Habitat Diversity in the Conservation of the Grassland Auchenorrhyncha (Homoptera: Cercopidae,Cicadellidae, Cixidae,Delphacidae) of Northern Britain

Definition of northern British grassland Auchenorrhyncha habitats was carried out using a classification based on analysis of data from 351 sites, involving 121 species, located between Greater Manchester and northern Scotland. Ten habitats were identified showing little influence of geographical position and exhibiting a basic upland-lowland trend. Other factors influencing habitat and species assemblage distribution were soil water, vegetation structure and land cover. An analysis of the species data with satellite-derived land cover data indicated that the lowland covers of tilled land, coast and urban and the upland covers of heath grassland and shrub heath were most important in affecting both species and assemblage distribution. The large-scale survey of grassland sites provided new information on both the ecology and distribution of individual Auchenorrhyncha species. Some were limited to specific habitat types but a considerable number were generalist species found in most or all of the 10 habitat types but showing preferences within upland to lowland or wet to dry site gradients. The ability to generate a subtle grassland Auchenorrhyncha habitat classification with large-scale survey results from standardised and reproducible sampling increases the potential for using habitat diversity for the conservation of grassland Auchenorrhyncha. Habitat preservation would also ensure that species richness (biodiversity) is maintained and that the habitats of rare species are conserved.     

Successional processes induced by fires on the northern offshore islands of New Zealand

A modern pollen-vegetation data set of 46 samples is presented from subantarctic Campbell Island, 600 km south of the New Zealand mainland. The sampled vegetation includes all major community types: maritime turf and grassland, sedge flushes, dwarf forest, scrub, cushion bog, tussock grassland, and high altitude graminoid turfs and tundra. Macrophyllous forbs—characteristic plants of subantarctic islands—are common throughout. Most taxa have highly restricted pollen dispersal, largely due to the short stature of the vegetation and the high proportion of insect-pollinated species. Percentages of pollen or spores of the dominant taxa have a significant positive correlation with the percent vegetation cover of the corresponding species, the exceptions being the widespread ferns Polystichum vestitum and Blechnum spp., and the ubiquitous macrophyllous forb, Bulbinella rossii .The relationship between the vegetation cover of a given taxon and its pollen representation was usually not strong enough to give confidence in a quantitative reconstruction based on pollen frequency alone. However, the broad vegetation groupings have characteristic pollen and spore spectra clearly related to the abundance of their dominant plant species. Detrended correspondence analysis of the pollen spectra grouped most sites according to their source vegetation type and generated a pattern similar to that of vegetation data analysed in a similar fashion in previous studies of the island. This study, together with recent work on Auckland Island pollen and spore representation, has resulted in a combined modem palynological data base of more than 100 sites for the New Zealand subantarctic islands.     

Airborne Grass Pollen Antigens in a Grassland as Studied by Immunoblotting with Anti-Lol p I Antibody

Airborne grass pollen in a grassland was measured using a Burkard air sampler and immunoblotting. The pollen antigens collected on the tape of the air sampler were transferred onto a nitrocellulose membrane, and visualized as purple spots by immunoenzymatic staining with anti-Lol p I antibody. These spots were quantified automatically using an image processing system. The airborne grass pollen antigens in the grassland where four species of grass were growing were prevalent mainly from evening to early morning during the first half of the pollen season, but in the daytime during the latter half of the season. The antigen dispersion time was confined to a few hours of the day. The flowering time of several grass species which were hydroponically cultured in the laboratory were also studied. Each species had a different flowering time. We believe that the time when an antigen is present during the day is related to the different flowering times of the various grass species growing in the grassland.     

Community reassembly: a test using limestone grassland in New Zealand

To examine community reassembly, we sampled grasslands on calcareous soil (4%–24% CaCO₃) in New Zealand that were largely composed of species introduced from Britain. We tested whether the British species had reassembled on New Zealand limestone into the same communities as they form on limestone in Britain. The vegetation of six New Zealand sites was sampled, each with ten 2 × 2 m quadrats that followed the standard methodology of the British National Vegetation Classification (NVC). Analysing species presence and cover using program TableFit with the full database of British species, the New Zealand species assemblages gave poor to moderate fits (40%–72%) to the communities of the NVC, and even then not to calcareous grassland, though one site did fit to a calcareous spring community. The poor fits can be partly attributed to the absence from New Zealand of many British calcareous-specialist species. On omitting from the NVC database all species not present in New Zealand, the fits increased somewhat to 48%–77%. Using this modified database, two sites fitted British calcareous grassland communities. These two sites are on thinner soil (<10 cm depth), under lower rainfall. Where fits were obtained to calcareous communities, the environment of the community in Britain matched very well that of the New Zealand site. It is concluded that environmental and perhaps biotic filtering has been strong enough in some sites to assort alien species into the same species assemblages as they form in their native range, indicating the Deterministic model of community structure. However, the absence of some species has prevented full reassembly.     

Extra petals in the buttercup (Ranunculus repens) provide a quick method to estimate the age of meadows

Background and Aims

There is a widely used crude method to estimate the age of hedgerows (Hooper''s rule) based on species'' richness. The aim of this study was to try and establish a similar field method for estimating the age of grasslands based on the accumulation of macro-somatic mutations.

Methods

A countrywide survey was carried out by the British public to investigate the relationship between grassland age and the number of Ranunculus repens (creeping buttercup) plants with extra petals. In addition the relationship between grassland age and R. repens pollen viability was also investigated.

Key Results

Each plant with flowers with additional petals in a sample of 100 was found to equate to approx. 7 years. A higher significant correlation was observed between pollen viability and population age; however, this is not amenable to providing field estimates.

Conclusions

The age of British grasslands can be easily and reliably estimated in the field by counting the number flowers with additional petals in R. repens in meadows up to 200 years old. An attempt to estimate the heritability of extra petals suggests that the phenotype results from the slow accumulation of somatic mutations in a species that primarily reproduces vegetatively.Key words: Conservation, meadow, Müller''s ratchet, petal number, pollen viability, Ranunculus repens, buttercup, somatic mutation     

Characteristics of Poaceae pollen grains as a tool to assess palaeoecological grassland dynamics in South America

Despite the dominance of grasslands during the last glacial period, especially in South America, the highly uniform morphology of Poaceae pollen grains has so far allowed only very few palynological studies based on Poaceae pollen. In our study we compare two methods of distinguishing between South American grassland ecosystems based on quantitative morphology of Poaceae pollen grains. We investigated data sets from Páramo in southern Ecuador, Campos de Altitude and Campos in south-eastern and southern Brazil as well as data sets from the Pampa in Argentina by measuring the pollen grain length, grain width, pore diameter and annulus width. Firstly we investigated the potential influence of chemical treatment of pollen grains on pollen grain size as well as the measurement setting for defining the boundary conditions for using Poaceae pollen grains in a palaeoecological investigation. Finally the measured pollen grain parameters were analyzed by comparison of average grain length using statistical tests. This approach reveals highly significant differences in average grain size between all grassland ecosystems. Assuming that a certain grain size range can be assigned to a certain Poaceae taxon, conclusions about differences and similarities in taxa composition can be derived. We used two methods of multivariate data analysis. One uses the pollen grain parameters directly for a Principle Component Analysis (PCA). The other is an already established method in grassland ecology which defines parameter based pollen grain types to investigate similarities between grassland ecosystems. Both approaches confirm the results of the grain length analysis. In this work we demonstrate that the method we developed has the potential to provide acquisition of so far inaccessible information on spatial and temporal patterns and dynamics of South American grasslands.     

Poaceae pollen grain size as a tool to distinguish past grasslands in South America: a new methodological approach

Despite the dominance of grasslands during the last glacial period, especially in South America, the highly uniform morphology of Poaceae pollen grains has so far allowed only very few palynological studies based on Poaceae pollen. In our study we compare two methods of distinguishing between South American grassland ecosystems based on quantitative morphology of Poaceae pollen grains. We investigated data sets from Páramo in southern Ecuador, Campos de Altitude and Campos in south-eastern and southern Brazil as well as data sets from the Pampa in Argentina by measuring the pollen grain length, grain width, pore diameter and annulus width. Firstly we investigated the potential influence of chemical treatment of pollen grains on pollen grain size as well as the measurement setting for defining the boundary conditions for using Poaceae pollen grains in a palaeoecological investigation. Finally the measured pollen grain parameters were analyzed by comparison of average grain length using statistical tests. This approach reveals highly significant differences in average grain size between all grassland ecosystems. Assuming that a certain grain size range can be assigned to a certain Poaceae taxon, conclusions about differences and similarities in taxa composition can be derived. We used two methods of multivariate data analysis. One uses the pollen grain parameters directly for a Principle Component Analysis (PCA). The other is an already established method in grassland ecology which defines parameter based pollen grain types to investigate similarities between grassland ecosystems. Both approaches confirm the results of the grain length analysis. In this work we demonstrate that the method we developed has the potential to provide acquisition of so far inaccessible information on spatial and temporal patterns and dynamics of South American grasslands.     

A new species of Spermacoce (Rubiaceae) from the Manika high plateau (Katanga; R. D. Congo)

Spermacoce manikensis , a new species from Katanga (R. D. Congo) is described and illustrated. Special attention is given to anatomy of the primary root and pollen and seed morphology. The species is mainly characterised by the geoxylic habit, large flowers arranged in two-or three-flowered inflorescences, and seeds with a basal elaiosome. It is closely related to Spermacoce stipularis , from which it differs in its more robust taproot and habit, longer leaves, larger fruits, and 10–13(-14)- aperturate pollen grains. The species is found on the Kalahari sands of the Manika high plateau in suffrutex grassland.     

The use of the mounds of Lasius fiavus in teaching some principles of ecological investigation

The mounds built by the yellow hill ant, Lasius fiavus, in British pastures have striking and consistent effects on the patterns of plant species. Winter annuals, for example, are often confined to the mounds, but many rosette perennials, though abundant in the surrounding grassland, are virtually absent from ant-hills. A comparison of the abundances of six or seven species on and off the mounds provides a field practical suitable for senior school students and undergraduates. The students discuss the sampling methods to be used, collect the data, analyse it, and discuss the factors which might influence the relative abundance of the plant species. This paper describes the practical in detail, and provides some background information about the ecology of the ants and some common grassland plants. Some suggestions for individual projects are put forward.     

Community Associations of Chalk Grassland Leafhoppers (Hemiptera: Auchenorrhyncha): Conclusions for Habitat Conservation

Due to their confinement to specific hostplants or restricted habitat types, Auchenorrhyncha have the potential to make suitable biological indicators to measure the quality of chalk grassland under different management practices for nature conservation. The Auchenorrhyncha data from a study designed to identify the factors influencing the invertebrate diversity of chalk grasslands in southern England was used to evaluate the potential use of this group of insects as biological indicators. Between 1998 and 2002 altogether 81 chalk grassland sites were sampled. Vegetation structure and composition were recorded, and Auchenorrhyncha were sampled at each site on three occasions in each of two seasons using a ‘Vortis’ suction sampler. Auchenorrhyncha assemblages were then linked to the different grassland plant communities occurring on chalk soils according to the British National Vegetation Classification (NVC). Altogether 96 Auchenorrhyncha species were recorded during the study. Using data on the frequency and dominance of species, as is commonly done for plant communities, it was possible to identify the preferential and differential species of distinct Auchenorrhyncha assemblages. Significant differences between the Auchenorrhyncha assemblages associated with the various chalk grassland plant communities of the NVC were observed down to a level of sub-communities. We conclude that data on Auchenorrhyncha assemblages can provide valuable information for the setting of conservation management priorities, where data on floristic composition alone may not be sufficient, providing additional information on aspects of vegetation structure and condition.     

Flower‐visitor networks only partially predict the function of pollen transport by bees

Mutualistic networks display distinct structural and organizational features such as nestedness, power‐law degree distribution and asymmetric dependencies. Attention is now focused on how these structural properties influence network function. Most plant‐pollinator networks are constructed using records of animals contacting flowers, which is based on the assumption that all visitors to flowers are pollinators; however, animals may visit flowers as nectar robbers, florivores, or to prey upon other visitors. To differentiate potential pollinator interactions from other interaction types, we examined individual bees that had visited flowers to detect if they carried pollen. Using these data, we constructed visitation and pollen‐transport networks for a spinifex‐dominated arid zone grassland. To determine how the structure of the visitation network reflects pollen transport, we compared the two networks using a null model approach to account for differences in network size. Differences in number of species, nestedness and connectance observed between the visitation and pollen‐transport networks were within expected ranges generated under the null model. The pollen‐transport network was more specialized, had lower interaction evenness, and fewer links compared to the visitation network. Almost half the number of species of the visitation network participated in the pollen‐transport network, and one‐third of unique visitation interactions resulted in pollen transport, highlighting that visitation does not always result in pollination. Floral visitor data indicate potential pollen transporters, but inferring pollination function from visitation networks needs to be performed cautiously as pollen transport resulted from both common and rare interactions, and depended on visitor identity. Although visitation and pollen‐transport networks are structurally similar, the function of all species cannot be predicted from the visitation network alone. Considering pollen transport in visitation networks is a simple first step towards determining pollinators from non‐pollinators. This is fundamental for understanding how network structure relates to network function.     

Evidence of spatial genetic structure in a California bunchgrass population

We investigated the scale of genetic variation of purple needlegrass (Nassella pulchra), a species commonly used in California for grassland restoration. Common garden and field data revealed evidence of genetic differentiation between two intermixed microhabitats characterized by differences in soil depth and community composition. We assessed the genetic variation within a single population using randomly amplified polymorphic DNA (RAPD) data collected from clusters of five individuals in 40 locations. We found no evidence for genetic structure at the whole population level. At smaller spatial scales, however, we found strong evidence that genetic subdivision of the population occurs at the level of the maternal neighborhood. We suggest that the interaction between widespread pollen dispersal and restricted seed dispersal may be the primary factor generating these results; panmictic pollen dispersal will make detection of genetic patterning difficult at larger spatial scales while limited seed dispersal will generate local genetic structure. As a result, the detection of population genetic structure will depend on the spatial scale of analysis. Local selection gradients related to topography and soil depth are also likely to play a role in structuring local genetic variation. Since N. pulchra is widely used in California in grassland and woodland habitat restoration, we suggest that, as a general rule, care should be exercised in transferring germplasm for the purposes of conservation when little is known about the within-population genetic subdivision of a plant species. Received: 23 December 1996 / Accepted: 20 May 1997     

Pollen and chromosome studies in Hieracium sect. Alpestria (Asteraceae)

Potential pollen viability has been investigated in British apomictic Hieracium section Alpestria and related species using Alexander's Stain. New and existing chromosome counts are given. Some species produced no pollen, others varied in the amount they produced, and a few species consistently produced reasonable quantities. The three triploid section Alpestria species produced very little or no pollen, whilst the amount produced by the tetraploids varied widely. Section Alpestria species had significantly lower pollen viability than the non‐section Alpestria species. The generally low pollen production in most species of section Alpestria is consistent with them being of hybrid origin.     

An Ecological Classification of British Butterflies: Ecological Attributes and Biotope Occupancy

We classify British butterflies using 136 non-biotope associated binary state ecological attributes describing all stages of butterfly life-cycles. Using cluster analysis we identify two groups of woodland species, a group occurring in tall open grassland, another group associated with short sward herb-rich grassland, and a ruderal group. Principal Component and Factor analyses (4 factor solution) are used to identify ecological attributes that determine species groupings. No single attribute or attribute type is responsible for the groupings, which are also insensitive to hostplant type. We use presence/absence data from Butterfly Monitoring Scheme transects in southern Britain to test our classification. On the basis of adult occurrences, similarities within two of the four groups identified from PCA are greater than between groups. Exclusivity between species pairs is also more frequent between groups than within groups. Species' ranges, distributions, biotope range, dispersal ability and recent decline in abundances differ between groups identified by their factor loadings. Ruderal species have large ranges, abundances, extensive mobilities and show little recent decline. The group associated with short sward grassland have the lowest mobilities, and the smallest distributions within their geographic ranges. True woodland species have the smallest biotope range, and the species associated with open areas have the second smallest decline in their distributions. Our ecological classification identifies characteristics of species that determine their habitat requirements and could serve to predict the response of species groups to environmental change on the basis of their ecological attributes. Our method may be of use in identifying the relative importance of ecological attributes of less-well studied taxa and be applicable in less well known geographic regions.     

Re-creating Semi-natural Communities: Effect of Sowing Rate on Establishment of Calcareous Grassland

Restoration ecologists are increasingly aware of the potential to re-create chalk grassland on abandoned farmland. Success is often hampered by lack of desirable species in the seed bank and by poor dispersal from nearby sites. In certain schemes, the input of seed may be essential. Locally collected seed is desirable but availability is limited. We examined whether lower sowing rates than currently recommended may be successfully utilized, facilitating more-efficient use of available seed. Experimental plots on former agricultural land were sown at different rates in a randomized complete block, and the vegetation was surveyed for two years. We compared species richness and cover for chalk grassland plants and weeds - species not associated with chalk grassland communities. Values for cover and abundance were matched with data for communities of the British National Vegetation Classification (NVC). Species richness for chalk grassland plants increased with sowing rate and with time, although after two years there was no significant difference between the treatments sown at 0.4, 1.0, and 4.0 grams of seed per square meter. Weed species decreased with increasing rate and time. After two seasons, the vegetation on all treatment plots was similar to that of recognized NVC chalk grassland communities, while the controls were dominated by weeds and showed signs of developing into species-poor grassland. Higher rates rapidly eliminated weeds, but even a small inoculum of seed seemed to significantly enhance establishment of desirable plants and to reduce weed cover. We conclude that lower sowing rates would enable the desired vegetation to become established successfully, under appropriate conditions and management regimes. Lower rates allow for the re-creation of sizable areas using local seed, and they minimize damage to donor sites.     

Pollen production in anemophilous species of the Poaceae family in Tetouan (NW Morocco)

Total pollen production per inflorescence and per square meter were studied in anemophilous species of the Poaceae family, to determine the relative contribution of each species to the total load of pollen released and to identify the species with the highest potential pollen emission. This was done by calculating the number of pollen grains per flower and per inflorescence and by estimating the density of inflorescences in an area of one square meter. Pollen production per inflorescence varied between 12,000 and 15 million grains, often being higher in the perennial species. Mountainous, grassland, riverside, and littoral areas contributed similarly to the amount of pollen grain production per square meter. Species such as Gaudinia fragilis, Hordeum murinum, Lolium rigidum, and Trisetaria panicea, with high pollen records, were found in all the studied areas. Aerobiological analysis will be undertaken to determine the main pollen season of Poaceae species. The results obtained are of great importance for respiratory health management in this region.     

The independent and combined effects of floral traits distinguishing two pollination ecotypes of a moth‐pollinated orchid

Identifying traits and agents of selection involved in local adaptation is important for understanding population divergence. In southern Sweden, the moth‐pollinated orchid Platanthera bifolia occurs as a woodland and a grassland ecotype that differ in dominating pollinators. The woodland ecotype is taller (expected to influence pollinator attraction) and produces flowers with longer spurs (expected to influence efficiency of pollen transfer) compared to the grassland ecotype. We examined whether plant height and spur length affect pollination and reproductive success in a woodland population, and whether effects are non‐additive, as expected for traits influencing two multiplicative components of pollen transfer. We reduced plant height and spur length to match trait values observed in the grassland ecotype and determined the effects on pollen removal, pollen receipt, and fruit production. In addition, to examine the effects of naturally occurring variation, we quantified pollinator‐mediated selection through pollen removal and seed production in the same population. Reductions of plant height and spur length decreased pollen removal, number of flowers receiving pollen, mean pollen receipt per pollinated flower, and fruit production per plant, but no significant interaction effect was detected. The selection analysis demonstrated pollinator‐mediated selection for taller plants via female fitness. However, there was no current selection mediated by pollinators on spur length, and pollen removal was not related to plant height or spur length. The results show that, although both traits are important for pollination success and female fitness in the woodland habitat, only plant height was sufficiently variable in the study population for current pollinator‐mediated selection to be detected. More generally, the results illustrate how a combination of experimental approaches can be used to identify both traits and agents of selection.     

Nectar usage in a southern Arizona hawkmoth community

Abstract.  1. Hawkmoths (Sphingidae) are important plant associates at two life-history stages: larvae are herbivorous, whereas adults are nectar feeders and often pollinators. The diversity and identities of plants used for nectar is poorly known, however.
2. This study takes a community-level approach to hawkmoth nectar plant usage in a semi-arid grassland habitat in southern Arizona, U.S.A.
3. Pollen carried on the proboscis was identified from over 700 individuals of 14 hawkmoth species attracted to lights over a 2-year period.
4. Two plant species dominated pollen loads, suggesting that hawkmoths use these species extensively as nectar sources: Datura wrightii (Solanaceae), a classic hawkmoth-pollinated plant, and Agave palmeri (Agavaceae), which is known to be used extensively by bats. Field surveys indicate that both species are relatively rare in the flowering community. Little or no pollen was present on the moths from the most common plant species in flower during the study.
5. The dominance of Agave in pollen loads suggests that this typically bat-pollinated species may be subsidising pollinator populations of the hawkmoth-pollinated flora.
6. Three groups of hawkmoths within this community are identified based on larval diets (reported in the literature) and adult diets (documented here): those that, at a given site, heavily exploit the same plant species at both life-history stages ( Manduca sexta and M. quinquemaculata ); those that have broad local associations at both life-history stages ( Hyles lineata ); and those that exhibit narrow but non-overlapping local associations at the two life-history stages (all other hawkmoths at this site).     

High dissimilarity within a multiyear annual record of pollen assemblages from a North American tallgrass prairie

Grassland vegetation varies in composition across North America and has been historically influenced by multiple biotic and abiotic drivers, including fire, herbivory, and topography. Yet, the amount of temporal and spatial variability exhibited among grassland pollen assemblages, and the influence of these biotic and abiotic drivers on pollen assemblage composition and diversity has been relatively understudied. Here, we examine 4 years of modern pollen assemblages collected from a series of 28 traps at the Konza Prairie Long‐Term Ecological Research Area in the Flint Hills of Kansas, with the aim of evaluating the influence of these drivers, as well as quantifying the amount of spatial and temporal variability in the pollen signatures of the tallgrass prairie biome. We include all terrestrial pollen taxa in our analyses while calculating four summative metrics of pollen diversity and composition – beta‐diversity, Shannon index, nonarboreal pollen percentage, and Ambrosia:Artemisia – and find different roles of fire, herbivory, and topography variables in relation to these pollen metrics. In addition, we find significant annual differences in the means of three of these metrics, particularly the year 2013 which experienced high precipitation relative to the other 3 years of data. To quantify spatial and temporal dissimilarity among the samples over the 4‐year study, we calculate pairwise squared‐chord distances (SCD). The SCD values indicate higher compositional dissimilarity across the traps (0.38 mean) among all years than within a single trap from year to year (0.31 mean), suggesting that grassland vegetation can have different pollen signatures across finely sampled space and time, and emphasizing the need for additional long‐term annual monitoring of grassland pollen.     

Pollinator-mediated competition between two congeners, Limnanthes douglasii subsp. rosea and L. alba (Limnanthaceae)

? Premise of Study: Pollinator visits are essential for reproduction in many plants, yet interspecific movements of pollinators can also lead to competitive interactions between coflowering species. Pollination-mediated reductions in fertility could potentially lead to exclusion of competing plant species, and may generate spatial variation in the associations among coflowering species across a landscape. ? Methods: I documented the potential for heterospecific pollen transfer to cause competitive interactions between two annual grassland species native to California, Limnanthes douglasii subsp. rosea and L. alba, two reproductively incompatible species that have broadly overlapping geographic ranges in the foothills of the Sierra Nevada. I observed pollinator movement in constructed arrays and controlled crosses in the greenhouse and field to investigate the consequences of heterospecific pollen transfer. ? Key Results: Pollinators move readily between species when they are presented together in experimental arrays. In the greenhouse, deposition of heterospecific pollen decreased fertility in both species. The decrease in seeds produced per flower was much more pronounced in L. d. rosea (90.6% reduction) than in L. alba (40.8% reduction). In field experiments, L. d. rosea plants that received pollen from heterospecific neighbors first showed >50% reduction in per-flower fertility. ? Conclusions: Under natural pollination conditions, heterospecific pollen transfer has the ability to decrease the fertility of L. d. rosea when it occurs at low frequency in mixed stands. Accordingly, pollinator-mediated competition may contribute to the locally disjunct distributions of these two species. It may also influence important restoration decisions in vernal pool habitats.