Small-scale patterns of species richness in Swedish semi-natural grasslands: the effects of community species pools

Temperate semi-natural grasslands are known for their high plant species richness at small spatial scales. We examined the variation in small-scale species richness in a sample of 63 sites from Swedish semi-natural grasslands, located as fragments in the modem landscape dominated by forest and agricultural land. Data were obtained from two spatial scales at each site. 1 dm² and 4 m². Using an analysis based on a Monte Carlo simulation, we found support for the species-pool hypothesis: a high species richness at the I dm- scale was associated with high species richness at the 4 m2 scale. The conclusion from this pattern analysis would, however, be considerably strengthened if we could reduce the likelihood that other mechanisms than sampling from species pools of unequal size influence the pattern of small-scale species richness. Additional analyses were made in order to identify such mechanisms. We examined whether four putative key traits: seed size, seed production, plant size and reproductive allocation were different among species at comparatively species-rich vs species-poor I dm' plots. We found only a little evidence for such differences. There was a weak tendency that species in the plots with high species richness possessed larger (and fewer) seeds than species from species-poor plots. Our results are congruent with the main prediction of the species pool model: variation in small-scale species richness (1 dm²- scale) is basically a result of sampling from unequally sized community species pools (4 m² scale). Variation in species richness between the 4 nr semi-natural grassland "patches" may thus be sought for among mechanisms operating al larger spatial scales than 4 m². We briefly discuss such mechanisms, based on other studies performed in the same study area.     

Two ways of trapping seeds in alpine environments, Lapland, Sweden

Seed dispersal in two habitats was investigated in the mid-alpine Latnjajaure valley, in northernmost Swedish Lapland. The seed rain size was measured using artificial seed traps and natural snowbed at a heath and a meadow site, c. 1000 m a.s.l. The average seed rain size trapped in the snowbeds varied from 18 seeds m^-2 at the heath site, to 96 seeds m-² at the meadow site. On average, the heath trap station had 177 seeds m-² and the meadow station, 218 seeds m². At each site, the vegetation was inventoried within a circular area with a radius of 20 m. Overall, the species assembly in the seed rain reflected the surrounding vegetation. However, there were additional species in the seed rain as well. At the seed trap station on the wind-exposed heath, 78% of the seeds originated from sites outside the homogeneous vegetation around the trap. These seeds were dispersed from locations at least 150 m from the trap site. By trapping a higher number of species than artificial traps and a higher number of seeds per unit area, snowbeds served as effective 'seed traps'. However, the location of the sampling spot in the snowbed is crucial due to a decrease in seed density towards the edge. In this study, with the aim of trapping the main dispersing species in the area, the optimal artificial seed trap size appears to be - 1.5–2.0 m². There was also a positive correlation between the mean July and August temperature, the seed rain size, and the number of species found. Thus, through brief visits to remote locations, the study of seed dispersal can be accomplished effectively when snowbeds are present on the landscape.     

Body weight distributions of European Hymenoptera

Species number–body weight distributions are generally thought to be skewed to the right. Hence it is assumed that the number of relatively small species is larger than the number of relatively large species. While this pattern is well documented in vertebrates, comparative studies on larger invertebrate taxa are still scarce. Here I show that the weight distributions of European Hymenoptera (based on 12 601 species body weight data compiled from major catalogues) do not exhibit a general trend towards right skewed species–body weight distributions. Skewness did not depend on the number of species per taxon. Species richness peaked at intermediate body weights irrespective of taxonomic level. Kernel density analysis revealed that hymenopteran taxa had between one and four peaks in their size distributions with larger taxa having fewer peaks. Within genus variability in body weight was allometrically related to mean body weight (σ²=μ^1.81) in line with a proportional rescaling pattern. These results call for a rethinking about the generality of current vertebrate centred models of body size evolution.     

19th century woodland structure controls stand-scale epiphyte diversity in present-day Scotland

Axioms developed from island biogeography theory (i.e. species–area relationships, effects of fragmentation and isolation) are central to the development of conservation strategy. Within this context, the 'extinction debt' hypothesis brings into question an often assumed relationship between species richness and present-day spatial habitat structure (i.e. extent, fragmentation), suggesting instead that the richness and composition of biological communities may lag behind spatial changes in habitat. We examined evidence for an extinction debt among epiphytic lichens, a highly diverse biological group of significant conservation concern. Using sites in Scotland, we compared epiphyte species richness in smaller-scale habitat units (aspen stands) to larger-scale woodland structure (extent and fragmentation) measured at two spatial scales (1 km² and 4 km²) and for two timeframes, modern (1990s to 2000s) and historic (1860s to 1880s). Species richness was positively related to woodland extent and negatively related to woodland fragmentation; however, richness was explained better by historic woodland structure at a 1-km² scale, than by modern woodland structure. The results indicate: (1) a coupling of stand-scale epiphyte assembly and dynamics of the wider woodland ecosystem, and (2) a significant lag in the response of epiphyte species richness to habitat spatial structure. However, the effect of spatial habitat structure is different between species groups with contrasting traits. The effect of decreasing woodland extent on epiphyte richness is generally more severe for microlichens (comprising a greater number of rare and specialist species) than the more generalist macrolichens.     

Seed banks in Danish deciduous forests: species composition, seed influx and distribution pattern in soil

Samples of soil from six forest stands of different age and composition were collected, and the seedlings emerging during germination experiments were recorded Total seed bank densities varied between 2230 and 15,630 seeds m⁻² The seed bank size was positively correlated with the yearly seed influx, which constituted from 400 to 2720 seeds m⁻² Juncus effusus dominated the seed banks with generally 2000 to 3300 seeds m⁻², to exceptionally 10820 seeds m⁻² Seeds of Rubus idaeus, Betula pubescens and Carex spp were also abundant in the soil Similar numbers of species were present in the seed banks and vegetation, but only 7-48% were species in common Anemochorous species were more frequent in soil than above-ground, while zoochorous species were rare in soil but abundant in the vegetation A measure of the occurrence of seeds in relation to soil depth was designed Seeds of species with short-term survival in soil (e g Anemone nemorosa, Epilobium angustifolium, Oxalis acetosella and Stellaria holostea) were confined to the top soil, while many well known long-term survivors (e g Carex piluhfera, Poa annua, Rubus idaeus and Stellaria media) were found predominantly deep in the soil The horizontal distribution of seeds of herbaceous species forming short-term seed banks was largely random, while seeds of herbs capable of long-term survival in soil were significantly aggregated Except for one old and species-poor site, the dormant flora indicated generally high levels of disturbance in the past     

The environmental determinants of the insular butterfly faunas of the British Isles

A multiple regression analysis was performed upon selected environmental variables for a series of islands in the British Isles, to establish their effects upon the size of the butterfly fauna, measured as he number of species regularly breeding, S_B .
So that the data be normally distributed, the regression analyses were performed upon log₁₀ transformed data only, with the data for outliers, mainland Britain and Ireland, the two largest islands, excluded.
Most highly correlated with the number of butterfly species breeding upon an island is the number breeding within a 25 km radius of the nearest point of the mainland, r ²=0.5941, followed by the correlations with the latitude of the mid-point of the island, r ²=0.5541, the number of plant species comprising the island Hora, r ²=0.5225, and the distance separating the island from the mainland, r ²=0.4514.
A partial correlation analysis confirms the importance of the parameters distance separating the island from the mainland, D ₁, and the size of the faunal source S _F , and rejects the importance of the size of the flora and the latitude of the island. This is further confirmed by the results of a step-wise regression analysis, the two variables D ₁ and S_F accounting for 66% of the variation of the butterfly fauna.
If an alternative measure of isolation, D ₂, which allows for the geographical clumping of islands, is combined with the variable S_F , then 69% of the variation of the butterfly fauna is accounted for.     

Elevated CO2 concentration, nitrogen use, and seed production in annual plants

Elevated atmospheric CO₂ concentration ([CO₂]) stimulates seed mass production in many species, but the extent of stimulation shows large variation among species. We examined (1) whether seed production is enhanced more in species with lower seed nitrogen concentrations, and (2) whether seed production is enhanced by elevated [CO₂] when the plant uses more N for seed production. We grew 11 annuals in open top chambers that have different [CO₂] conditions (ambient: 370 μmol mol⁻¹, elevated: 700 μmol mol⁻¹). Elevated [CO₂] significantly increased seed production in six out of 11 species with a large interspecific variation (0.84–2.12, elevated/ambient [CO₂]). Seed nitrogen concentration was not correlated with the enhancement of seed production by elevated [CO₂]. The enhancement of seed production was strongly correlated with the enhancement of seed nitrogen per plant caused by increased N acquisition during the reproductive period. In particular, legume species tended to acquire more N and produced more seeds at elevated [CO₂] than non-nitrogen fixing species. Elevated [CO₂] little affected seed [N] in all species. We conclude that seed production is limited primarily by nitrogen availability and will be enhanced by elevated [CO₂] only when the plant is able to increase nitrogen acquisition.     

Pollen transfer patterns in a distylous plant with overlapping pollen-size distributions

In this study we quantify the relative rates and amounts of between-morph (legitimate) cross-pollination, within-morph (illegitimate) cross-pollination and self-pollination in a natural population of distylous Gaertnera vaginata (Rubiaceae) endemic to the oceanic island of La Réunion in the Indian Ocean. Since pollen size distributions of each morph overlap we develop a method, based on chi-squared comparisons of observed to expected pollen size distributions, to quantify pollen transfer proficiencies. This method, unlike previous methods, does not assume an equal chance of pollen transfer for pollen grains in the region of overlap. The probability that a single pollen grain from a short-styled plant undergoes legitimate pollen transfer to long-styled flowers (4.19×10⁻²) was more than twice that for illegitimate transfer to other short-styled plants (1.56×10⁻²). In contrast, the probability that a single pollen grain from a long-styled flower undergoes legitimate pollen transfer to short-styled flowers (1.65×10⁻²) was no greater than that of illegitimate pollen transfer to other long-styled plants (2.15×10⁻²). The transfer proficiencies of individual pollen grains are thus highly asymmetric. However, because long-styled flowers produce significantly more pollen, the ratio of legitimate to illegitimate pollen on the stigmas of short-styled flowers (1.85) markedly exceeded that for long-styled flowers (1.11). Short-styled flowers incurred significantly higher levels of self-pollination than long-styled plants, emphasizing the importance for self-pollination to be removed before comparing stigmatic pollen loads. The method we propose could be applied to other distylous species with pollen size overlap between morphs and thus allow for a more widespread study of the pollination ecology of heterostylous species.     

Seedling recruitment in semi-natural pastures: the effects of disturbance, seed size, phenology and seed bank

We examined recruitment patterns in semi-natural pastures and their relationships to disturbance and species composition at different spatial scales (0.01 m² up to 4 m²) of both the vegetation and the seed bank. Possible associations between seedling recruitment and phenology were also studied. The study was performed in four pastures with different management history. Seedling recruitment was generally enhanced by disturbance, with the greatest effect for small-seeded species that germinate in the autumn. The local species-pool contributed to a large extent to seedling recruitment; 83 % of the recruited species were found in the vegetation, whereas 44 % of the recruited species were found in the seed bank. There were a total of 88 species found in the vegetation at different spatial scales; 61 % were recorded at the smallest spatial scale (0.01 m²). The area sampled at this scale comprised 1.25% of the area examined, indicating a small scale structuring of diversity in the pastures. The species number in the vegetation varied between 7 and 14 for the smallest scale (0.01 m²), and between 23 and 42 for the largest scale (4 m²), in the four pastures. The species number in the seed bank was less variable. Like most studies of perennial grasslands, we found no close correspondence between species distribution in the vegetation and in the seed bank. Two of the pastures had a joint management history and exhibited a close similarity of species in the seed bank, despite that one of the pastures was abandoned already in the beginning of this century.     

Fast locomotion of some African ungulates

Ten species of ungulate were filmed, galloping in their natural habitat. They ranged in size from Thomson's gazelle (about 20 kg) to giraffe (about 1000 kg). They were pursued to make them run as fast as possible. The films have been analysed to determine speed, stride frequency, stride and step lengths, and duty factors. The dependence of these quantities on body size is discussed.  

Summary:

Fast locomotion of zebra, giraffe, warthog and seven species of Bovidae has been studied. The animals were filmed from a pursuing vehicle while galloping in their natural habitat.
Stride frequency was more closely correlated with limb length (represented by hip height) than with body mass. Mean stride frequency was proportional to (hip height)^-0·51 and maximum stride frequency to (hip height) ^-0·63.
Maximum speed was between 10 and 14 m s ^-1 for all species except buffalo (7 m s ^-1). It was not significantly correlated with body mass.
Since the small species ran at least as fast as the large ones they attained higher Froude numbers. Relative stride length was approximately 1·8 (Froude number)^0·39 for all species, irrespective of size. Relative step length was approximately 0·65 (Froude number)^0·2, both for the fore feet and for the hind ones. The vertical forces exerted by the feet are proportional to (body weight)×(Froude number)^0·2 so the forces at maximum speed are larger multiples of body weight for small species than for large ones.     

Influence of habitat characteristics on small mammals in a Mexican high-altitude grassland

Small mammals in a high-altitude grassland area close to Mexico City were studied. Populations of 10 species were censused using live traps in 48 sample quadrats. Within each quadrat, vegetation characterization, including complete floristic listings, cover values for species and layers and values of habitat modification, were assessed. Habitats were described according to plant communities identified using ordination and classification methods. Nine different plant communities were obtained. Densities and abundance of all small mammal species were calculated for each of the habitats classified. Peromyscus alsloni was the most abundant species in all habitats, reaching maximum densities of 55 ha⁻¹ in pine forest with dense ground and herb layer. Peromyscus melanotis also occurred in all habitats but at lower densities (maximum 29 ha⁻¹). Reithrodontomys megalotis was found in all habitats except in tall dense grassland. Densities for this species were generally low (1-9 ha⁻¹) but reached 19 ha⁻¹ in short dense grassland. All other species were largely absent from 4–8 habitats and showed very low densities (0.75–4 ha⁻¹). The densities of the more abundant species were largely correlated with more open habitats and higher indices of habitat modification. Lower altitude grassland habitats have a greater abundance of small mammals and a higher species richness than the medium and higher altitude, physiognomically more complex habitats. Species richness was highest in tall pine-alder forest with a species-rich, dense herb layer and lowest in pine forest with dense ground and herb layers. Species richness was positively correlated with overall small mammal density.     

Plant responses to livestock grazing frequency in an Australian temperate grassland

Livestock grazing is often thought to enhance native plant species co-existence in remnant grasslands but may also favour exotic invaders. Recommendations for appropriate grazing strategies are needed, for which an understanding of the response of plant species is necessary. We explored the response of plant species and plant functional groups to grazing in temperate grassland of the Monaro Tablelands of south-east Australia by comparing species abundance in adjacent areas that differed in livestock grazing regime (minimal, infrequent and frequent). We also examined whether species with similar responses to grazing share certain traits and consider whether these traits might provide a useful method of assessing grazing impact. At the scale measured (0.25 m²), an infrequent grazing regime maximised plant species co-existence in these grasslands due to widespread invasion by exotic plant species at infrequent grazing intensity. Many native species declined in abundance when grazing frequency increased from minimal to infrequent. Annuals invaded under infrequent grazing while perennials declined most strongly under high frequency grazing. Low levels of grazing apparently reduce cover and create sites suitable for seed recruitment whereas more frequent grazing reduces the persistence of perennials. While there was a tendency for native species to be more susceptible to grazing impact than exotics, plant traits, in particular longevity (perennial, annual) provided a better prediction of the response of plants to grazing. Although a few native plant species persisted at high grazing frequency, even infrequent livestock grazing may not be appropriate for the conservation of many native perennial grassland species. Targeted reductions in grazing frequency may be necessary to enable the long-term coexistence of grazing susceptible species.     

Taxonomic partitioning shedding light on the diversification of parasite communities

Power relations were used for 297 parasite communities of fish, bird and mammal hosts between the number of genera (G) and the number of species (S) per community to test for the relative importance of different processes in their diversification. For the bird hosts the relation was G∝S^0.92, and for fish hosts it was G∝S^0.97, but for mammal hosts, the number of parasite genera increases more slowly as a function of species richness (G∝S^0.83), suggesting, among other possibilities, a higher rate of within-host parasite speciation in mammals than in other vertebrates.     

Possible Involvement of K^＋/Na^＋ in Assessing the Seed Vigor Index

More substances leaked from a higher-vigor seed sample than from a lower-vigor sample. This indicates that, in some cases, electric conductivity does not represent seed vigor level very well, especially for high-vigor seeds. Results from germination, germination index, leachate conductivity, and the ratio of K^＋/Na^＋ from three-seed lots of Chinese cabbage （Brassica pekinensis （Louv.） Rupr） showed that K^＋/Na^＋ correlated well with germination and germination index. The ability of K^＋/Na^＋ to indicate well changes in vigor was further supported by investigation in soybean （Glycine max （L.） Merr.） seeds and another cultivar of Chinese cabbage seeds. Thus, seed leakage of K^＋/Na^＋ can accurately indicate seed vigor, whereas the conductivity test failed to do so. Furthermore, K^＋/Na^＋ showed up bigger quantitative differences in vigor level than did the conductivity test. This findings provide a more sensitive and accurate index for the assessment of seed vigor. The mechanisms of Na^＋ and K^＋ ion transport are also discussed.     

Lichen recolonization in London under conditions of rapidly falling sulphur dioxide levels, and the concept of zone skipping

HAWKSWORTH, D. L. & McMANUS, P. M., 1989. Lichen recolonization in London under conditions of rapidly falling sulphur dioxide levels, and the concept of zone skipping. The lichen flora on trees and wood at 50 sites in north-west London was examined in 1988 to determine what changes had taken place since 1980, a period when mean winter SO₂ levels had fallen from around 130 μg m^-3 to within the range 29–55 μg m^-3 in the study area. Forty-nine species were found. The fruticose lichens Evernia prunaslri and Ramalina farinacea now occur in many central London parks, and the yellow green foliose Parmelia caperata was discovered at 12 sites (previously only at one, Ruislip, in 1980). Comparable numbers and growth forms of species occurred throughout the area. An assemblage of zone 4–5 species (Hawksworth & Rose, 1970 Nature, 227: 145–148) tolerant of mean winter SO₂ levels of 60–70 μg m^-3 failed to penetrate central London, although more sensitive species of zones 6 and 7 had; this phenomenon, which has not previously been reported, is referred to as 'zone skipping' and is attributed to the speed at which mean sulphur dioxide levels in London have fallen. Species returning are those to be expected on the basis of previously-published field correlations with mean winter SO₂ levels. Twenty-five species not seen within 16 km of the centre of London on trees or wood during this century were found; of these, eight had not been seen during the last 200 years.     

Gastric evacuation rates and daily ration of piscivorous coho salmon, Oncorhynchus kisutch Walbaum

Effects of temperature and meal size on gastric evacuation rates of juvenile coho salmon, Oncorhynchus kisutch , consuming sockeye salmon, O. nerka , fry were examined and used in the estimation of daily meal, daily ration and number of fry consumed by coho in Chignik Lake, Alaska. Evacuation of fry by coho was best described by a negative exponential model (average R² = 0.93). A square root model also provided a good fit (average R² = 0·93), but the y-intercepts deviated more from the expected value than did the y-intercepts of the exponential model. The effect of temperature ( T , 5–13° C) and meal size (MS, 0·166–0·367 g) on the exponential evacuation rate (r_e, h^-1) could be described as
In the lake, coho fed continuously during the 24-h period in early June 1986 and 1987. Estimates of daily meal and ration of coho calculated by the Eggers method and the geometric mean of prey weight ranged from 0·224 to 0·435 g (2.1–4.4% body wt) depending on location and year. The Elliott & Persson method provided similar estimates of food consumption, whereas estimates based on the Pennington method and square root evacuation of prey differed from the exponential models. Sockeye fry represented 93% of the total prey weight. The average number of sockeye fry consumed per coho per 24 h, based on the arithmetic mean of prey weight, was 3·0–3·9 fry.     

Efficiency and allometry of seed production in grassland forbs: site and year comparative study in the Italian Eastern Pre-Alps

Forbs are important biodiversity components of grasslands and are often threatened by management intensification. As most forbs propagate predominantly by seed, knowledge of their seed regenerative traits would improve the conservation and restoration of forb-rich ecosystems. The main seed production traits of six forbs that are common in European species-rich grasslands were studied by collecting fertile shoots from different sites and over several years. Among sites and years, variability was high, particularly in the number of inflorescences per shoot, which affected ovule production more than any other trait. Relationships between inflorescence size and the number of ovules were mainly negatively allometric or almost so, with lower flower densities in larger inflorescences. The average ovule-to-seed transformation efficiency was 58%. There was significant variation among collections of the same species, and even more between species. Species with a low ovule-to-seed transformation efficiency generally exhibited compensatory, high seed viability. Large inflorescences had high ovule to seed utilization values, probably because of better nutrient conditions. Seed germinability (average, 30%) was much lower than seed viability (average, 54%); therefore, seed dormancy was an important feature of the species studied.     

Relationship between the seed rain and the establishment of vegetation in two areas abandoned after peat harvesting

In this study the number and species composition of diaspores dispersing into two newly abandoned peat harvesting areas in Finland were investigated. In an area abandoned six years earlier a total of 2978 living seeds m⁻², representing 18 species, was captured by using water-filled traps during two summers. In a one-year-old succession area the total number of living seeds m⁻² was 2241, representing 16 plant species. The soil seed banks were found to be empty of viable seeds. In neither area there appeared to be any relation between number of dispersing seeds and of plant individuals of the same species in the pioneer vegetation. It is suggested that unfavourable conditions on the soil surface for seed germination and seedling growth of most dispersed species are majors factor in determining the structure of the established vegetation.     

How important are species richness,species evenness and interspecific differences to productivity? A mathematical model

We present a theoretical model to quantify the influence of diversity on productivity and nutrient acquisition in plant communities during exponential growth. The model fractionates diversity into three components, namely species richness ( S ), species evenness ( E ) and the degree of difference between species ( D ). The influence of each of these components is assessed individually: S is varied by changing the number of species, E by changing their population size, and D by changing the range of species traits critical to productivity (specific nutrient uptake rate, Σ _r , or nutrient use efficiency, NUE ). D was quantified as the coefficient of variation of Σ _r or NUE . All three components of diversity enhance the biomass and nutrient stocks in the community, but the response patterns are different. Species richness has a saturating influence, whereas effects of E and D are linear and exponential, respectively. In all cases the non-linear dependence of productivity and nutrient acquisition on Σ _r and NUE during exponential growth was the single mechanism underlying these effects. This causes the presence of plants with extreme traits to promote productivity, and S , E and D all affect the abundance and/or intensity of these extremes. The model offers a framework to explain differences between experimental observations, and suggests a concept of diversity where S and E are structural components and D a qualitative or functional component, which modulates the influence of the two others. We propose to explicitly recognise D as an integral constituent of plant diversity in future studies.