bias in the estimates. We concluded that the count is needed at least six times (once a month) and the estimates were improved by separating the multivoltine species from the univoltine species. Intensive counting during the flight period of the univoltine species was recommended.     

Effects of restoration with cattle grazing on plant species composition and richness of semi-natural grasslands

Restoration of semi-natural grasslands by cattle grazing is among the most practical options for reversing the decline of northern European floristic diversity, but no studies on this subject are available. In this work the success of restoration of abandoned, privately owned mesic semi-natural grasslands by farmers receiving support from the EU agri-environmental support scheme was studied in southwestern Finland. Three kinds of grasslands were compared: old (continuously cattle grazed), new (cattle grazing restarted 3–8 years ago) and abandoned pastures (grazing terminated >10 years ago). Plant species composition of the three pasture types was floristically different in multivariate analyses (non-metric multidimensional scaling). Total species richness, richness of grassland plants, indicator plants and rare plants were highest in old and lowest in abandoned pastures in all studied spatial scales (0.25–0.8 ha, 1 and 0.01 m²). The results were congruent with different scales and species list definitions, suggesting that species density scale (1 m²) can be used as a partial surrogate for large scale species richness. Species richness of new pastures was 20% higher on 0.25–0.8 ha, 40–50% higher on 1 m² and 30% higher on the 0.01 m² scale compared to abandoned grasslands. Rare species showed insignificant response to resumed grazing. Despite problems in management quality, this study showed promising results of restoration of abandoned grasslands by cattle grazing on private farms. However, populations of several rare grassland plants may not recover with present cattle grazing regimes. Management regulations in the agri-environmental support scheme need to be defined more precisely for successful restoration.     

Landscape composition and habitat area affects butterfly species richness in semi-natural grasslands

During the last 50 years, the distribution and abundance of many European butterfly species associated with semi-natural grasslands have declined. This may be the result of deteriorating habitat quality, but habitat loss, resulting in decreasing area and increasing isolation of remaining habitat, is also predicted to result in reduced species richness. To investigate the effects of habitat loss on species richness, we surveyed butterflies in semi-natural grasslands of similar quality and structure, but situated in landscapes of different habitat composition. Using spatially explicit habitat data, we selected one large (6–10 ha) and one small (0.5–2 ha) grassland site (pasture) in each of 24 non-overlapping 28.2 km² landscapes belonging to three categories differing in the proportion of the area that consisted of semi-natural grasslands. After controlling for local habitat quality, species richness was higher in grassland sites situated in landscapes consisting of a high proportion of grasslands. Species richness was also higher in larger grassland sites, and this effect was more pronounced for sedentary than for mobile species. However, the number of species for a given area did not differ between large and small grasslands. There was also a significant relationship between butterfly species richness and habitat quality in the form of vegetation height and abundance of flowers. In contrast, butterfly density was not related to landscape composition or grassland size. When species respond differently to habitat area or landscape composition this leads to effects on community structure, and nestedness analysis showed that depauperate communities were subsets of richer ones. Both grassland area and landscape composition may have contributed to this pattern, implying that small habitat fragments and landscapes with low proportions of habitat are both likely to mainly contain common generalist species. Based on these results, conservation efforts should aim at preserving landscapes with high proportions of the focal habitat.     

Low congruence between the diversity of Waxcap (Hygrocybe spp.) fungi and vascular plants in semi-natural grasslands

The diversity of fungi in semi-natural grasslands is poorly known, partly due to difficulties in species identification in the field but also because there are few specialists available. Diversity assessments of grassland fungi would be facilitated if a potential surrogate group for fungal diversity could be identified. The aim of this study was to assess whether plant diversity patterns in semi-natural grasslands are congruent with diversity patterns of Waxcap (Hygrocybe spp.) fungi. Waxcaps, together with several other groups of fungi (e.g. the genera Entoloma, Dermoloma, Camarophyllopsis, and the families Clavariaceae and Geoglossaceae) have habitat requirements similar to many plants typical for semi-natural grasslands and they are all threatened by ceased management and nutrient enrichment. Diversity data from 31 semi-natural grasslands in southern Sweden were used to examine if there is congruence in species richness, nestedness, β-diversity and species composition between plants and Hygrocybe fungi. Species richness of Hygrocybe was significantly positively correlated with plant richness, although the relationship was not strong (r²=0.14). Both plant and Hygrocybe species composition was significantly nested, i.e. species-poor sites contain a subset of species from species-rich sites, which suggests that rare species mostly occur in the species-rich sites. A species similarity analysis between the grassland sites showed that there is low overlap between species composition of plants and Hygrocybe, indicating that conservation decisions based solely on plants may not fulfil the requirements of the Hygrocybe species. The conclusion is that there is low congruence between plant and Hygrocybe species diversity, making plants a poor surrogate group for Hygrocybe fungi, and probably also for other grassland fungi.     

Grassland versus non-grassland bird abundance and diversity in managed grasslands: local,landscape and regional scale effects

Declines of West European farmland birds have been associated with intensive agricultural practices, while in Central and Eastern European countries grasslands still harbour a diverse and unique bird community. However, in these countries comparative studies on the effects of agricultural intensity on biodiversity are virtually missing. We compared bird communities of paired extensively and intensively grazed cattle pastures in three different regions of the Hungarian Great Plain. The influence of grazing intensity, landscape and regional effects were tested on the abundance and species richness of two ecological groups of bird species (grassland and non-grassland birds), as well as on the abundance of the three commonest grassland bird species (Skylark, Yellow wagtail, Corn bunting) in linear mixed models. We found significant effects of grazing intensity on the abundance of grassland birds, which were more abundant on the extensive sites, whereas no effects were found on non-grassland birds. This could be explained by a closer dependence of grassland birds on grasslands for nesting and foraging, whereas non-grassland birds only used grasslands opportunistically for foraging. Landscape effect was shown on grassland bird abundance, but not on non-grassland birds. The regions did affect only the species richness of grassland birds. At species level, the effect of management was significant for the three commonest grassland species, which were more abundant on the extensive fields in all regions. Additionally, on Skylark abundance landscape and regional effects were also shown. These findings suggest that conservation of biodiversity in agricultural systems requires the consideration of landscape perspective to apply the most adequate management.     

Vegetation composition of Lolium perenne-dominated grasslands under organic and conventional farming

Agricultural intensification has caused a decline of semi-natural grasslands and loss of botanical diversity, making agricultural fields dominated by Lolium perenne the main grassland system in large areas of Europe. Increased insight into the factors determining their vegetation composition and plant species richness is needed to improve the effectiveness of agri-environment schemes and to substantiate the benefits of organic over conventional agriculture. Our aims were (1) to determine the difference in vegetation composition (including species richness) between Lolium perenne-dominated fields of conventional and organic farms in a case study region in The Netherlands, and (2) to identify the soil and management related drivers behind this difference.We collected vegetation, soil and management data in grasslands of dairy farms under conventional or organic management (45 fields in total), and used multivariate statistics to determine the effect of fertilisation rates, grazing and cutting regime, and soil properties on plant species composition. In a next step we determined to what extent these abiotic drivers differ between organic and conventional farms.On average the organic fields appeared to have a c. 30% higher plant species richness compared to the conventional ones. Vegetation composition was most strongly influenced by groundwater level and nitrogen and phosphorus fertilisation rates, and to a lesser extent by plant-available soil potassium, mowing date, total soil nitrogen, potassium fertilisation rate, age of each field, and livestock (cow or sheep). In general, a low fertilisation rate, high groundwater level, late mowing, low level of plant-available potassium, high level of total soil nitrogen, old fields and the presence of sheep promote a high species richness. However, of these variables only nitrogen fertilisation rate and groundwater level differ significantly between the organic and conventional farms and are therefore likely to be the abiotic drivers of the difference in species richness and vegetation composition between the farm types. Of these two, the difference in nitrogen fertilisation rate is a direct result of a difference in management philosophy, but the difference in groundwater level is not. We hypothesize that the latter difference is caused by economic drivers, whereby a less productive soil is an incentive for a changeover to organic farming. If this is the case indeed, the application of agri-environment schemes would be most effective in less productive (and naturally more species-rich) sites.     

Changes in dry matter and nutrient yields during the restoration of species-rich grasslands

Abstract. In two moderately productive grasslands that had been withdrawn from agricultural use, one on humic sandy soil and one on heavy clay, the decrease in dry matter, N-, P- and K-yields were measured for 14 and 11 years respectively, after fertilization had been stopped. The grasslands were mown twice a year. The yields and concentrations of nutrients were compared with those of already unproductive grasslands under the same growing conditions. On sandy soil, the annual dry matter yield decreased from 10.2 to 6.5 ton / ha in four years. After nine years, the dry matter (4.1 ton / ha) and K (20.5 kg / ha) yields were as low as those of the unproductive grassland. At that time yields and concentrations of N and P were still greater. The low K- concentration of 0.50 % and the lack of response upon N fertilizing indicate that the supply of K limits biomass production. On clay, during the first three years, the dry matter yield decreased from 10.2 to 5.0 ton / ha, after 6 yr the production increased again. The low N-concentration of 1.52 % after 10 yr indicates that N is likely to limit biomass production, but the low P-concentration of 0.14% indicates that P shortage can also be expected. In the lower productive grassland on sand the number of species that invaded was higher.     

Effects of ants on plant diversity in semi-natural grasslands

Ants (L. niger and L. flavus) build conspicuous mounds that are covered with vegetation. The aim of this study was to investigate whether the vegetation on ant mounds in semi-natural grasslands differed from that around the mounds. Another aim was to investigate whether the changes in the vegetation on ant mounds were influenced by grazing management or by habitat characteristics, semi-dry versus moist. Here, the total number of plant species and total plant cover were lower on ant mounds than in patches off-mound. The plant cover of perennials that form rosettes was twice as high on mounds inhabited by L. niger than on those inhabited by L. flavus. Only a few plant species were restricted to either ant mounds or adjacent field and the effects of ants on the plant diversity in semi-natural grasslands seemed to be low. Grazing management did not affect the differences in the vegetation on ant mounds and in equal-sized patches off-mound, whereas habitat characteristics affected ant-induced changes in vegetation cover of some plant species.     

Study of the relationships between weather conditions and the marathon race,and of meteorotropic effects on distance runners

The relationships between the results of Beijing International Marathon Races and the corresponding weather conditions are analysed quantitatively. There are obvious interrelationships between the marathon results and weather factors such as air temperature, wet bulb temperature and human biometeorological indices. For example, the correlation coefficient between the average times of the top ten finishers and temperature isr=0.8910. The meteorological conditions are classified into three categories, suitable, moderate and unsuitable for running a marathon race, and the optimum meteorological index is given. Also the concept of a meteorological result, i.e. the part of the actual performances fluctuating with the changes in weather conditions, is presented. This plays an important role in some kinds of sports such as marathon racing. Finally, the results of physiological tests are given, which illustrate the physiological reactions of long- and middle-distance runners to the surrounding temperature.     

Relative influence of size,connectivity and disturbance history on plant species richness and assemblages in fragmented grasslands

Questions: What is the relative influence of size, connectivity and disturbance history on plant species richness and assemblages of fragmented grasslands? What is the contribution of small fragments to the conservation of native species pool of the region? Location: Tandilia's Range, Southern Pampa, Argentina. Methods: Cover of plants was registered within 24 fragments of tall‐tussock grassland remnants within an agricultural landscape using modified Whittaker nested sampling. We analysed the influence of site variables related to disturbance history (canopy height, litter thickness) and fragment variables (size, connectivity) on species richness (asymptotic species richness, slope of the species–area curve) as well as on species assemblages by multiple regressions analysis and canonical correspondence analyses, respectively. Cumulative area was used for analysing whether small fragments or large fragments are more important to species diversity in the landscape. Results: Asymptotic species richness was significantly influenced by site variables, in particular by Paspalum quadrifarium's canopy height, but not by fragment variables. Species assemblages were also affected by site variables (12.2% of total variation), but no additional portion of the species assemblage variability was significantly explained by fragment size and connectivity. Sampling of several small fragments rendered more exotic and native species than sampling of few large fragments of the same total area. Conclusions: Our results agree with previous studies reporting low sensitivity of species diversity to size and isolation of grassland fragments in fragmented landscapes and high sensitivity of species diversity to local variables. The higher capture of regional native species pool by small grassland fragments than by few larger ones of equivalent accumulated area highlights the value of small fragments for conservation.     

Physical and chemical effects of litter on plant establishment in semi-natural grasslands

A field experiment was conducted in a semi-natural grassland to study the interspecific variation in the effect of litter on seedling emergence and establishment and separate physical from chemical effects. Seeds of seven forb species were sown in plots subjected to either litter amendment (0, 400 or 900 g m⁻²) or water extracts of litter (corresponding to 400 and 900 g litter m⁻²). In addition, an extract was treated with activated carbon to estimate the possible effects of secondary chemical compounds. The response to plant litter differed amongst species: negative, neutral and positive responses were observed. Anthriscus sylvestris was the only species with a strong positive response to litter. We found no consistent relation between seed size and response to plant litter. Physical effects of litter were generally stronger than chemical effects. However, water extract of litter inhibited emergence in three species. Activated carbon removed the negative effect of the litter extract, which suggests that the effect was caused by an inhibitory chemical compound rather than by increased competition in response to nutrients added via the extract. The balance between facilitative and inhibitory effects of litter depended on species identity and litter quantity. Facilitative effects dominated at low and intermediate quantities of litter, and inhibitory effects at high litter quantities. One species, Campanula rotundifolia, showed a switch from positive to negative responses with increasing quantities of litter. However, we found no general threshold for litter quantity valid across species.     

Changes of species richness pattern in mountain grasslands: abandonment versus restoration

Changes in plant species richness at various spatial scales were investigated by manipulative experiment in mountain grasslands. The aim of the research was to compare changes in species richness in newly abandoned sites and sites where restoration measures were applied after 20 years of abandonment. The plots were located in two vegetation types with different moisture regime. Species richness decreased significantly after abandonment, mainly at the finest spatial scale of 10 × 10 cm. There was significant increase of species richness on restored sites, but it was apparent mainly at a larger scale. However, even 4 years of regular mowing were not sufficient to restore species richness to the level typical for traditionally managed grasslands in the region. No significant difference was found in the performance of the 2 contrasting vegetation types (wet and dry) in relation to management measures. A significant difference in scale-dependent species richness was only observed. The dry type had a steeper species-area curve, with a lower number of species at the finest spatial scale. According to the results of the experiment, mountain grasslands are very vulnerable habitats, losing their conservation value quickly after abandonment. Restoration is possible due to an extensive species pool in the region, but return to the original species richness at all spatial scales is quite a long process.     

Conservation potential of semi-natural habitats for birds in intensively-used agricultural landscapes

Agricultural intensification resulted in substantial loss of farmland biodiversity. Semi-natural habitats may be viewed as potential buffers of these adverse impacts, but a rigorous assessment of their capacity for supporting farmland biodiversity is lacking. In this study, we explored conservation potential of two different types of semi-natural habitats for birds in intensively-used agricultural landscapes – farmland hedges (i.e., linear strips of shrubby and tree vegetation) and open scrubland (i.e., scattered shrubs and abandoned orchards). Specifically, we tested whether the abundance and species richness of birds differ between these habitats considering various species traits, such as habitat affinity (i.e., forest, farmland and urban species), diet specialization (i.e., animal eaters, plant eaters, and omnivores) and conservation status (Species of European Conservation Concern). We found that open scrubland hosted on average 37.9 bird species and 122.6 individuals per 1 km² of the transect, whereas farmland hedges hosted only 19 species and 61.8 individuals per 1 km² of the transect. However, results have substantially changed if we considered the area of suitable habitat into account. More specifically, open scrubland hosted more bird species and individuals when we considered open habitat species and the area of open habitats, whereas farmland hedges had higher species diversity and individuals of woodland bird species when we considered the area of woodland habitats. Similarly, analyses of habitat affiliations of individual species corresponded to the whole-community patterns; and revealed that several woodland bird species were mainly associated with farmland hedges (e.g., Chaffinch Fringilla coelebs, Common Nightingale Luscinia megarhynchos and Blackcap Sylvia atricapilla), whereas the open scrubland was preferred by open habitat bird species (e.g., Corn Bunting Emberiza calandra, Quail Coturnix coturnix and Skylark Alauda arvensis). These results demonstrate that semi-natural habitats, both open scrubland and farmland hedges, have large potential for promotion and conservation of bird communities within intensively used agricultural landscapes, as both may have represented suitable habitats for species with different ecological requirements. Therefore, management measures focused on the enlargement of the area of these habitats, in combination with suitable management (e.g., regulating the progress of natural succession in open scrubland; increasing structural diversity of existing farmland hedges), may substantially contribute to bird conservation within agricultural landscapes.     

Patterns in the structure of grassland butterfly communities along a gradient of human disturbance: further analysis based on the generalist/specialist concept

Kitahara and Fujii, in 1994, analyzed the butterfly communities along a gradient of human disturbance by applying the generalist/specialist concept. Butterfly species were classified into generalist or specialist species based on their voltinism (seasonal time dimension) and potential larval resource breadth (food dimension). The community structure and species composition showed the systematic changes along the gradient. To verify the generality of those trends, we monitored five grassland butterfly communities with varying degrees of human disturbance twice a month during 1985 by the line transect method at the foot of Mt. Fuji, central Japan, and analyzed their structure in a manner similar to that employed by Kitahara and Fujii. Most results were consistent with the patterns recognized by Kitahara and Fujii. The route (community) order based on increasing human disturbance was strongly and negatively correlated with butterfly species richness but with neither butterfly species diversity (H′) nor evenness (J′). Also, the degree of human disturbance was significantly and negatively correlated with the number of specialist species, but not with that of generalists, in a community. Butterfly species richness was more strongly correlated with the number of specialist species than with that of generalists. Our analyses also showed that the generalist species were distributed more widely over the communities than were the specialists. However, in contrast to the trend revealed by Kitahara and Fujii, there was no significant difference in the population densities and in the spatial population variability between the two species groups. As a whole, our analyses confirmed the consistency of most community patterns detected by Kitahara and Fujii. The causes of the inconsistencies in some patterns were thought to be mainly the present habitat conditions with a relatively short growing season at high altitudes. Received: October 19, 1999 / Accepted: June 5, 2000     

Flood events overrule fertiliser effects on biomass production and species richness in riverine grasslands

Question: Do severe winter flood events lift the nutrient limitation of biomass production in a river floodplain? How does this affect plant species richness? How long do the effects last? Location: Floodplain grassland on calcareous sandy loam near river Rhine in The Netherlands. Methods: Plots were fertilised with four treatments (control, N, P, N+P) for 21 years; plant species composition, vegetation biomass and tissue nutrient concentrations were determined every year between 1985 and 2005. Results: Fertilisation with N generally increased biomass production and reduced species richness, but these effects varied over time. During the first four years of the experiment, biomass production appeared to be co‐limited by N and P, while N fertilisation dramatically reduced plant species richness; these effects became weaker subsequently. Following two extreme winter floods in 1993–94 and 1994–95 and a drought in spring 1996, the effects of fertilisation disappeared between 1998 and 2001 and then appeared again. Flooding caused an overall reduction in species richness (from c. 24 to 15 species m^‐2) and an increase in biomass production, which were only partly reversed after ten years. Conclusions: Long time series are necessary to understand vegetation dynamics and nutrient limitation in river floodplains, since they are influenced by occasional flood and drought events, whose effects may persist for more than ten years. A future increase in flooding frequency might be detrimental to species richness in floodplain grasslands.     

Reproductive strategies of female butterflies: variation in and constraints on fecundity

ABSTRACT.

• 1 This study first examines the reproductive strategy of female Speyeria mormonia Edwards (Lepidoptera: Nymphalidae):
• 2 Egg weight and number laid per day decrease with age.
• 3 Survival and daily egg number may be affected by temperature; mean daily egg weight is not affected by temperature.
• 4 Daily egg number is not correlated with body size. In the central range of body size, egg weight is also not correlated with body size. However, exceptionally large or small females lay respectively heavier or lighter eggs than average.
• 5 A simple trade-off between offspring size and number does not occur within females on a daily basis, or among females averaged over their lifespans.
• 6 Fat body resources are depleted at a rate independent of body size.
• 7 Females are essentially monogamous.
• 8 Age-specific fecundity data reported here for S.mormonia are next compared with data for other Lepidoptera with different adult feeding habits and egg maturation patterns, and hence different possibilities for adult feeding to play a role in egg production. Based on these comparisons, I propose that the shape of the age-specific fecundity curve for each species under optimal conditions is constrained by the potential importance of adult nutrients in egg production.