Succession of bee communities on fallows

Wild bee communities were studied on one- to five-year-old set-aside fields with naturally developed vegetation (n = 20). and old orchard meadows (n = 4) to analyse effects of secondary succession on species diversity, resource use and associated life history traits. General theory predicts a steady increase of species richness with age of succession. In contrast, we found a first maximum in species richness of bees on two-year-old set-aside fields and a second on old meadows. Successional changes of bee communities were related to changes of vegetation. The transition from pioneer successional stages, dominated by annuals, to early successional stages, dominated by perennials, resulted in the highest species richness of flowering plants in the second year within the first five years of succession. Species richness of flowering plants was the best predictor variable for species richness of bees, whereas the cover of flowering plants correlated with the abundance of bees. Annual plants were visited more often and perennials less often than expected from their flower cover. Halictidae tended to prefer flowers of annuals, whereas Megachilidae. Apidae and Anthophoridae significantly preferred perennials. In departure from successional theory, body size, proportion of specialised bees and proportion of parasitic bees did not significantly increase with successional age, but number of generations and the proportion of soil-nesting bees decreased with successional age. Comparison of different management types showed that set-aside fields with naturally developed vegetation supported much more specialised and endangered bee species than set-aside fields sown with Phacelia tanacetifolia.     

Pollen dispersal and genetic variation in an early‐successional forest herb in a peri‐urban forest

In order to conserve forest plant species under the particularly high constraints that represent urban surroundings, it is necessary to identify the key factors for population persistence. This study examined within‐ and between‐population pollen dispersal using fluorescent dye as pollen analogue, and genetic variation and structure using 15 allozyme loci in Centaurium erythraea, an insect‐pollinated, early‐successional forest biennial herb occurring in a peri‐urban forest (Brussels urban zone, Belgium). Dye dispersal showed an exponential decay distribution, with most dye transfers occurring at short distances (<15 m), and only a few long‐distance events (up to 743 m). Flowers of C. erythraea are mainly visited by Syrphids (Diptera) and small bees, which are usually considered as short‐distance pollen dispersers, and occasionally by bumblebees, which are usually longer‐distance pollen dispersers. Small and large dye source populations differed in dye deposition patterns. The populations showed low genetic diversity, high inbreeding coefficients (F_IS) and high genetic differentiation (F_ST), suggesting restricted gene flow, which can be expected for an early‐successional biennial species with a predominantly selfing breeding system and fluctuating population sizes. The positive relationship between recruitment rate and allelic richness and expected heterozygosity, and the absence of significant correlations between genetic variation and population size suggest seedling recruitment from the seed bank, contributing to maintain genetic diversity. Long‐distance dye dispersal events indicate pollinator movements along urban forest path and road verges. These landscape elements might therefore have a potential conservation value by contributing to connectivity of early‐successional species populations located in patchy open habitats.     

Effects of vegetation, corridor width and regional land use on early successional birds on powerline corridors

Powerline rights-of-way (ROWs) often provide habitat for early successional bird species that have suffered long-term population declines in eastern North America. To determine how the abundance of shrubland birds varies with habitat within ROW corridors and with land use patterns surrounding corridors, we ran Poisson regression models on data from 93 plots on ROWs and compared regression coefficients. We also determined nest success rates on a 1-km stretch of ROW. Seven species of shrubland birds were common in powerline corridors. However, the nest success rates for prairie warbler (Dendroica discolor) and field sparrow (Spizella pusilla) were <21%, which is too low to compensate for estimated annual mortality. Some shrubland bird species were more abundant on narrower ROWs or at sites with lower vegetation or particular types of vegetation, indicating that vegetation management could be refined to favor species of high conservation priority. Also, several species were more abundant in ROWs traversing unfragmented forest than those near residential areas or farmland, indicating that corridors in heavily forested regions may provide better habitat for these species. In the area where we monitored nests, brood parasitism by brown-headed cowbirds (Molothrus ater) occurred more frequently close to a residential area. Although ROWs support dense populations of shrubland birds, those in more heavily developed landscapes may constitute sink habitat. ROWs in extensive forests may contribute more to sustaining populations of early successional birds, and thus may be the best targets for habitat management.     

Seed Bank and Vegetation Development of Sandy Grasslands After Goose Breeding

Four hypotheses were tested using long-term observations of vegetation development (12 years) and present-day seed bank data in a sandy grassland area overgrazed by domestic geese: i) Gap regeneration is crucial in maintaining species richness; thus, closed vegetation of the lower sites prevents continuous establishment of short-lived species. ii) Short-lived, early successional species comprise most of the seed banks and late successional perennials have at most sparse seed banks. iii) Composition of seed banks is more similar to pioneer vegetation than to later successional stages. iv) The similarity is higher between vegetation and seed banks in the upper-positioned plots than in the closed, lower-positioned ones. Two sites, located in the upper part of dune slopes, and another two, positioned on the lower part, were studied. In each site five 2?×?2 m permanent plots were surveyed between 1991 and 2002. Percentage cover was estimated three times a year. In the last study year, soil seed banks were sampled. Two vertical segments (0–5, 5–10 cm) were separately analyzed. The seedling emergence method was applied on concentrated samples. We found that the vegetation developed from open, annual dominated weedy assemblages to grasslands dominated by perennial graminoids. In the lower-positioned sites perennial clonal grasses (Cynodon dactylon, Poa angustifolia and P. pratensis) formed more closed vegetation, which was accompanied by lower species richness compared to the upper-positioned sites. Seed density varied between 10,300 and 40,900 seeds/m². Significantly higher seed densities were found in upper sites than in the lower ones. Annuals and short-lived perennial dicots comprised most of the seed bank. The dominant perennial graminoids also built up dense seed banks. We found a low to medium similarity between vegetation and the seed bank; similarity was the highest with the vegetation of the 1994–1998 period. In the upper sites the similarity between seed bank and the vegetation of the last studied years was also high. The vertical position had a significant effect on regeneration after overgrazing. The large cover of grasses in lower sites decreased species richness and it also decreased the seed density preventing the seed bank formation of annuals and short-lived perennials. Here, further management practices are needed to increase the species richness.     

Influence of late season fire on early successional vegetation of an Oklahoma prairie

Abstract. The study of vegetation dynamics in tallgrass prairie in response to fire has focused on dormant season fire in late successional prairies. Our objective was to determine if late season fire of varying frequency results in divergent successional patterns in an early successional tallgrass prairie disturbed by grazing and cultivation. Specifically, we evaluated the influence of late‐summer fires of varying frequency on community composition and species richness. We collected vegetation and environmental data on two sites burned in the late growing‐season at varying frequencies. These communities differed in composition depending primarily on edaphic factors, time since the last burn, and year‐to‐year variation. We interpret the time effect as related to changes in species composition accompanying plant succession that followed disturbance either from cropping and heavy grazing on the loamy site or heavy grazing on the shallow site. Other unidentified factors also have a role in vegetation dynamics on this prairie. Community composition and species richness were not consistently responsive to frequency of growing‐season fires.     

Mycobionts of <Emphasis Type="Italic">Salix herbacea</Emphasis> on a glacier forefront in the Austrian Alps

Dwarf willows (e.g. Salix herbacea) are among the earliest ectomycorrhizal (EM) plants colonising primary successional sites such as glacier forefronts in the Tyrolean Alps. EM of S. herbacea were sampled at the Rotmoos glacier forefront (Otz Valley, Austria) three times a year during the growing season and once a year during winter when plants were covered with snow in 2005 and 2006. EM were investigated using morphological methods and by sequencing the rDNA ITS region. The degree of EM mycorrhization was high throughout both years (93%). We distinguished 21 EM morphotypes and identified 19 fungal species. Cenococcum geophilum, Sebacina spp., Tomentella spp. and Cortinarius spp. dominated the mycobiont community of S. herbacea. The observed species richness in this about 150-year-old soil was at least 59% of the estimated species richness. Fungal communities differed significantly between consecutive years, and spatial heterogeneity was high. These differences made it difficult to detect seasonal impacts. Abundances of C. geophilum EM increased throughout the 2-year sampling period. Sebacina incrustans EM were very abundant in 2005, but nearly disappeared in 2006, whilst its fruitbodies were still frequent in the sampling area. This suggests that the mycorrhizae were displaced from the roots by an outcompeting species, whereas the mycelium was still present in the soil.     

The genetic diversity of rhizobia associated with Dalea purpurea Vent. in fragmented grasslands of west-central Minnesota

The increase in human population and the spread of agriculture over the past 150 years have transformed the landscape in west-central Minnesota into a mosaic of agricultural fields and urban land, leaving only remnants of the once dominant prairie ecosystem. Limited natural habitat in this fragmented landscape threatens the diversity and abundance of native legumes and could impact the size and function of associated belowground microbial populations. In this study, BOXA1R PCR and 16S rRNA gene sequence analyses were used to assess the genetic diversity of rhizobia associated with Dalea purpurea (Vent.) in nine prairie remnants ranging in size from 0.04 to 3.5 ha. The variation in soil properties was also determined. While 53 different genotypes of rhizobia were identified, four of these accounted for 84% of the 1029 rhizobia characterized using BOXA1R PCR. Representatives from three of the four dominant genotypes had a 16S rRNA gene sequence similar to that of Rhizobium gallicum, with two of these genotypes recovered at all sites. The fourth genotype was similar to that of Rhizobium etli and occurred with frequency at only two sites. Rhizobium genotype richness and site area were positively correlated. The implications of these results are discussed.     

Effects of population succession on demographic and genetic processes: predictions and tests in the daylily Hemerocallis thunbergii (Liliaceae)

Spatial genetic structure within plant populations is influenced by variation in demographic processes through space and time, including a population's successional status. To determine how demographic structure and fine-scale genetic structure (FSGS) change with stages in a population's successional history, we studied Hemerocallis thunbergii (Liliaceae), a nocturnal flowering and hawkmoth-pollinated herbaceous perennial with rapid population turnover dynamics. We examined nine populations assigned to three successive stages of population succession: expansion, maturation, and senescence. We developed stage-specific expectations for within-population demographic and genetic structure, and then for each population quantified the spatial aggregation of individuals and genotypes using spatial autocorrelation methods (nonaccumulative O-ring and kinship statistics, respectively), and at the landscape level measured inbreeding and genetic structure using Wright's F-statistics. Analyses using the O-ring statistic revealed significant aggregation of individuals at short spatial scales in expanding and senescing populations, in particular, which may reflect restricted seed dispersal around maternal individuals combined with relatively low local population densities at these stages. Significant FSGS was found for three of four expanding, no mature, and only one senescing population, a pattern generally consistent with expectations of successional processes. Although allozyme genetic diversity was high within populations (mean %P = 78.9 and H(E) = 0.281), landscape-level differentiation among sites was also high (F(ST) = 0.166) and all populations exhibited a significant deficit of heterozygotes relative to Hardy-Weinberg expectations (range F = 0.201-0.424, mean F(IS) = 0.321). Within populations, F was not correlated with the degree of FSGS, thus suggesting inbreeding due primarily to selfing as opposed to mating among close relatives in spatially structured populations. Our results demonstrate considerable variation in the spatial distribution of individuals and patterns and magnitude of FSGS in H. thunbergii populations across the landscape. This variation is generally consistent with succession-stage-specific differences in ecological processes operating within these populations.     

Clonal Diversity in Introduced Populations of An Asian Sea Anemone in North America

Previous reports hypothesized that introduced populations of the Asian sea anemone Diadumene lineata (Verill, 1870 (Presented 1869) Communications of the Essex Institution 6: 51–104), which reproduces by fission, are often monoclonal or to be composed of few clones. To test this hypothesis, sea anemones were collected from thirteen sites in three non-native regions and one native region: Chesapeake Bay, New England, central California, and Japan. The internal transcribed spacer (ITS) region separating nuclear ribosomal RNA genes was amplified from each individual using PCR and surveyed for DNA sequence variation using single strand conformational polymorphism analysis (SSCP). Fifty-six distinct electrophoretic banding patterns were found in 268 anemones, and each pattern was considered a different genotype. The number of genotypes in a population ranged from one to thirteen. Only one sample (York River, Chesapeake Bay, n = 10) was monoclonal, although six populations were dominated (>50%) by single genotype. Only four genotypes were found in more than one population, and these were confined to single regions. Walker Creek, California was sampled in 1995 and 1997 and no genotypes were found in both years, suggesting rapid shifts in genotype frequency. We conclude that multiple genotypes of D. lineata have invaded North America and that the primary importance of clonal growth for introduced populations is the production of colonizing propagules.     

Small seed bank in grasslands and tree plantations in former grassland sites in the South Brazilian highlands

The soil seed bank can be an important source for vegetation regeneration, and data on the similarity between aboveground vegetation and the seed bank can provide information about successional pathways after disturbances or land-use change. We conducted this study in natural grasslands in the subtropical highland region in southern Brazil. We evaluated the effect of silviculture on richness, density, and composition of the seed bank at former grassland sites converted to pine plantations 25 years ago. We worked at six grassland sites and three pine plantation sites and used the seedling emergence method. Seed bank density and richness in grasslands were lower than those reported in similar environments in other regions. Species richness and density varied considerably within each vegetation type; therefore, richness and density were not statistically significant, while composition varied among vegetation types. In terms of species, the pine plantation seed bank was a small subset of the grassland seed bank. Seeds of typical grassland species were missing in the pine plantation, but also had only low abundances in the grassland, and similarity of seed bank and vegetation were low (less than 20%). The low seed density found in this study, including in grasslands areas, indicates that regeneration of species from the soil seed bank likely is of a limited role for the maintenance of plant populations after disturbances in this system. Our data further suggest that natural regeneration after tree planting in grasslands is reduced due to seed limitation.     

Colonization of a host tree by herbivorous insects under a changing climate

Climate warming has been predicted to increase the abundance of herbivorous insects. Together with concurrent poleward shifts in many insect species this may increase herbivore pressure on plants. However, the manner in which plants at higher latitudes become colonized by herbivorous insects in the future is unknown. We established a translocation experiment using 26 micropropagated silver birch Betula pendula genotypes from six populations originating from 60°N to 67°N, to study the susceptibility of the translocated birches to local herbivores. The birches were planted at three different latitudes in Finland (60°N, 62°N and 67°N). We studied the effect of source population and latitudinal translocation on herbivore density, species richness, and community composition among the genotypes growing in the same environmental conditions in two years; 2011 and 2012. The source population explained the variation in the herbivore density only in 2012, whereas latitudinal translocation did not affect herbivore density. Variation in species richness was not explained by the source population or by the latitudinal translocation. At two of the study sites, the similarity of the herbivore communities among the populations decreased with increasing latitudinal distance of the source populations, possibly because birch populations that grow geographically closer to each other are genetically more similar, and therefore support a more similar composition of the arthropod community. All birch genotypes were colonized by local herbivores, suggesting that as herbivores shift their ranges polewards, they are able to colonize novel host‐plant genotypes. This enables compositional changes in insect communities on their host plants in the future, which in turn, might affect total herbivory and eventually, plant growth.     

Genetic diversity and population structure in the Barrens Topminnow (<Emphasis Type="Italic">Fundulus julisia</Emphasis>): implications for conservation and management of a critically endangered species

The Barrens Topminnow (Fundulus julisia) has undergone a rapid and dramatic decline. In the 1980s, at least twenty localities with Barrens Topminnows were known to exist in the Barrens Plateau region of middle Tennessee; currently only three areas with natural (not stocked) populations remain. The long-term survival of the Barrens Topminnow will depend entirely on effective management and conservation efforts. Captive propagation and stocking of captive-reared juveniles to suitable habitats have successfully established a handful of self-sustaining populations. However, very little is known about the genetic composition of source and introduced populations including levels of genetic diversity and structuring of genetic variation. Here we use both mitochondrial sequence data and genotypes from 14 microsatellite loci to examine patterns of genetic variation among ten sites, including all sites with natural populations and a subset of sites with introduced (stocked) populations of this species. Mitochondrial sequence analysis reveals extremely low levels of variation within populations and fixed differences between drainages. Microsatellite genotype data shows higher levels of genetic variability and a molecular signature consistent with a recent history of population bottlenecks. Measures of genetic diversity at microsatellite loci including allelic richness are similar within source and introduced populations. Bayesian assignment tests and analysis of molecular variation (AMOVA) support two distinct populations, consistent with drainage boundaries. Results from AMOVA analysis also suggest low levels of genetic connectivity between isolated populations within the same drainage. Here we propose two distinct evolutionary significant units (ESUs) and two management units that reflect this population substructure and warrant consideration in future management efforts.     

Genetic uniformity characterizes the invasive spread of water hyacinth (Eichhornia crassipes), a clonal aquatic plant

Aquatic plant invasions are often associated with long‐distance dispersal of vegetative propagules and prolific clonal reproduction. These reproductive features combined with genetic bottlenecks have the potential to severely limit genetic diversity in invasive populations. To investigate this question we conducted a global scale population genetic survey using amplified fragment length polymorphism markers of the world’s most successful aquatic plant invader –Eichhornia crassipes (water hyacinth). We sampled 1140 ramets from 54 populations from the native (South America) and introduced range (Asia, Africa, Europe, North America, Central America and the Caribbean). Although we detected 49 clones, introduced populations exhibited very low genetic diversity and little differentiation compared with those from the native range, and ～80% of introduced populations were composed of a single clone. A widespread clone (‘W’) detected in two Peruvian populations accounted for 70.9% of the individuals sampled and dominated in 74.5% of the introduced populations. However, samples from Bangladesh and Indonesia were composed of different genotypes, implicating multiple introductions to the introduced range. Nine of 47 introduced populations contained clonal diversity suggesting that sexual recruitment occurs in some invasive sites where environmental conditions favour seedling establishment. The global patterns of genetic diversity in E. crassipes likely result from severe genetic bottlenecks during colonization and prolific clonal propagation. The prevalence of the ‘W’ genotype throughout the invasive range may be explained by stochastic sampling, or possibly because of pre‐adaptation of the ‘W’ genotype to tolerate low temperatures.     

Characteristics of Golden‐winged Warbler territories in plant communities associated with regenerating forest and abandoned agricultural fields

In the Appalachian portion of their breeding range, Golden‐winged Warblers (Vermivora chrysoptera) nest in shrubland and regenerating forest communities created and maintained by disturbance. Because populations of Golden‐winged Warblers have exhibited precipitous declines in population throughout their Appalachian breeding range, management activities that create or maintain early successional habitat are a priority for many natural resource agencies and their conservation partners. Within these early successional habitats, however, additional information is still needed concerning the relative importance of different vegetation features in selection of breeding territories by Golden‐winged Warblers. Our objective, therefore, was to use logistic regression to estimate the probability of territory‐level occupancy by Golden‐winged Warblers in north‐central Pennsylvania at two sites, each with its own early successional community, based on vegetation characteristics. Our communities were composed of shrublands and regenerating forest sites resulting from two disturbances: agriculture and forest fire. Despite differences in vegetation structure, portions of both study areas (regenerating forest and old field) supported territorial Golden‐winged Warblers. Probability of territory occupancy by Golden‐winged Warblers increased with percent blackberry (Rubus) cover in the regenerating forest community, and decreased as basal area and distance to microedge increased (i.e., as vegetation patchiness decreased) in both communities. These habitat features have also been found to influence other aspects of Golden‐winged Warbler breeding ecology such as nest‐site selection, pairing success, and territory abundance. Vegetation features influencing Golden‐winged Warbler territory establishment can differ among shrubland and regenerating forest communities, and management decisions and outcomes may be affected by these differences. Our study provides a starting point for a more comprehensive hypothesis‐driven occupancy survey to investigate features of the territories of Golden‐winged Warblers across a broader geographic range and in different vegetation communities.     

Effects of disturbance intensity and frequency on early old‐field succession

Abstract. Early old‐field succession provides a model system for examining vegetation response to disturbance frequency and intensity within a manageable time scale. Disturbance frequency and intensity can interact with colonization and competition to influence relative abundance of earlier and later successional species and determine, respectively, how often and how far succession can be reset. We tested the joint effects of disturbance frequency and intensity on vegetation response (species richness, abundance, canopy structure) during the first six years of succession by clipping the dominant species (D) or all species (T) in spring and fall of each year (S), once per year in summer (Y1), each two years in summer (Y2), or each four years in summer (Y4). Vegetation response reflected disturbance effects on expansion of a later monospecific dominant perennial herb, Solidago altissima, and persistence of the early, richer flora of annuals. A more abundant and taller top Solidago canopy developed on plots clipped each 2 yr or less frequently. Plots clipped yearly or seasonally were richer, but had less abundant, shorter, and differently stratified canopy. Disturbance mediated the relative abundance of early and later successional species; however, frequency and intensity effects were not completely congruent. Persistence of a richer early successional flora increased through the most frequent disturbance (S), and was magnified by disturbance intensity. Disturbance as extreme as clipping all vegetation twice yearly did not cause a drop in species richness, but maintained the early successional community over the first six years of succession. We conclude that clipping disturbance influenced the rate of succession, but the early community could rebound through the range of disturbance frequency and intensity tested.     

Soil seed bank dynamics in post-fire heathland succession in south-eastern Australia

Soil seed banks can exert a strong influence on the path of vegetation succession following fire, with species varying in their capacity to persist in the seed bank over time, leading to changes in seed bank composition and propagules available for post-fire colonisation. This study examined the effect of time since fire on soil seed bank dynamics in a chronosequence of seven sites spanning 26 years in a south-eastern Australian sand heathland. No significant change was evident in the species richness and density of the germinable soil seed bank, but species composition differed significantly among young (0–6 years since fire), intermediate (10–17 years since fire) and old-aged (24–26 years since fire) sites (using presence/absence data). No significant trend was observed in the similarity between the extant vegetation and the soil seed bank with time since fire. A total of 32% of the species recorded in the soil seed bank were not present in the above-ground vegetation at the same site, which suggests that species requiring fire for germination may be present in the seed bank. Most species present in the extant vegetation were not recorded (63%) or were in very low abundances in the soil seed bank (29%). The mode of regeneration appears to be the major determinant of species absence in the soil seed bank, as 66% of species occurring in the extant vegetation but not in the seed bank have the capacity to regenerate by resprouting. This study shows that a major shift in the successional pathway after fire due to altered seed bank composition is unlikely in this vegetation; most species not recorded in the seed bank are either resprouters (obligate or facultative) or serotinous, suggesting that they will readily regenerate following fire. Unless fire frequencies are high and kill fire-sensitive obligate seeders before they reach maturity, the chance that the soil seed bank could substantially alter vegetation composition within the study area after fire is low. However, it is unclear how successional pathways may alter in response to severe fires with the potential to kill both seeders and resprouters.     

Are plant populations seed‐limited? A review of seed sowing experiments

We define seed limitation to be an increase in population size following seed addition. Here, we briefly consider how theoretical models deal with seed limitation and how seed sowing experiments can be used to unravel the extent of seed limitation in natural systems. We review two types of seed addition experiments: seed augmentation studies where seeds are added to existing populations; and seed introductions where seeds are sown in unoccupied sites. Overall, approximately 50% of seed augmentation experiments show evidence of seed limitation. These studies show that seed limitation tends to occur more commonly in early successional habitats and in early successional species. Most of the studies have concentrated on simply categorising populations as seed- or microsite-limited, but we believe that seed sowing experiments could be used to reveal much more about community structure, and we discuss possible future directions. In 53% of introduction studies (where seeds were sown at sites from which the species was known to be absent) the introduced species was recorded in at least one of the experimental sites following sowing. However, of the subset of studies where both seedlings and adult plants were recorded, 64% of sites contained seedlings while only 23% contained adults. This implies that, for many species, conditions for establishment are more stringent than conditions for germination. The successful establishment of plants in unoccupied patches indicates the potential for immigration to enhance local diversity (the spatial mass effect). Few studies continued monitoring for long enough to determine whether or not self-sustaining populations were successfully established, and no study attempted to link introduction sites to a putative natural source of propagules, or considered the dynamics of the metapopulation as a whole.     

Variation and covariation among life-history traits in Rumex acetosella from a successional old-field gradient

In this study morphological variation and the potential for competition to affect biomass and seedling selection of the families of five populations of Rumex acetosella L. sampled along a successional old-field gradient have been investigated. Seeds from 25 families were submitted to four competitive regimes: no competition (one plant per pot), medium competition (two plants/ pot taking plants from the same population), high within-population competition (four individuals from the same population in a pot) and high between-population competition (four individuals from two different populations in a pot). Eight traits were analysed after 3 months of growth for variation among families within populations. A significant difference among families within the two older populations was recorded for sexual biomass and related components. High sensitivity of these traits to density was observed in all populations except the youngest, suggesting specialization to particular environmental conditions in late successional populations, and a good adaptive capacity to buffer environmental variation in the pioneer population. Little significant interaction between competitive regimes and families within populations was found, i.e. genotypes within each population showed little variation in their response to environmental variation. Genotypic variance decreased with increasing competitive conditions for the majority of the traits. However, the percentage of variance in sexual reproduction explained by family was stable among treatments. Tradeoffs between vegetative reproduction and sexual reproduction were recorded at the population level along the successional gradient, with increasing competitive conditions. As succession proceeds, we observed a decrease in sexual reproduction and an increase in vegetative reproduction. At the family level, correlation among traits were similar when plants were grown in the absence of competition and at high density, with a significant negative correlation between sexual reproduction and vegetative reproduction. For both sprout number and sexual biomass, the performance of families grown under all the treatments was positively correlated. Together these results indicate allocational constraints on the reproductive biology of R. acetosella that may be favoured by natural selection and have influenced population differentiation along the successional gradient. However, they also revealed that the potential exists for evolutionary specialization through plasticity, in response to variation in environmental conditions.     

Seed dispersal by wind,birds, and bats between Philippine montane rainforest and successional vegetation

In the moist Neotropics, vertebrate frugivores have a much greater role in the dispersal of forest and successional woody plants than wind, and bats rather than birds play the dominant role in dispersing early successional species. I investigated whether these patterns also occurred in a Philippine montane rainforest and adjacent successional vegetation. I also asked whether seed mass was related to probability of dispersal between habitats. A greater number of woody species and stems in the forest produced vertebrate-dispersed seeds than wind-dispersed seeds. Although input of forest seeds into the successional area was dominated by vertebrate-dispersed seeds in terms of species richness, wind-dispersed seeds landed in densities 15 times higher. Frugivorous birds dispersed more forest seeds and species into the successional area than bats, and more successional seeds and species into the forest. As expected, seed input declined with distance from source habitat. Low input of forest seeds into the successional area at the farthest distance sampled, 40 m from forest edge, particularly for vertebrate-dispersed seeds, suggests very limited dispersal out of forest even into a habitat in which woody successional vegetation provides perches and fruit resources. For species of vertebrate-dispersed successional seeds, probability of dispersal into forest declined significantly with seed mass.     

Plants and insects in early old-field succession: comparison of an English site and an American site

The plant and insect communities of early, secondary succession beginning with bare ground in an Old World site (southern Britain) and a New World site (Iowa, U.S.A.) shared a number of characteristics. Both sites showed similar temporal patterns of plant species cover and species richness, although overall richness was greater at the Old World site. Annuals dominated at both sites during the first year of succession and were largely replaced by perennials in the second year. Monocotyledons were more abundant at the Old World site, especially in the second year. The two sites differed markedly in the contribution of native and introduced plant species, with the Old World site dominated by natives and the New World site by alien plant species. Insect herbivore load was greater at the Old World site, when expressed in terms of structural complexity of the vegetation, suggesting that there may be major differences in the influence of herbivores on the direction and rate of succession at the two sites.