Rapid recovery of soil arthropod assemblages after exotic plantation tree removal from hydromorphic soils in a grassland‐timber production mosaic

Many wetland systems are being lost or degraded by human activities such as plantation forestry. Therefore, efforts to restore these wetland systems are important for biodiversity recovery. We assess the recovery of arthropod assemblages that occupy hydromorphic grassland topsoil and leaf litter after the removal of exotic pine trees. We sampled arthropods in three biotopes (natural untransformed hydromorphic grasslands, restored hydromorphic grasslands, and commercial pine plantations) replicated across a large‐scale timber‐grassland mosaic in the KwaZulu‐Natal Midlands, South Africa. In the restored sites, overall species richness, as well as species richness of spiders, ants, and orthopterans, was significantly higher than in plantations, and was as high as in natural, untransformed sites. Additionally, overall assemblage structure along with spider, beetle, ant, and orthopteran assemblages showed no significant differences between restored and natural grasslands. Therefore, pine tree removal enables recovery of these arthropod taxa to levels similar to those in natural hydromorphic grassland. Recovery was rapid, with the assemblages in some restored sites resembling those in untransformed sites after only 6 years, indicating a high level of resilience and recovery in these systems. Contrary to expectations, time since pine removal had a negative effect on arthropod recovery. This was due to the strongly negative effect of alien invasive American bramble (Rubus cuneifolius), which was most common in older restored sites, causing deviation from their restoration trajectory. The potential for arthropod recovery in these hydromorphic grasslands is high, but successful restoration is dependent on ongoing appropriate grassland management, especially removal of bramble.     

Limited mate availability decreases reproductive success of fragmented populations of Linnaea borealis, a rare, clonal self-incompatible plant

Background and Aims

Small populations of rare plant species are increasingly reported to have high levels of reproductive failure. The objective of this study was to understand the principal constraints on sexual reproduction in small fragmented populations of a rare clonal self-incompatible plant.

Methods

The pollinator spectrum, diversity of flower colour, natural pollination and fruit-set levels of L. borealis were examined in Scotland. Artificially crossed seed production was compared within and between different flower colour types and patches.

Key Results

Linnaea borealis was pollinated by a diverse spectrum of insect species and the principal pollinators were muscid, syrphid and empid flies which mostly moved only small distances (<0·25 m) between flowers when foraging. Natural pollination levels were high, indicating high pollinator effectiveness, but fruit set was very low in most patches. Flower colour diversity was low in most patches and only those with a diversity of flower colour types had high fruiting success. Pollination experiments showed L. borealis to be highly self-incompatible and artificial crosses within and between patches and flower colour types confirmed that low fruit success was the result of a lack of compatible mates and limited pollen movement between them. Evidence of isolation from pollen exchange was apparent at as little as 6 m and severe at 30 m and beyond.

Conclusions

Limited mate availability and isolation from pollen exchange compromise the reproductive success of fragmented populations of L. borealis in Scotland. A diversity of compatible mates situated within close proximity (<6 m) is the key requirement to ensure high natural fruiting success. This study emphasizes that an understanding of the breeding system, pollinator spectrum and potential for interconnectivity via pollinator movement are fundamental to identify isolation distances and to establish when conservation intervention is necessary for rare species.Key words: Linnaea borealis, clonal, self-incompatible, reproductive failure, fragmented populations, isolation, pollination     

Responses of brambles,Rubus vestitus,to herbivory

We investigated the effect of fertilisation and multiple episodes of simulated herbivory on the prickliness of bramble, Rubus vestitus. The prickliness of fertilised, uncut plants was significantly greater than that of plants in all other treatments. Our results indicate that prickle production is constrained by resource availability, with brambles allocating resources to growth under intense herbivory. Isozyme electrophoresis of plants collected from Wytham Woods, Oxford, UK, failed to detect any variation, suggesting that observed variation in prickliness may be primarily the result of phenotypic plasticity. In our experiments, however, we observed significant variation in prickliness among putative genotypes.     

Impacts of water stress removal and disturbance regimes on Mediterranean dry grasslands diversity and succession

Our goal was to disentangle the effects of stress removal and disturbance on plant communities of a Mediterranean rangeland, La Crau (southeastern France). We compared undisturbed reference steppe vegetation with vegetation impacted by changes in land use such as earlier phases of cultivation (dating back 20 years) and/or current water infiltrations (revealed by the presence of Brachypodium ph?nicoïdes), since the establishment of adjacent hay meadows. We considered plots with and without brambles (Rubus ulmifolius), an indigenous shrub species that had colonized the area after the land-use changes. We monitored the composition and measured the taxonomic richness and richness of functional groups, evenness and similarities of plant communities. The species richness of the undisturbed community was significantly higher than that of all disturbed plant communities. Although cultivation led to the dominance of ruderal species, the removal of water stress had a stronger negative impact, enabling the establishment of herbaceous competitor species such as B. ph?nicoïdes. The dominance of this species resulted in a significant decline in species richness and evenness after water stress removal. The presence of brambles correlating with former cultivation and/or current water infiltration did not have a significant impact on plant species richness in the vicinity of bramble bushes, although it significantly modified the composition of the adjacent herbaceous vegetation. Our study highlights again the low resilience of Mediterranean dry grasslands after disturbance. While both the disturbance and the water stress removal resulted in changes within the plant community, our findings reveal a stronger impact of the water stress removal. Water infiltration led to decreased plant species richness and evenness because the greater availability of water favored competitor species over the stress-tolerant xeric species. Therefore, for restoring the original steppe species richness, the priority will be to control water infiltrations, even before any scrub-clearing is undertaken to control bramble colonization.     

Two alien invasive acacias in Italy: Differences and similarities in their flowering and insect visitors

In Italy, alien acacias have been introduced for ornamental and reforestation purposes, and some species became invasive occupying patches of the Mediterranean landscape. On the Island of Elba (Central Italy), Acacia dealbata and A. pycnantha form dense stands at short distance, showing an impressive massive flowering at the end of the winter/early spring. Our aim was to investigate the behaviour of the two species in relation to the flowering features, from phenology to floral characteristics, and their replay to the observed flower visitors. Differences between the two species emerged on all the parameters considered. A. pycnantha peak of flowering occurred later than A. dealbata and showed larger flower heads (FHs), more flowers/head, stamens/flower and polyads. On A. dealbata, we recorded longer racemes and more FHs/raceme, determining a more flower-dense crown. Even if contacts with flower visitors were generally low on both species, A. dealbata showed a more heterogeneous visitor assemblage. Both acacias species interacted with local generalist pollinators, as bumblebees and honey bees. Flower handling and resource collection strategy by the honey bee indicate a long-term relationship between the bee and the acacias, with bees investing longer time on the larger A. pycnantha flower heads.     

Massive floral display affects insect visits but not pollinator‐mediated pollen transfer in Rhododendron ferrugineum

Fragmentation of natural vegetation creates one of the largest threats to plant–pollinator interactions. Although fragmentation impacts on plant populations have been explored in many, mainly herbaceous, species, the response of wild mass‐flowering species is poorly known. Here, we studied 28 heathland patches dominated by the mass‐flowering shrub Rhododendron ferrugineum, each presenting different R. ferrugineum floral display sizes (total inflorescence number per patch) and patch isolation (median distance to the three nearest patches). We assessed the impacts of these two factors on (i) heathland patch visitor assemblage (considering R. ferrugineum versus surrounding community) and (ii) R. ferrugineum flower visitation rate and pollen transfer limitation (comparing seed set from emasculated to pollen‐supplemented flowers). We found that diversity and abundance of bees visiting R. ferrugineum in heathland patches significantly decreased with decreasing R. ferrugineum floral display, while overall visitor density per patch and flower visitation rate increased. Moreover, a decrease in massive floral display and increase in patch isolation resulted in reduced visitor density in the surrounding community. Even in patches with few individuals, we found disproportionate visitor abundance in R. ferrugineum compared to the surrounding community. Finally, pollen transfer limitation in R. ferrugineum was neither affected by visitation rate nor by patch attributes. By disproportionally attracting pollinators from co‐flowering species, and probably promoting geitonogamous pollen transfer, the mass‐flowering trait appears adequate to compensate, in terms of conspecific pollen transfer, for the decrease in visitor diversity and abundance and in mate availability, which usually result from population fragmentation.     

Morphological and molecular identification to secure cultivar maintenance and management of self-sterile Rubus arcticus

Background and Aims

Preservation of cultivar purity creates a particular challenge for plants that are self-incompatible, require insects for cross-pollination, and have easily germinating seeds and vigorously spreading rhizomes. As the fields must be planted with mixed populations, and a balance must be maintained between the cultivars to achieve effective pollination, methods for field monitoring of the relative density of different cultivars must be practical. Furthermore, a DNA-based method is needed for cultivar verification in the collections and outside of the growing season. The aim of this study was to develop both types of methods for Rubus arcticus (arctic bramble).

Methods

Morphological parameters were measured from six cultivars grown on three farms. Observations from the flowers and fruits included: petal and sepal number, flower diameter, arrangement of petals, size of calyx in relation to corolla, fruit weight, yield and soluble sugars. Observations from the leaves included: width and height of middle leaflet, shape of the base of terminal leaflet, shape of terminal leaflet, leaf margin serration and fingertip touch. The applicability of simple sequence repeat (SSR) or microsatellite DNA markers developed for red raspberry was tested on eight arctic bramble cultivars.

Key Results and Conclusions

Morphological and molecular identification methods were developed for R. arcticus. The best morphological characteristics were the length-to-width ratio of the middle leaflet and leaf margin serration. A particular characteristic, fingertip touch, was shown by electron microscopy to be related to the density and quality of the leaf hairs. Red raspberry SSR marker no. 126 proved to be applicable for differentiation of the eight arctic bramble cultivars tested. These identification methods are critical to secure the maintenance and management of R. arcticus. However, the challenges faced and approaches taken are equally applicable to other species with similar biology.     

501. Rubus Calophyllus

Summary. A rare bramble from the eastern Himalaya, Rubus calophyllus C.B. Clarke is illustrated. Its distribution and habitat are described and related species are discussed.     

Temporal variations in the linkages between plants and flower visitors and the pollination success of Primula modesta along the snowmelt gradient

Although pollination networks between plants and flower visitors are diverse and flexible, seed production of many plant species is restricted by pollen limitation. Obligate outcrossers often suffer from low pollinator activity or severe interspecific competition for pollinator acquisition among co-flowering species. This study focused on seasonal changes in plant–flower visitor linkages in an alpine ecosystem and examined whether and how this seasonality affected the seed-set of Primula modesta, a self-incompatible distylous herb having long-tubed flowers. First, we recorded the linkages between plants and flower visitors along the snowmelt gradient. Then, pollination experiment was conducted to estimate the degree of pollen limitation over the course of flowering season of P. modesta. Flower visitors were classified by their tongue length based on the morphological matching with P. modesta flowers. As the season progressed, plant–visitor linkages became more diverse and generalized, and the visitation frequency to P. modesta flowers increased. In the later part of the season, however, the seed set of P. modesta was significantly reduced due to severe pollen limitation, presumably because of increased competition for long-tongued pollinators among co-flowering species. The present study revealed that pollinator availability for specialist species may be restricted even when plant–visitor linkages are diverse and generalized as a whole. In the case of P. modesta, morphological matching and competition for pollinators might be the main factors explaining this discrepancy.     

Rubus canduliger, a new regional species from the Netherlands, with notes on the range structure and dynamics of brambles (Rubus, Rosaceae)

Rubus canduliger is described as a new regional bramble species belonging to the seriesDiscolores. It has been recognized in the Netherlands as a distinct species for more than 70 years but identified as the central EuropeanR. grabowskii. The latter, however, differs by its tapering prickles, regular leaves with longer petiolule of the terminal leaflet, broader, often pyramidal inflorescence, and white or pink flowers with carpels hairy at the top.R. canduliger has a well-defined range north of the Rhine river around Arnhem where it occurs in urban and other disturbed habitats. Historical events, e.g. transport of young trees in the 16th century may explain most records south of the Rhine river, 25–50 km outside the main range. The ecology and phytosociology of the new bramble are discussed. Its distribution is compared toR. trichanthus, another regional species with a similar range asR. canduliger, but occurring in more natural habitats. In pseudogamous European blackberries (Rubus sectionRubus =R. fruticosus agg.), range size is considered to be related to the age of the species. Several widespread species have disjunct distribution ranges that are suggested to be the result of long-distance dispersal by migratory birds but unintentional anthropogenic transport of seeds may be involved as well. We hypothesize that in northwestern Europe fragmentation of continuous ranges of widespread species was caused by ongoing deforestation and subsequent degradation (acidification) of originally base-rich sandy and loamy soils from the Iron Age onward. Many regional species, however, have originated and spread in the resulting man-made landscapes. A classification of range types of brambles differing in size and internal structure is presented.     

Community‐wide impacts of early season herbivory on flower visitors on tall goldenrod

1. The flower visitor community consists not only of pollinators but also of non‐pollinators, such as florivores, thieves and predators that attack flower visitors. Although there is increasing evidence that early‐season foliar herbivory influences pollinator visitation through changes in floral traits, few studies have explored indirect effects of foliar herbivory on community structure of the flower visitors. We examined how early‐season foliar herbivory influences the flower visitor community established in late season. 2. We conducted an inoculation experiment using a lacebug (Corythucha marmorata), which is a predominantly herbivorous insect attacking leaves of tall goldenrod (Solidago altissima) in Japan. 3. Flower abundance significantly decreased when damaged by the lacebug. The numbers of pollinators, florivores and thieves were positively correlated with flower abundance, whereas predators were not. In response to flower abundance, florivores decreased on damaged plants. On the other hand, thieves increased on damaged plants, and pollinators and predators did not differ between damaged and undamaged plants. 4. When effects of flower abundance were excluded, foliar herbivory still influenced florivores negatively and thieves positively. This implies that factors besides flower abundance may have affected the numbers of florivores and thieves. 5. Community composition of flower visitors on damaged plants significantly differed from undamaged plants, although overall abundance, taxonomic richness and taxonomic evenness were unaffected by foliar herbivory in the early season. It is important to recognise that only evaluating species diversity and overall abundance may fail to detect the significant consequence of early‐season herbivory on the flower visitor community.     

Non-Additive Effects of Genotypic Diversity Increase Floral Abundance and Abundance of Floral Visitors

Background

In the emerging field of community and ecosystem genetics, genetic variation and diversity in dominant plant species have been shown to play fundamental roles in maintaining biodiversity and ecosystem function. However, the importance of intraspecific genetic variation and diversity to floral abundance and pollinator visitation has received little attention.

Methodology/Principal Findings

Using an experimental common garden that manipulated genotypic diversity (the number of distinct genotypes per plot) of Solidago altissima, we document that genotypic diversity of a dominant plant can indirectly influence flower visitor abundance. Across two years, we found that 1) plant genotype explained 45% and 92% of the variation in flower visitor abundance in 2007 and 2008, respectively; and 2) plant genotypic diversity had a positive and non-additive effect on floral abundance and the abundance of flower visitors, as plots established with multiple genotypes produced 25% more flowers and received 45% more flower visits than would be expected under an additive model.

Conclusions/Significance

These results provide evidence that declines in genotypic diversity may be an important but little considered factor for understanding plant-pollinator dynamics, with implications for the global decline in pollinators due to reduced plant diversity in both agricultural and natural ecosystems.     

Pollination biology of fruit-bearing hedgerow plants and the role of flower-visiting insects in fruit-set

Background and Aims

In the UK, the flowers of fruit-bearing hedgerow plants provide a succession of pollen and nectar for flower-visiting insects for much of the year. The fruits of hedgerow plants are a source of winter food for frugivorous birds on farmland. It is unclear whether recent declines in pollinator populations are likely to threaten fruit-set and hence food supply for birds. The present study investigates the pollination biology of five common hedgerow plants: blackthorn (Prunus spinosa), hawthorn (Crataegus monogyna), dog rose (Rosa canina), bramble (Rubus fruticosus) and ivy (Hedera helix).

Methods

The requirement for insect pollination was investigated initially by excluding insects from flowers by using mesh bags and comparing immature and mature fruit-set with those of open-pollinated flowers. Those plants that showed a requirement for insect pollination were then tested to compare fruit-set under two additional pollination service scenarios: (1) reduced pollination, with insects excluded from flowers bagged for part of the flowering period, and (2) supplemental pollination, with flowers hand cross-pollinated to test for pollen limitation.

Key Results

The proportions of flowers setting fruit in blackthorn, hawthorn and ivy were significantly reduced when insects were excluded from flowers by using mesh bags, whereas fruit-set in bramble and dog rose were unaffected. Restricting the exposure of flowers to pollinators had no significant effect on fruit-set. However, blackthorn and hawthorn were found to be pollen-limited, suggesting that the pollination service was inadequate in the study area.

Conclusions

Ensuring strong populations of insect pollinators may be essential to guarantee a winter fruit supply for birds in UK hedgerows.Key words: Blackthorn, bramble, Crataegus monogyna, frugivorous birds, hawthorn, Hedera helix, hedgerows, ivy, insect pollination, Prunus spinosa, Rubus fruticosus, Rosa canina     

Relative abundance of an invasive alien plant affects native pollination processes

One major characteristic of invasive alien species is their occurrence at high abundances in their new habitat. Flowering invasive plant species that are visited by native insects and overlap with native plant species in their pollinators may facilitate or disrupt native flower visitation and fertilisation by forming large, dense populations with high numbers of flowers and copious rewards. We investigated the direction of such a proposed effect for the alien invasive Rhododendron ponticum in Irish habitats. Flower visitation, conspecific and alien pollen deposition, fruit and seed set were measured in a self-compatible native focal plant, Digitalis purpurea, and compared between field sites that contained different relative abundances of R. ponticum. Flower visitation was significantly lower at higher alien relative plant abundances than at lower abundances or in the absence of the alien. Native flowers experienced a significant decrease in conspecific pollen deposition with increasing alien abundance. Heterospecific pollen transfer was very low in all field sites but increased significantly with increasing relative R. ponticum abundance. However, lower flower visitation and lower conspecific pollen transfer did not alter reproductive success of D. purpurea. Our study shows that indirect interactions between alien and native plants for pollination can be modified by population characteristics (such as relative abundance) in a similar way as interactions among native plant species. In D. purpurea, only certain aspects of pollination and reproduction were affected by high alien abundances which is probably a result of high resilience due to a self-compatible breeding system. Native species that are more susceptible to pollen limitation are more likely to experience fitness disadvantages in habitats with high relative alien plant abundances.     

Benefits and limitations of isolated floral patches in a pollinator restoration project in Arizona

This study examined invertebrate floral visitor responses to floral richness, floral abundance, and distance between floral patches within a newly planted pollinator restoration habitat in an arid ecosystem in central Arizona, United States. We created a pollinator habitat experiment consisting of a large central garden (11‐m diameter) surrounded by concentric rings of smaller habitat patches (1‐m diameter), separated from one another by 1, 8, 13, and 21 m, respectively, and including four flowering species. We observed plant and visitor interactions via structured 10‐minute flower visitation observations over a 3‐month period. Key findings included: (1) each plant species interacted with a variety of flower visitors, but flower visitor groups differed only marginally among the plant species; (2) floral patches outside the central garden exhibited reduced quantities of floral structures; and (3) number of floral structures per patch, but not isolation of floral patches within the habitat, affected the number of visitors and visitor taxa richness. For practitioners and land managers looking to restore pollination systems in arid ecosystems with low establishment via seeding, the results of this study suggest that installing species‐rich and florally abundant patches of flowering plant species within a habitat could efficiently support plant‐pollinator interactions.     

Altitudinal variation in fertility and vegetative growth in the invasive plant Rubus alceifolius Poiret (Rosaceae), on Réunion island.

Rubus alceifolius Poiret (Rosaceae) was introduced to the island of Réunion in the southeastern Indian Ocean about 1850 and is now highly invasive. This bramble, native from southeastern Asia and Malaysia, has invaded a wide variety of habitats (lowland rainforest, mountain and submountain rainforest, Acacia heterophylla rainforest) from sea level to 1700 m. It is suspected to be monoclonal so, its remarkable success may be due in part to great phenotypic plasticity. On Réunion, bud, flower, fruit and seed production, the duration of the flowering period and the importance of the seed bank were found to be negatively correlated with elevation (50-1500 m a.s.l.). At a lowland site, fruit production in mature stands averaged between 30 and 80 fruits/m² during 1999 and 2000. No fruit set occurred above 1100 m. This fruit production pattern was similar over two years. Although the number of leaves per unit area is similar along the whole gradient studied, decrease of fruit set in upland areas might be compensated for by an increase in vegetative growth. Temperature variation is very sharp along the elevation gradient and may control the fruit and the seed production. Fruit production allows establishment of new populations all around the island via bird dissemination. Once established, R. alceifolius maintains dense patches that can grow vegetatively. Our results may be relevant for eradication programs that should take into account variation in reproductive strategy in lowland vs. highland habitats.     

Comparison of defence and performance traits between one widespread clone and native populations in a major invasive plant species

Aim

The success of invasive species in their introduced range is often assumed to result from evolutionary changes in defence and growth traits, or as a response to more favourable conditions. The latter is assumed particularly for species exhibiting low, or even no, sexual reproduction in the introduced range.

Location and Methods

Here, we compared Japanese (native range) and French (introduced range) populations of Fallopia japonica under common growth conditions in a glasshouse. We measured height, aboveground and belowground mass, stem stiffness, leaf toughness and secondary metabolites found in hydroalcoholic extracts of rhizomes of F. japonica, as well as the competitive response of Rubus caesius (a co‐occurring native species in the invaded range) in the presence of F. japonica from both ranges.

Results

Aboveground biomass, height, stem stiffness and composition of secondary metabolites were not significantly different between the two ranges, showing that increased aboveground vigour observed in situ in France is probably the result of a plastic response following the release of abiotic or biotic constraints from the native range. On the other hand, belowground mass, effect on R. caesius, and leaf toughness were all higher in French populations, suggesting increases in competitive ability and defence mechanisms. These differences between France and Japan may be explained either by post‐introduction evolution or by the introduction in Europe, in nineteenth century, of an exceptionally vigorous clone (pre‐adaptation).

Main conclusions

Our results provide evidence that the high vigour of this major invasive species in its introduced range is probably due to both a response to more favourable conditions and rapid evolution.

     

Identification and characterization of flowering genes in kiwifruit: sequence conservation and role in kiwifruit flower development

Background  

Flower development in kiwifruit (Actinidia spp.) is initiated in the first growing season, when undifferentiated primordia are established in latent shoot buds. These primordia can differentiate into flowers in the second growing season, after the winter dormancy period and upon accumulation of adequate winter chilling. Kiwifruit is an important horticultural crop, yet little is known about the molecular regulation of flower development.