Staying in touch : how highly specialised moth pollinators track host plant phenology in unpredictable climates

Background: For specialised pollinators, the synchrony of plant and pollinator life history is critical to the persistence of pollinator populations. This is even more critical in nursery pollination, where pollinators are obligately dependant on female host plant flowers for oviposition sites. Epicephala moths (Gracillariidae) form highly specialised nursery pollination mutualisms with Phyllanthaceae plants. Several hundred Phyllanthaceae are estimated to be exclusively pollinated by highly specific Epicephala moths, making these mutualisms an outstanding example of plant–insect coevolution. However, there have been no studies of how Epicephala moths synchronise their activity with host plant flowering or persist through periods when flowers are absent. Such knowledge is critical to understanding the ecology and evolutionary stability of these mutualisms. We surveyed multiple populations of both Breynia oblongifolia (Phyllanthaceae) and it’s Epicephala pollinators for over two years to determine their phenology and modelled the environmental factors that underpin their interactions. Results: The abundance of flowers and fruits was highly variable and strongly linked to local rainfall and photoperiod. Unlike male flowers and fruits, female flowers were present throughout the entire year, including winter. Fruit abundance was a significant predictor of adult Epicephala activity, suggesting that eggs or early instar larvae diapause within dormant female flowers and emerge as fruits mature. Searches of overwintering female flowers confirmed that many contained pollen and diapausing pollinators. We also observed diapause in Epicephala prior to pupation, finding that 12% (9/78) of larvae emerging from fruits in the autumn entered an extended diapause for 38–48 weeks. The remaining autumn emerging larvae pupated directly without diapause, suggesting a possible bet-hedging strategy. Conclusions: Epicephala appear to use diapause at multiple stages in their lifecycle to survive variable host plant phenology. Furthermore, moth abundance was predicted by the same environmental variables as male flowers, suggesting that moths track flowering through temperature. These adaptations may thereby mitigate against unpredictability in the timing of fruiting and flowering because of variable rainfall. It remains to be seen how widespread egg diapause and pre-pupal diapause may be within Epicephala moths, and, furthermore, to what degree these traits may have facilitated the evolution of these highly diverse mutualisms

Changes in plant species abundance alter the multifunctionality and functional space of heathland ecosystems

Though it is well established that species composition affects ecosystem function, the way in which species combine to control overall ecosystem functioning is still debated. In experimental mesocosms, we planted three functionally distinct dry-heath species in varying proportions and measured multiple ecosystem properties related to nutrient cycling and carbon storage (hereafter functions). Overall ecosystem functioning was described as the main axes of variation in ecosystem functioning (functional space) and the proportion of ecosystem functions at high levels; for example, fast carbon and nutrient cycling (cluster-based multifunctionality). The first functional space axis, related to nitrogen availability, was driven by plant species abundance, particularly that of legumes, which strongly affected many individual functions. The second, related to total plant biomass and woodiness, was mostly driven by the abundance of dwarf shrubs. Similarly, cluster-based multifunctionality was related to the initial abundance of all species, but particularly the legume. Interactions between species also affected ecosystem multifunctionality, but these effects were smaller in magnitude. These results indicate that species interactions could play a secondary role to species abundance and identity in driving the overall ecosystem functioning of heathlands, but also that axes of variation in functional space are clearly linked to plant functional composition

Tree traits and microclimatic conditions determine cooling benefits of urban trees

Trees play a key role in mitigating urban heat by cooling the local environment. This study evaluated the extent to which street trees can reduce sub-canopy air temperature relative to ambient conditions (DT), and how DT relates to tree traits and microclimatic variables. Air temperature under the canopies of 10 species was recorded within residential areas in Western Sydney, Australia, during summer 2019–2020. Tree and canopy traits, namely tree height, specific leaf area, leaf dry matter content, leaf area index, crown width and the Huber value (the ratio of sapwood area to leaf area) were then measured for all species. Species differed significantly in their DT values, with peak cooling (maximum DT 3.9 C) observed between 9–10 am and sub-canopy warming (i.e., positive DT values) typically occurring during afternoon and overnight. Trees with high LAI and wider canopies were associated with the greatest daytime cooling benefits and lower levels of nighttime warming. DT was also negatively related to windspeed and vapor pressure deficit, and positively to solar irradiance. This study provides valuable information on how tree characteristics and microclimate influence potential cooling benefits that may aid planning decisions on the use of trees to mitigate heat in urban landscapes

Drivers of the microbial metabolic quotient across global grasslands

Aim: The microbial metabolic quotient (MMQ; mg CO2-C/mg MBC/h), defined as the amount of microbial CO2 respired (MR; mg CO2-C/kg soil/h) per unit of microbial biomass C (MBC; mg C/kg soil), is a key parameter for understanding the microbial regulation of the carbon (C) cycle, including soil C sequestration. Here, we experimentally tested hypotheses about the individual and interactive effects of multiple nutrient addition (nitrogen + phosphorus + potassium + micronutrients) and herbivore exclusion on MR, MBC and MMQ across 23 sites (five continents). Our sites encompassed a wide range of edaphoclimatic conditions; thus, we assessed which edaphoclimatic variables affected MMQ the most and how they interacted with our treatments. Location: Australia, Asia, Europe, North/South America. Time period: 2015–2016. Major taxa: Soil microbes. Methods: Soils were collected from plots with established experimental treatments. MR was assessed in a 5-week laboratory incubation without glucose addition, MBC via substrate-induced respiration. MMQ was calculated as MR/MBC and corrected for soil temperatures (MMQsoil). Using linear mixed effects models (LMMs) and structural equation models (SEMs), we analysed how edaphoclimatic characteristics and treatments interactively affected MMQsoil. Results: MMQsoil was higher in locations with higher mean annual temperature, lower water holding capacity and lower soil organic C concentration, but did not respond to our treatments across sites as neither MR nor MBC changed. We attributed this relative homeostasis to our treatments to the modulating influence of edaphoclimatic variables. For example, herbivore exclusion, regardless of fertilization, led to greater MMQsoil only at sites with lower soil organic C (< 1.7%). Main conclusions: Our results pinpoint the main variables related to MMQsoil across grasslands and emphasize the importance of the local edaphoclimatic conditions in controlling the response of the C cycle to anthropogenic stressors. By testing hypotheses about MMQsoil across global edaphoclimatic gradients, this work also helps to align the conflicting results of prior studies

Temporal trends in twig growth of Fagus sylvatica L. and their relationships with environmental factors

Dendrochronology and twig growth studies provide useful historical information about tree growth and permit analyses of the effects of measured environmental parameters on such growth. Recent growth histories of currently healthy and unhealthy beech {Fagus sylvatica L.) trees were reconstructed for a number of sites in southern England, by measuring annual twig extension growth back to 1960. The relationships between twig growth and a number of climatic (and one pollution) variables were then determined, using multiple regression analyses.A substantial reduction in twig growth was found in 1976 and 1977 for all trees, indicating the dramatic effect of the severe drought that was experienced throughout much of southern England during 1976. After this time, healthy trees at all sites quickly regained their pre-1976 growth rates. In contrast, unhealthy trees at two sites failed to recover and still show greatly reduced rates of growth. At two additional sites, while the effect of the 1976 drought is clear, the growth rates of unhealthy trees have been significantly lower than their healthy neighbours since before 1960. Soil type and tree age failed to provide an adequate explanation for the lower than expected growth rates of unhealthy trees seen at almost all sites.At most sites, a significant proportion of the variance in twig growth is explained by combinations of climatic variables. In most such cases, growth rates decreased as the variable concerned increased. Ozone was included in the best fit model for unhealthy trees at two sites. However, since this pollutant accounted for only a small amount of the model variance and the ozone data were of poor spatial resolution, care must be taken over the interpretation of this result

Functional relationships with N deposition differ according to stand maturity in Calluna-dominated Heathland

Plant and soil bio(chemical) indicators are increasingly used to provide information on N deposition inputs and effects in a wide range of ecosystem types. However, many factors, including climate and site management history, have the potential to influence bioindicator relationships with N due to nutrient export and changing vegetation nutrient demands. We surveyed 33 heathlands in England, along a gradient of background N deposition (7.2–24.5 kg ha−1 year−1), using Calluna vulgaris growth phase as a proxy for time since last management. Our survey confirmed soil nutrient accumulation with increasing time since management. Foliar N and phosphorus (P) concentrations in pioneer- and mature-phase vegetation significantly increased with N deposition. Significant interactions between climate and N deposition were also evident with, for example, higher foliar P concentrations in pioneer-phase vegetation at sites with higher temperatures and N deposition rates. Although oxidized N appeared more significant than reduced N, overall there were more, stronger relationships with total N deposition; suggesting efforts to control all emissions of N (i.e., both oxidized and reduced forms) will have ecological benefits

Direct and indirect effects of roads and road vehicles on the plant community composition of calcareous grasslands

Exposure of plants to vehicle exhaust emissions and road-induced changes to soil biogeochemistry and hydrology can lead to shifts in plant composition in calcareous grasslands. Mixed effects models were used to identify relationships between plant community composition and a suite of measured and modelled environmental variables along transects away from roads at eight calcareous grasslands. Ellenberg pH, moisture and nitrogen (N) scores increased nearer roadsides, however, only Ellenberg N scores were associated with their respective measured or modelled values highlighting NO2 deposition as a likely driver of change. Forb abundance and diversity increases nearer roadsides were also associated with NO2 deposition, with increases seen in the abundance and diversity of typical edge species rather than species characteristic of calcareous grasslands. Grazing, removal of invasive species and the use of barriers to intercept transport-derived air pollution may help to reduce the detrimental effects of roads across these diverse but threatened landscapes

Field-scale evaluation of effects of nitrogen deposition on the functioning of heathland ecosystems

Summary 1. Plant communities that are adapted to low levels of nutrient availability are particularly sensitive to the effects of elevated nitrogen (N) deposition. Although site-level manipulation experiments have been used widely to examine the effects of N deposition on (semi-)natural ecosystems under controlled conditions, relatively few studies have investigated field-scale impacts of elevated ambient N deposition on native plant communities and ecosystems. 2. This study investigates the relationship between atmospheric N deposition, plant and soil nutrient status and microbial enzyme activities at 32 lowland heathland sites across the UK. Sites were chosen to cover a range of N deposition rates (13.3-30.8 kg N ha−1 year−1), geographical areas and different geologies. 3. Significant relationships were found between rates of N deposition (total, reduced and oxidized) and concentrations of N and phosphorus (P) in Calluna shoots, litter and soil; relationships were generally stronger with total and reduced forms of N, compared with oxidized N. Litter and humus layers were deeper at sites receiving higher atmospheric N inputs, suggesting increased rates of soil carbon and N accumulation, despite higher levels of phenol-oxidase activity (implying faster rates of organic matter decomposition) at these sites. 4. The combination of elevated plant and soil P concentrations at sites receiving greater N inputs suggests strongly that N is increasing the availability and uptake rates of P, in addition to N. Furthermore, significant interactions between temperature and N deposition on indices relating to productivity and the turnover and uptake of nutrients highlight the influence of climate on ecosystem response to N deposition. 5. Synthesis: Field-scale evidence of changes in rates of nutrient cycling, organic matter accumulation and plant biochemistry suggests that ambient levels of N deposition are affecting the functioning of many heathland ecosystems and that the magnitude of these effects is also influenced by climate. Since such changes are known to be associated with reduced resistance to environmental stress and loss of plant diversity, current (and predicted future) levels of N deposition are likely to have important implications for the conservation and long-term sustainability of nutrient-poor ecosystems, particularly in the face of climate change

Nutrient relations and root mycorrhizal status of healthy and declining beech (Fagus sylvatica L.) in southern Britain

Many studies have been carried out to investigate the wide scale problem of forest decline that is affecting vast numbers of trees throughout Europe and North American. Evidence of nutritional disturbances, both above- and below-ground, has been found for a variety of species. Furthermore, it has been suggested that differences in above-ground tree health are often reflected below ground, particularly in the root systems of declining trees. An investigation of root vitality, soil and leaf chemistry of healthy and unhealthy beech (Fagus sylvatica L.) was carried out at a number of sites in southern Britain, covering a range of different soil types. At each site, healthy and unhealthy trees growing in very close proximity were compared in order to avoid large scale environmental differences that might be expected when comparing stands of trees at different locations. Healthy trees were found to have significantly greater proportions of live mycorrhizal roots than their unhealthy neighbours at most of the sites investigated. In addition, significant differences in soil chemistry were found between trees in contrasting health, with healthy trees generally growing in soil containing higher concentrations of calcium, magnesium and potassium and lower aluminium/calcium ratios. Very few differences were found in leaf nutrient content, suggesting that soil chemistry is not yet limiting nutrient uptake. The importance of root system differences in the tolerance of unfavourable edaphic and climatic conditions is discussed with respect to tree health.Many studies have been carried out to investigate the wide scale problem of forest decline that is affecting vast numbers of trees throughout Europe and North American. Evidence of nutritional disturbances, both above- and below-ground, has been found for a variety of species. Furthermore, it has been suggested that differences in above-ground tree health are often reflected below ground, particularly in the root systems of declining trees. An investigation of root vitality, soil and leaf chemistry of healthy and unhealthy beech (Fagus sylvatica L.) was carried out at a number of sites in southern Britain, covering a range of different soil types. At each site, healthy and unhealthy trees growing in very close proximity were compared in order to avoid large scale environmental differences that might be expected when comparing stands of trees at different locations. Healthy trees were found to have significantly greater proportions of live mycorrhizal roots than their unhealthy neighbours at most of the sites investigated. In addition, significant differences in soil chemistry were found between trees in contrasting health, with healthy trees generally growing in soil containing higher concentrations of calcium, magnesium and potassium and lower aluminum/calcium ratios. Very few differences were found in leaf nutrient content, suggesting that soil chemistry is not yet limiting nutrient uptake. The importance of root system differences in the tolerance of unfavourable edaphic and climatic conditions is discussed with respect to tree health

Effects of within-tree flowering asynchrony on the dynamics of seed and wasp production in an Australian fig species

Within-tree flowering asynchrony in figs, which may allow pollinating wasps to avoid the risks of dispersal in inclement conditions, has been predicted as a trait to be favoured in highly seasonal environments. Comparisons of such asynchronous figs with better-known species that exhibit within-tree synchrony might also be expected to reveal differences in the outcome of the conflict between pollinator wasp and fig seed production, and the dynamics of non-pollinating wasps. This paper presents data on wasp and seed production in Ficus rubiginosa Desf. ex Vent., an asynchronous species that occurs in the highly seasonal environment of south-eastern Australia. In contrast to recent studies of figs showing within-tree flowering synchrony, syconium size was the main determinant of wasp and seed production in F. rubiginosa. Non-pollinating wasps were highly prevalent but occurred in low numbers and appeared to have relatively little impact on pollinator wasp or fig seed production. Data on flower positions revealed that non-pollinating wasps occurred almost exclusively in the outer layer of flowers, while pollinators were more abundant in the inner flower layer, which may represent an area of enemy-free space. The ratio of seeds to female pollinator wasps, an index of fig sex allocation, was more seed-biased than in several New World fig species that exhibit within-tree synchrony. This last result supports the idea that within-tree fruiting asynchrony permits a degree of self-pollination in F. rubiginosa