Bryophyte biomass and species richness on the Park Grass Experiment,Rothamsted, UK

The relationships between bryophyte biomass and species richness and soil pH, nutrient applications and vascular plant biomass and species richness were analyzed for the Park Grass Experiment (Rothamsted, UK). The study examined the abundance of bryophytes in relation to long-term fertilizer and lime application and to fertilizer treatments recently being ceased on some plots. The probability of bryophytes being present on a plot increased with increasing soil pH, and on plots at soil pH 3.3–4.5, the lowest values in this experiment, there were virtually no mosses present. Total bryophyte biomass decreased with increasing vascular plant biomass and vascular plant richness. Both bryophyte biomass and species richness showed a curvilinear response to soil pH. Bryophyte biomass was markedly increased on plots where nitrogen (N) fertilization had recently been ceased. The abundance of the common bryophyte species showed individualistic responses to treatments. N had a negative effect on the abundance of Brachythecium rutabulum. Increasing soil pH, and the application of phosphorus (P) and potassium (K) fertilizer together, had a positive effect on Eurhynchium praelongum. This species was also negatively affected by N, but tolerated larger amounts of it (100–150 kg ha^–1 N) than B. rutabulum. An ephemeral moss, Bryum subapiculatum, had a unimodal response to soil pH but showed no response to N, P, K or other explanatory variables.     

Determinants of species richness in southern African fig wasp assemblages

Summary We investigated the species richness of 24 fig wasp (Hymenoptera) assemblages associated with southern African fig trees (Ficus species, Moraceae). Assemblage sizes ranged between 3 and 30 species on different host tree species, with parasitoids slightly outnumbering gall-forming phytophages. Ten potential taxonomic, geographic and ecological determinants of assemblage richness were examined. Galler richness differed significantly between taxonomic sub-groups of Ficus and was significantly correlated with several ecological characteristics of the host trees, but there was no species-area effect. Parasitoid richness was strongly correlated with galler richness. We conclude that both ecological and historical factors have combined to determine the numbers of species that form fig wasp assemblages.     

Determinants of avian species richness at different spatial scales

ABSTRACT. Studies of factors influencing avian biodiversity yield very different results depending on the spatial scale at which species richness is calculated. Ecological studies at small spatial scales (plot size 0.0025–0.4 km²) emphasize the importance of habitat diversity, whereas biogeographical studies at large spatial scales (quadrat size 400–50,000 km²) emphasize variables related to available energy such as temperature. In order to bridge the gap between those two approaches the bird atlas data set of Lake Constance was used to study factors determining avian species diversity at the intermediate spatial scales of landscapes (quadrat size 4–36 km²). At these spatial scales bird species richness was influenced by habitat diversity and not by variables related to available energy probably because, at the landscape scale, variation in available energy is small. Changing quadrat size between 4 and 36 km², but keeping the geographical extension of the study constant resulted in profound changes in the degree to which the amount of different habitat types was correlated with species richness. This suggests that high species diversity is achieved by different management regimes depending on the spatial scale at which species richness is calculated. However, generally, avian species diversity seems to be determined by spatial heterogeneity at the corresponding spatial scale. Thus, protecting the diversity of landscapes and ecosystems appears to ensure also high levels of species diversity.     

Determinants of species richness patterns in the Netherlands across multiple taxonomic groups

We examined the species richness patterns of five different species groups (mosses, reptiles and amphibians, grasshoppers and crickets, dragonflies, and hoverflies) in the Netherlands (41,500 km²) using sampling units of 5 × 5 km. We compared the spatial patterns of species richness of the five groups using Spearman’s rank correlation and used a stepwise multiple regression generalized linear modelling (GLM) approach to assess their relation with a set of 36 environmental variables, selected because they can be related to the several hypotheses on biodiversity patterns. Species richness patterns of the five groups were to a certain extent congruent. Our data suggest that environmental heterogeneity (in particular habitat heterogeneity) is one of the major determinants of variation in species richness within these five groups. We found that for taxonomic groups comprising a low number of species, our regression model explained more of the variability in species richness than for taxonomic groups with a large number of species.     

Reinforcement of reproductive isolation between adjacent populations in the Park Grass Experiment

It has been debated, ever since Charles Darwin and Alfred Russell Wallace disagreed about the matter, whether natural selection plays a role in reinforcing reproductive isolation during the earliest stages of speciation. Recent theory suggests that it can do so, but until now the empirical evidence has conspicuously lacked a case in which reinforcement has actually been observed to split a population. We show that this has occurred at least once in populations of the grass Anthoxanthum odoratum growing in the Park Grass Experiment where flowering time has shifted at the boundaries between plots. As a consequence, gene flow via pollen has been severely limited and adjacent populations that had a common origin at the start of the experiment in 1856 have now diverged at neutral marker loci.     

Determinants of species richness, endemism and turnover in European longhorn beetles

This study assessed the diversity patterns of a large family of beetles, Cerambycidae, in Europe and tested the following hypotheses: 1) richness gradients of this hyperdiverse taxon are driven by water and energy variables; 2) endemism is explained by the same factors, but variation between areas also reflects post‐glacial re‐colonization processes; and 3) faunal composition is determined by the same climatic variables and, therefore, beta diversity (species turnover) is related to richness gradients. Species richness, endemism and beta diversity were modelled using inventories of 37 European territories, built from a database containing the distributions of 609 species. Area, spatial position, and nine topographical and climatic variables were used as predictors in regression and constrained analysis of principal coordinates modelling. Species richness was mostly explained by a temperature gradient, which produced a south‐to‐north decreasing richness gradient. Endemism followed the same pattern, but was also determined by longitudinal variation, peaking in the southwestern and southeastern corners of the continent. Faunal turnover was explained by an important purely spatial pattern and a spatially structured environmental gradient. Thus, contrary to other groups, cerambycid richness was mostly explained by environmental energy, but not by water availability. Endemism was concentrated in the Iberian and Greek peninsulas, but not in Italy. Thus, the latter area may have been the major source of post‐glacial re‐colonization for European longhorn beetles or, otherwise, a poor refuge during glaciations. Turnover patterns were independent of the richness gradient, because northern faunas are nested in southern ones. Turnover, in contrast to richness, was driven by both the independent effects of climate and geographic constraints that might reflect dispersal limitation or stochastic colonization events, suggesting that richness gradients are more environmentally deterministic phenomena than turnover patterns.     

Evidence for constraint on species coexistence in vegetation of the Park Grass experiment

Repeated patterns, of a type that would be expected to result from limitations to species coexistence (i.e. assembly rules) were sought in the Park Grass experiment. This classical grassland experiment was sampled in two years, using replicated biomass samples. Variance in a number of measures was examined, and compared to the variance expected under appropriate null models, the latter based on assumptions of no interactions between species. In each case, an assembly rule would result in low variance. Examining variance in species richness between quadrats within a treatment, there was no indication of constraint on species co-occurrences; variance in richness was actually greater than expected under the null model, attributable to environmental variation or perhaps positive interactions between species. However, there was control on biomass, evidenced by variance in total biomass (i.e. over all species) within a treatment being significantly lower than expected under the null model. There was no indication of community structure based on guilds (i.e. functional types). Although there was in 1991 some, non-significant, indication of a constant proportion of species from the legume guild, there was no sign of such an effect in 1992. Searches for intrinsic guilds failed to converge. There was no indication at all of constancy in the proportional representation of guilds by biomass. Thus, there is good evidence for competitive control on plant growth, but none for control of species occurrences. There is no convincing evidence for guild structure in this community at the scale sampled. Possible conflict is discussed between the existence of evidence for temporal stability but the absence of evidence for spatial uniformity. It is concluded that most of the mechanisms proposed for temporal stability will not necessarily lead to control on spatial variation. For many mechanisms, this would depend on the spatial scale examined.     

Determinants of tapeworm species richness in elasmobranch fishes: untangling environmental and phylogenetic influences

Parasite species richness is a fundamental characteristic of host species and varies substantially among host communities. Hypotheses aiming to explain observed patterns of richness are numerous, and none is universal. In this study, we use tapeworm parasites of elasmobranch fishes to examine the phylogenetic and environmental influences on the variation in species richness for this specific system. Tapeworms are the most diverse group of helminths to infect elasmobranchs. Elasmobranchs are cosmopolitan in distribution and their tapeworm parasites are remarkably host specific; therefore, making this an ideal system in which to examine global patterns in species diversity. Here, we 1) quantify the tapeworm richness in elasmobranch fishes, 2) identify the host features correlated with tapeworm richness, and 3) determine whether tapeworm richness follows a latitudinal gradient. The individual and combined effects of host size, factors associated with water temperatures (influenced by latitude and depth), host habitat, and type of elasmobranch (shark or batoid) on measures of species diversity were assessed using general linear models. These analyses included tapeworm host records for 317 different elasmobranch species (124 species were included in our analyses) and were conducted with and without taking into account phylogenetic relationships between host species. Since sharks and batoids differ substantially in body form, analyses were repeated for each host subset. On average, batoids harboured significantly more tapeworm species than shark hosts. Tapeworm richness in sharks was influenced by median depth, whereas no predictor variable included in our models could adequately account for interspecific variation in tapeworm richness in batoid hosts. The taxonomic diversity of tapeworm assemblages of sharks and batoids was influenced by median depth and median latitude, respectively. When the influence of host phylogeny is accounted for, larger hosts harbour a greater tapeworm richness, whereas hosts exploiting wider latitudinal ranges harbour more taxonomically distinct tapeworm assemblages. Species richness and taxonomic diversity of tapeworm assemblages in elasmobranch fishes are influenced by different evolutionary pressures, including host phylogenetic relationships, space constraints and geographical area. Our results suggest that ca 3600 tapeworm species have yet to be described from elasmobranch fishes.     

Relationship between grass species richness and ecosystem stability in Kruger National Park, South Africa

The relationship between grass species richness and ecosystem stability was investigated in the Kruger National Park. A total of 135 489 individual grasses were identified from 189 sites spread across 19 485 km² of savanna biome, making this one of the largest studies of its kind. At each site, grass percentage abundance and standing crop were measured at one year intervals to provide an index of ecosystem function. A severe drought that affected the region between 1991 and 1993 provided a convenient perturbation. At the height of the drought, mean species richness declined by 12.7% while standing crop declined by 38.1%, from 3199 to 1979 kg ha^?1. Percentage grass abundance declined to 87.5% of its pre‐drought value. After the drought had passed species richness, standing crop and percentage abundance recovered to 92.1%, 113.8% and 92.8% of their pre‐perturbation values, respectively. Statistical analysis of these responses revealed that grass assemblages of low species richness were more resistant to drought than assemblages of high species richness. Species‐poor sites also showed better recovery from perturbation after the drought had passed. These findings suggest that ecosystem stability may be negatively related to grass species richness in South African savanna grasslands.     

Testing the distinctness of shoot ionomes of angiosperm families using the Rothamsted Park Grass Continuous Hay Experiment

? The ionome is the elemental composition of a tissue or organism. Phylogenetic variation in the ionomes of plant shoots has been widely reported based on controlled experiments, vegetation surveys and literature meta-analyses. However, environmental effects on phylogenetic variation in shoot ionomes have not been quantified. This study tests the hypothesis that phylogenetic variation in shoot ionomes is robust to environmental perturbation and that plant families can be distinguished by their shoot ionomes. ? Herbage was sampled from six subplots of the Rothamsted Park Grass Experiment. Subplots had received contrasting fertilizer treatments since 1856. Herbage was separated into its constituent species (n?=?21) and concentrations of eleven mineral elements were determined in dried shoot material. ? Shoot concentrations of calcium (Ca), zinc (Zn), manganese (Mn), magnesium (Mg) and sodium (Na) showed significant variation associated with plant species, and responded similarly to fertilizer treatments in diverse plant species. Species?×?treatment interactions were indicated for phosphorus (P), potassium (K), nickel (Ni), copper (Cu) and iron (Fe). Plant families could be distinguished by their shoot ionomes. The most informative elements for discriminant analysis were Ca?>?Mg?>?Ni?>?S?>?Na?>?Zn?>?K?>?Cu?>?Fe?>?Mn?>?P. ? Whilst shoot ionomes were sensitive to fertilizer treatment, phylogenetic variation in a subset of the shoot ionome (Ca, Zn, Mn, Mg) was robust to this environmental perturbation.     

Soil moisture mediates association between the winter North Atlantic Oscillation and summer growth in the Park Grass Experiment

Several aspects of terrestrial ecosystems are known to be associated with the North Atlantic Oscillation (NAO) through effects of the NAO on winter climate, but recently the winter NAO has also been shown to be correlated with the following summer climate, including drought. Since drought is a major factor determining grassland primary productivity, the hypothesis was tested that the winter NAO is associated with summer herbage growth through soil moisture availability, using data from the Park Grass Experiment at Rothamsted, UK between 1960 and 1999. The herbage growth rate, mean daily rainfall, mean daily potential evapotranspiration (PE) and the mean and maximum potential soil moisture deficit (PSMD) were calculated between the two annual cuts in early summer and autumn for the unlimed, unfertilized plots. Mean and maximum PSMD were more highly correlated than rainfall or PE with herbage growth rate. Regression analysis showed that the natural logarithm of the herbage growth rate approximately halved for a 250 mm increase in maximum PSMD over the range 50-485 mm. The maximum PSMD was moderately correlated with the preceding winter NAO, with a positive winter NAO index associated with greater maximum PSMD. A positive winter NAO index was also associated with low herbage growth rate, accounting for 22% of the interannual variation in the growth rate. It was concluded that the association between the winter NAO and summer herbage growth rate is mediated by the PSMD in summer.     

Effects of spatial heterogeneity on butterfly species richness in Rocky Mountain National Park, CO, USA

We investigated butterfly responses to plot-level characteristics (plant species richness, vegetation height, and range in NDVI [normalized difference vegetation index]) and spatial heterogeneity in topography and landscape patterns (composition and configuration) at multiple spatial scales. Stratified random sampling was used to collect data on butterfly species richness from seventy-six 20 × 50 m plots. The plant species richness and average vegetation height data were collected from 76 modified-Whittaker plots overlaid on 76 butterfly plots. Spatial heterogeneity around sample plots was quantified by measuring topographic variables and landscape metrics at eight spatial extents (radii of 300, 600 to 2,400 m). The number of butterfly species recorded was strongly positively correlated with plant species richness, proportion of shrubland and mean patch size of shrubland. Patterns in butterfly species richness were negatively correlated with other variables including mean patch size, average vegetation height, elevation, and range in NDVI. The best predictive model selected using Akaike’s Information Criterion corrected for small sample size (AIC_c), explained 62% of the variation in butterfly species richness at the 2,100 m spatial extent. Average vegetation height and mean patch size were among the best predictors of butterfly species richness. The models that included plot-level information and topographic variables explained relatively less variation in butterfly species richness, and were improved significantly after including landscape metrics. Our results suggest that spatial heterogeneity greatly influences patterns in butterfly species richness, and that it should be explicitly considered in conservation and management actions.     

Determinants of bird species richness: role of climate and vegetation structure at a regional scale

We examined the respective roles of climate and vegetation structure on geographical variation in bird species richness. The Province of Buenos Aires (central-eastern Argentina) was divided into 146 squares of 50 km on a side. For each square we evaluated the number of bird species, the value of thirteen climatic variables, and the value of a vegetation strata index. The climatic matrix was analyzed by Principal Component Analysis (PCA), and the first factors resulting from PCA were considered as multifactorial climatic gradients. Simple and Partial Correlation Analysis among bird species richness, vegetation strata, and the first two factors derived from PCA (65% of total variation) indicated that bird richness distribution was determined by the availability of vegetation strata, associated with different vegetation types that, at the same time, were influenced by the climatic conditions summarized in the first climatic factor (a gradient of precipitation, relative humidity, annual termical amplitude, and frost occurrence). This relationships reflect the complexity of factors that can act directly as well as indirectly on the geographical patterns in species richness. Also, we evaluated the importance of study scale comparing our results with previous studies at macrogeographic and local scales, found out that the vegetation structure was the principal determinant of bird species richness at this three geographical scales.     

Determinants of species richness in generalist and specialist Mediterranean butterflies: the negative synergistic forces of climate and habitat change

Although it is well established that butterfly richness is affected by climate and human factors (e.g. habitat disturbance and degradation) at different spatial scales, the drivers behind these changes vary greatly according to the geographical region and the ecology of the species concerned. It is essential that this variation be understood if trends in diversity are to be predicted with any degree of confidence under a scenario of global change. Here we examine patterns of butterfly species richness among groups differing in degree of habitat specialization, diet breadth and mobility in the north‐west Mediterranean Basin, a European hotspot for this taxon. We analyze a large number of butterfly communities and take into consideration the main potential drivers, that include climatic, geographic and resource variables, landscape structure and human environmental impact at different spatial scales. Our study shows that both climatic and anthropogenic factors play an important role in determining butterfly species richness in the north‐west Mediterranean Basin, but that their relative impact differs between specialist and generalist groups. At lower altitudes, water availability, a product of the interplay between temperature and rainfall, and negative effects of temperature appear as the most determinant factors. Maximum diversity was observed at mid‐altitudes, which reveals the importance from a conservation point of view of Mediterranean mountain ranges. The results suggest serious population declines in specialist species restricted to mountain areas as a result of climate warming in combination with habitat loss caused by the abandonment of grazing and mowing. They also suggest negative trends for generalist species due to an increase in aridity in combination with an increase in intensification of human land use in lowland areas. Such synergies are expected to lead to rapid declines in Mediterranean butterfly populations in the coming years, thereby posing a severe threat for the conservation of European biodiversity.