Relationship between nectar production and seasonal patterns of density and nesting of resident honeyeaters in heathland near Sydney

Some individual honeyeaters were repeatedly seen near the same location over at least a 2 day period and were considered resident in the present study. Such resident honeyeaters (mostly New Holland and White-cheeked Honeyeaters) were present on the heathland study areas from about February until about October. Production of nectar energy is negligible prior to April and low after October. It is hypothesized that the density of residents is determined by their ability to obtain energy in nearby habitats while establishing nesting sites in the heathland in February and March. Nesting tended to occur between April and July when there was sufficient production of nectar-energy.     

Seasonal pattern of abundance of honeyeaters and their resources in heathland areas near Sydney

Abundances of honeyeaters, flowers and flying insects, the daily nectar-energy production per flower and the average size of flying insects were estimated every three weeks for circles of radius 20 m located in three heathland areas near Sydney. Seasonal fluctuations in honeyeater density showed no apparent relationship with seasonal fluctuations in nectar-energy productivity or in biomass of flying insects. Variation between circles in honeyeater density was also unrelated to spatial variation in energy productivity and insect biomass. The relatively low incidence of nectar-feeding and high incidence of flying exhibited by birds observed during troughs in nectar-energy production suggest that many of these birds are transient and that their density may consequently be unrelated to local conditions.     

Banksia nectar and pollen: Dietary items affecting the abundance of the common blossom bat,Syconycteris australis,in southeastern Australia

Abstract The relationship between the abundance of the common blossom bat (Syconycteris australis), a nectar/pollen specialist, and the availability of its food sources was investigated across nine Banksia integrifolia heathland sites in coastal New South Wales, Australia. An index of bat abundance was more highly correlated with direct estimates of food density (productive inflorescences) than indirect estimates of food (Banksia trees and total inflorescences). Furthermore, standing crops of nectar-energy per hectare were better predictors of bat abundance than pollen-nitrogen per hectare. Adult male mass did not differ significantly across sites of differing food availability, suggesting that the corresponding increase in bat abundance with food reduces the amount of fat an individual can deposit. Although males and females responded similarly to food, adult numbers were more strongly correlated with food supply and they were more likely to be recaptured than juveniles. Energy requirements of blossom bats exceeded availability in four out of nine sites. Flying foxes were heard feeding in large numbers at the remaining five sites, where energy was superabundant for S. australis. In contrast to the conclusions of previous studies of megabats, these results provide the basis for a prediction that nectar-energy, rather than pollen-nitrogen, is the primary limiting factor for populations of the blossom bat.     

Use of heathland and adjoining forest by honeyeaters: Results of a radiotracking study

Abstract New Holland (Phylidonyris novaehollandiae) and White-cheeked (Phylidonyris nigra) Honeyeaters that are resident in heathland during February and March, when there is negligible nectar production in that habitat, are expected to forage for nectar at that time in the adjacent open forest habitat. As nectar production in the heathland increases from then to June, while that in the forest decreases, the proportion of time spent by these birds in the forest should decrease over the same period. Radiotracking revealed that during February the residents did, as expected, spend time feeding on nectar in the forest and more time was spent in the forest in February than in April or June. However, the residents spent time away from the heathland throughout the year and spent more time away in June than in April. The resident honeyeaters spent about 80% of their time on our heathland grid overall. Radiotracking also indicated that there was no sharp distinction in behaviour between residents and other birds because birds that had been resident in the past but were not considered to be so at the time (absent residents), birds that would later be considered resident (future residents) and some birds that were never resident, all spent about half their time on the grid. The area that included 90% of heathland locations, estimated by radiotracking, for P. novaehollandiae and P. nigra averaged 0.48 ha per bird.     

Nectar depletion and its implications for honeyeaters in heathland near Sydney

Several researchers have attempted to calculate whether depression of nectar resources by Australian honeyeaters is likely to limit their densities. Such calculations can be misleading, however, and do not directly test whether birds depress nectar availability. I monitored changes in nectar availability during the 8–9 months that honeyeaters bred in heathland near Sydney, and caged inflorescences to test whether nectar availability was being depressed by birds. There were pronounced seasonal changes in nectar availability in each of 2 years, and caging substantially increased the amounts of nectar in inflorescences during months when nectar production was low. The effects of caging must have resulted from exclusion of honeyeaters, as: (i) open-ended cage controls showed that the effects of caging resulted from exclusion of foragers, not from artifacts of caging; (ii) day-only and night-only caging showed that nectar was depleted only during the day: and (iii) observations showed that cages did not exclude any diurnal foragers other than honeyeaters. Resident honeyeaters spent more time foraging during months when nectar was scarce, implying that the rates at which they could obtain nectar were affected by changes in nectar availability. It is therefore possible that the depletion of nectar by honeyeaters could have limited their densities. However. I argue that such limitation could only be inferred safely if nectar-supplementation experiments showed survival and/or reproduction to be limited by nectar availability.     

Reproductive success of Woodlarks Lullula arborea in traditional and recently colonized habitats

Capsule Set-aside schemes have allowed breeding Woodlarks to colonize farmland, but heathland and forestry habitats remain the stronghold.

Aims To determine which habitats provide the best conditions for breeding Woodlarks and whether a buffer effect is operating, with density increasing faster in the poorer quality habitat as the population grows.

Methods Habitat colonization was examined and breeding success compared among heathland, clear-felled and young conifer plantations, and farmland set-aside.

Results Woodlarks on heathland and forestry habitats had similar clutch sizes and nesting success, but clutch sizes may be lower on farmland. Heathland was recolonized when population density was low in forest habitats, while farmland was colonized when density was increasing, and areas close to forest were preferentially occupied. Woodlarks breeding on farmland preferred set-aside stubbles to other field types.

Conclusion Forestry and heathland habitats are similar in quality for breeding Woodlarks, with no evidence for a buffer effect. Farmland set-aside may be suboptimal but the area available is much greater than the area of forest or heathland, and could therefore make a significant contribution to the conservation of the Woodlark population. However, set-aside should not be seen as an alternative to the conservation of forest and heathland.     

Consistent mixing of near and distant resources in foraging bouts by the solitary mason bee Osmia lignaria

Female bees are usually confronted with a choice among severalflower species that differ in their location and abundance withinthe community, and in the efficiency with which their pollenand nectar can be harvested. We investigated the effects ofdistance and flower density of two flower species on pollencollection by providing nest locations for the mason bee Osmialignaria in natural settings. Distance weakly affected pollenuse; on average, bees nesting near a flower species tended tocollect more of its pollen than did bees nesting at a greaterdistance. Flower density did not predictably impact pollen use,and use did not track changes in density during the season.Bees consistently mixed pollen from more distant species, despitesubstantial added foraging costs, and also mixed when one specieswas an order of magnitude less abundant than the other. Beesrequire nectar as well as pollen to feed their offspring, andour preliminary data suggest that the efficiencies of pollenand nectar collection are inversely related between the twoflower species, which would favor visitation to both species.Bees appear to collect some pollen from the low-pollen, high-nectarplant while visiting it for nectar. Thus, a nectar-collectingconstraint may favor collecting pollen from mixtures of species.     

Yearly variation in seasonal patterns of honeyeater abundance,flower density and nectar production in heathland near Sydney

Despite a consistent seasonal pattern of daily production of nectar-energy per unit area with peak production around the end of May, the seasonal pattern of honeyeater abundance is not consistent between years and does not correspond to the pattern of nectar-energy production.     

Recruitment of grey and red-legged partridges (Perdix perdix and Alectoris rufa) in relation to population density and habitat

The recruitment of first-year grey and red-legged partridges into subsequent breeding populations was investigated on 17 study areas in Britain.
The density of birds recruited per unit area increased with the density of birds available in their first autumn for potential recruitment in the following spring but decreased with adult density. The relationship between recruitment and adult density, while always negative, differed significantly between areas. These differences were correlated with variation in the amount of nesting habitat available to both species of partridge and with variation in nesting habitat quality. Recruitment of grey partridges was related to length of available nesting habitat, the amount of residual dead grass in the nesting cover, and earth bank height at the base of the nesting cover. Recruitment of red-legged partridges was also related to length of nesting habitat, together with the amount of nettle in the nesting cover.
It is suggested that the attractiveness of different areas to settling recruits is determined by the amount and quality of nesting habitat available.     

Pollinator service only depends on nectar production rates in sparse populations

As predicted, approach rate by bumblebees is strongly related to the nectar production rate of Echium vulgare plants in a sparse population, while in a dense population such a relationship is completely absent. These findings are confirmed by additional experiments with potted plants that were placed inside and outside a natural population. The results suggest that the direction of selection on nectar production may vary in space or time depending on population density. Such variation may help to explain the large genetic variation we found earlier for E. vulgare in our study area.     

Age and breeding success of a Golden Eagle Aquila chrysaetos population in southeastern Spain

Brood production rate of Grey Partridges Perdix perdix was studied in three areas in Poland in the years 1986–90, by comparing the number of pairs and the number of family coveys after the breeding period. In the study areas, linear permanent nesting cover was mapped and measured. In these areas the brood production rate declined with increasing spring pair density. A positive effect of the occurrence of nesting cover on brood production rate was found. An increase in the population density mostly occurred through occupying parts of the study area avoided earlier, thus presumably less suitable for Partridges. Pairs occupying these less suitable areas, characterized by less abundant nesting cover, had a lower brood production rate, whereas in the preferred areas this rate did not decline with increasing mean population density.     

Effects of Habitat Quality and Isolation on the Colonization of Restored Heathlands by Butterflies

In areas with intensive land use, such as the Netherlands, habitat fragmentation and loss of habitat quality due to eutrophication and drainage are major threats to the preservation of species‐rich communities of heathland and acid grassland. Restoration of such nutrient‐poor habitats may be carried out by removing the topsoil from ex‐arable land, in order to lower the nutrient levels. However, the establishment of target plant communities is known to be fragmentary. The current study shows that this also applies to butterflies. Ten years after topsoil removal in eight study areas, on average, only 3.5 of 10 characteristic heathland species were recorded on the sites. Species that did colonize had a significantly lower density than in the source populations. Our study indicates that although isolation effects were limiting colonization, poor habitat quality was the main limiting factor, mainly due to lack of host plants, hydrological conditions, and, to a lesser extent, lack of nectar plants and excessive residual nutrient levels. An experiment with the introduction of cut heather in one study area showed a significantly higher abundance of both target and nontarget butterflies in manipulated sites than in control sites. It can be concluded that habitat restoration by topsoil removal can be successful for butterflies of especially wet heathland habitats, provided that source populations are at close range and care is taken that complete plant communities are restored.     

Sources of variation in floral nectar production rate in Epilobium canum (Onagraceae): implications for natural selection

Sources of variation in floral nectar production were investigated in a natural population of Epilobium canum (Onagraceae), a hummingbird-pollinated herbaceous shrub. Field measurements showed significant phenotypic variation among plants in floral nectar production rates. Average variance among flowers within plants was approximately one-third to one-half as great as variance among plants, with coefficients of variation among flowers ranging from 6.5% to 116.7%. A greenhouse experiment using clonally propagated ramets from field plants showed significant genetic variation for nectar production rates; broad sense heritability was estimated to have a maximum value of 0.64. In the greenhouse, plants grown under low water or low light conditions produced approximately 25% less nectar on average than those grown under control conditions. However, significant genotype-environment interactions indicated that genets differed in their responses to the changes in conditions. Rank correlations for genet mean nectar production rates across environmental conditions were low, and in two out of three comparisons were not different from zero. It is concluded that although the opportunity for natural selection on nectar production rates exits in this population, the response to selection will likely be slow, and the opportunity for selection of a narrow-optimum nectar production phenotype may be limited. Received: 9 January 1996 / Accepted: 18 December 1996     

Heritability of nectar production in Echium vulgare

Heritabilities of nectar production in the wild species Echium vulgare were estimated as realised heritability under controlled and field conditions. The nectar production of offspring from high- and low-nectar-producing parents was significantly different in both controlled and field conditions, indicating that nectar production is in part genetically determined. The present study is the first one to report a genetic component of variation of nectar production in a wild plant species in the field. Heritability estimated under controlled conditions was 0.13 and therewith less than the heritability estimated under field conditions, which amounted to 0.26. Offspring of high-nectar-producing plants produced comparable amounts of nectar in the growth chamber (1.28 microl) and in the field (1.22 microl). In contrast, the nectar production of offspring of low-nectar-producing plants was significantly higher in the growth chamber (0.95 microl) than in the field (0.55 microl), indicating a genotype by environment interaction. The level of heritability of nectar production was dependent on the environment. Under less favourable conditions, like those in the field, heritability of nectar production increased. Nectar production was not correlated with any of the vegetative or reproductive traits measured, and hence no costs of nectar production could be detected. Results obtained stress the importance of field measurements in determining heritabilities.     

Population variation in plant traits associated with ant attraction and herbivory in Chamaecrista fasciculata (Fabaceae)

The benefits of ant–plant–herbivore interactions for the plant depend on the abundance of ants and herbivores and the selective pressures these arthropods exert. In plants bearing extrafloral nectaries (EFN), different mean trait values may be selected for by different populations in response to local herbivore pressure, ultimately resulting in the evolution of differences in plant traits that attract ants as defensive agents against herbivory. To determine if variation in traits that mediate ant–plant interactions reflect herbivore selective pressures, we quantified intra- and inter-population variation in plant traits for eight populations of the EFN-bearing annual Chamaecrista fasciculata (Michx.) (Fabaceae). Censuses in rural and urban areas of Missouri and Illinois (USA) showed population differences in ant attendance and herbivore pressure. Seeds were collected from each population, and plants were grown in a common greenhouse environment to measure sugar production, nectar volume and composition, EFN size and time of emergence, leaf pubescence, and leaf quality throughout plant development. Populations varied mainly in terms of nectary size, sugar production, and nectar volume, but to a lesser degree in leaf pubescence. Populations of C. fasciculata within urban areas (low in insect abundance) had small nectaries and the lowest nectar production. There was a positive correlation across populations between herbivore density and leaf damage by those herbivores on the one hand and sugar production and nectar volume on the other. These results, in conjunction with lack of evidence for maternally based environmental effects, suggest that population differences in herbivore damage have promoted differential evolution of EFN-related traits among populations. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

ADAPTIVE SIGNIFICANCE OF IPOMOPSIS AGGREGATA NECTAR PRODUCTION: OBSERVATION AND EXPERIMENT IN THE FIELD

Production of floral nectar is generally thought to be an adaptation that increases plant fitness by altering pollinator behavior, and therefore pollination success. To test this hypothesis, I investigated the effects of floral nectar production rate on pollination success of the hermaphroditic plant Ipomopsis aggregata (Polemoniaceae). Success through male function (estimated by the export of fluorescent dyes) was significantly greater for plants with naturally high nectar production rates than for nearby plants with low nectar production rates, whereas success through female function (receipt of fluorescent dye) was unrelated to nectar production rate. Experimental addition of artificial nectar also produced a significant increase in male function success and no increase in several estimates of female function success. Observations confirmed that hummingbirds probed a larger proportion of flowers on plants that received supplemental nectar, as they do in response to natural variation in nectar production. The concordance of results across these observational and experimental studies indicates that nectar production acts primarily to increase pollination success through male function for this species.     

Plant reproductive strategies vary under low and high pollinator densities

Long‐term variation in the population density of honey bees Apis mellifera across landscapes has been shown to correlate with variation in the floral traits of plant populations in these landscapes, suggesting that variations in pollinator population density and foraging rates can drive floral trait evolution of their host plants. However, it remained to be determined whether this variation in plant traits is associated with adaptive variation in plant reproductive strategies under conditions of high and low pollinator densities. Here we conducted a reciprocal transplant experiment to examine how this variation in floral traits, under conditions of either high and low pollinator density, impacted seed production in the Tibetan lotus Saussurea nigrescens. In 2014 and 2015, we recorded the floral traits, pollinator visitation rates, and seed production of S. nigrescens populations grown in both home sites and foreign sites, where sites varied in honey bee population density. Our results demonstrated that the floral traits reflected those of their original population, regardless of their current location. However, seed production varied with both population origin and transplant site. Seed number was positively correlated with flower abundance in the pollinator‐rich sites, but with nectar production in the pollinator‐poor sites. Pollinator visitation rate was also positively correlated with flower number at pollinator‐rich sites, and with nectar volume at pollinator‐poor sites. Overall, the local genotype had higher seed production than nonlocal genotypes in home sites. However, when pollen is hand‐supplemented, plants from pollinator‐rich populations had higher seed production than plants from pollinator‐poor populations, regardless of whether they were transplanted to pollinator‐rich or ‐poor sites. These results suggest that the plant genotypic differences primarily drive variation in pollinator attraction, and this ultimately drives variation in seed: ovule ratio. Thus, our results suggest that flowering plant species use different reproductive strategies to respond to high or low pollinator densities.     

Seasonal change in offspring sex and size in Dawson's burrowing bees (Amegilla dawsoni) (Hymenoptera: Anthophorini)

Abstract.  1. Nesting females of Dawson's burrowing bees, Amegilla dawsoni , produce a large size class of offspring, which includes daughters and major sons, and a small size class, which consists entirely of minor sons averaging half the weight of their larger siblings. Female allocation patterns change over the flight season such that the initial pattern of producing daughters shifts toward the production of both daughters and major sons in the middle of the season, and then the production of primarily minor sons in the latter part of the nesting season.
2. In Dawson's burrowing bees, this pattern is correlated with declines in pollen and nectar availability as the nesting season progresses as well as a heightened risk of dying before the final brood cell is completed. Here, the relation between these factors and the provisioning tactics of nesting Dawson's burrowing bees is discussed.     

Nectar yeasts in Delphinium nuttallianum (Ranunculaceae) and their effects on nectar quality

Microorganisms colonize the nectar of many angiosperms. Variable diversity and spatio-temporal dynamics of nectar-inhabiting microorganisms (e.g., yeasts) may drive variation in nectar sugar composition and subsequent plant–pollinator interactions. We assessed yeast frequency of occurrence and density in the nectar of the perennial herb, Delphinium nuttallianum, across multiple spatio-temporal scales, including flower lifetime and sex-phase transition, flowering season, populations, and years. We tested the hypothesis that pollinators vector yeasts by comparing densities between virgin flowers and those open to visitation. Finally, we identified yeasts using molecular methods and tested for an association between yeast density and nectar composition using ultra-performance liquid chromatography. Yeasts were frequent colonists of Delphinium nectar, occurring in all populations and years sampled. Yeast frequency of occurrence and density varied across most spatio-temporal scales examined. Pollinators were vectors of yeast: virgin flowers remained yeast-free, while those open to visitation became inoculated. Nectar samples were species-poor, with a majority colonized by Metschnikowia reukaufii. Finally, increasing yeast density was correlated with a decrease in sucrose and an increase in monosaccharides. Our results document that yeasts form species-poor communities in populations of this hermaphroditic perennial, in addition to highlighting their spatio-temporal dynamics and effects on nectar quality. Spatio-temporal variation in frequency of occurrence, density, and changes in nectar may have important implications for the nature and strength of interactions between Delphinium and its pollinators.