Non-native plants affect generalist pollinator diet overlap and foraging behavior indirectly,via impacts on native plant abundance

Flowering invasive plants can have dramatic effects on the resource landscape available to pollinators. Because many pollinators exhibit behavioral plasticity in response to competitor or resource density, this in turn can result in impacts on ecological processes such as pollination and plant reproduction. We examine how interactions between five common generalist eusocial bees change across an invasion gradient by examining how bee abundance and diet overlap changed with variation in both invasive plant abundance and competitor abundance in a temperate oak-savannah ecosystem. Specifically we focus on the bumblebees Bombus bifarius, B. mixtus, B. melanopygus and B. vosnesenskii, as well as the non-native honeybee Apis mellifera, and their interactions with the native flowering plants Camassia quamash, Camassia liechtlinii, and the invasive shrub Cytisus scoparius. We further examine whether changes in pollinator visits to the invasive and two common native plants can explain changes in diet overlap. Abundance of the invasive plant and other common floral resources had strong impacts on focal bee abundance, with certain species more likely to be present at highly invaded sites. This may be because highly invaded sites tended to be embedded in forested landscapes where those bees are common. Diet overlap was most affected by abundance of a common native plant, rather than the invasive plant, with diet overlap increasing non-linearly with abundance of the native plant. Furthermore, Apis mellifera, did not appear to have direct competitive effects on native bumblebees in this habitat. However, visit patterns suggest that bees most abundant at highly invaded sites may compete for access to native resources. Thus the impacts of this invasive plant on our focal bee species may be primarily indirect, via its’ competitive effects on native plants.     

Analysis of pollinator foraging: tests for non-random behaviour

1. A standardized protocol for analysing the behaviour of pollinators foraging on more than one plant type (species or morph) is needed.
2. A protocol is presented in which the first step is to test whether foraging trips are homogeneous in the frequency of visits to each plant type, or whether there are two or more groups of pollinators with different foraging preferences.
3. Tests for foraging preference and constancy in the sequence of plants visited then should be made separately for each homogeneous group.
4. A hypothetical example is given in which ignoring heterogeneity of preference would lead to the acceptance of a false null hypothesis of random behaviour.
5. A real example is given in which heterogeneity of preference was the only non-random aspect of pollinator behaviour (no significant positive constancy or overall preference), while transfer of powdered dye particles (pollen analogues) was assortative among floral morphs.
6. Heterogeneity of preference is probable, as many factors may cause individual pollinators to forage differently on the same patch of plants, and may be sufficient to produce non-random pollen transfer among plant morphs.     

Plant population dynamics, pollinator foraging, and the selection of self-fertilization

Many flowering plants rely on pollinators, self-fertilization, or both for reproduction. We model the consequences of these features for plant population dynamics and mating system evolution. Our mating systems-based population dynamics model includes an Allee effect. This often leads to an extinction threshold, defined as a density below which population densities decrease. Reliance on generalist pollinators who primarily visit higher density plant species increases the extinction threshold, whereas autonomous modes of selfing decrease and can eliminate the threshold. Generalist pollinators visiting higher density plant species coupled with autonomous selfing may introduce an effect where populations decreasing in density below the extinction threshold may nonetheless persist through selfing. The extinction threshold and selfing at low density result in populations where individuals adopting a single reproductive strategy exhibit mating systems that depend on population density. The ecological and evolutionary analyses provide a mechanism where prior selfing evolves even though inbreeding depression is greater than one-half. Simultaneous consideration of ecological and evolutionary dynamics confirms unusual features (e.g., evolution into extinction or abrupt increases in population density) implicit in our separate consideration of ecological and evolutionary scenarios. Our analysis has consequences for understanding pollen limitation, reproductive assurance, and the evolution of mating systems.     

Trapline foraging by bumble bees: III. Temporal patterns of visitation and foraging success at single plants

We analyzed the temporal structure of visitation by bumble beeworkers to a single Penstemon strictus plant growing in an arrayof conspecifics. When tested against a null distribution usinga randomization model, the observed pattern of arrivals forthe whole group of bees was random, but departures were clusteredin time. Certain individuals visited the plant repeatedly andfrequently throughout the day. These showed significantly regulararrival and departure schedules, which were likely producedby traplining. We explored whether these more frequent and regularforagers gained a higher reward than random or incidental plantvisitors. Using an analytical model, Possingham predicted thata dominating forager that visited a simple, renewing resourcein a regular pattern would garner higher and less variable rewardsthan random visitors. Inspired by these results, but interestedin plant-level visitation, we constructed a simulation modelof resource dynamics for a multiflowered plant with high visitation.The model incorporates the observed visitation schedules ofall bees and independent reward dynamics for each flower onthe plant. We calculated the rewards that observed bees wouldhave collected given a range of resource-renewal parameters.More frequent visitors did not return to the plant when whole-plantresource levels were higher, but these visitors did get greaterrewards. Their increased reward resulted from greater foragingefficiency, primarily through selecting (on average) more rewardingflowers than those selected by less frequent, random visitors.     

Field margins, foraging distances and their impacts on nesting pollinator success

The areas of wild land around the edges of agricultural fields are a vital resource for many species. These include insect pollinators, to whom field margins provide both nest sites and important resources (especially when adjacent crops are not in flower). Nesting pollinators travel relatively short distances from the nest to forage: most species of bee are known to travel less than two kilometres away. In order to ensure that these pollinators have sufficient areas of wild land within reach of their nests, agricultural landscapes need to be designed to accommodate the limited travelling distances of nesting pollinators. We used a spatially-explicit modelling approach to consider whether increasing the width of wild strips of land within the agricultural landscape will enhance the amount of wild resources available to a nesting pollinator, and if it would impact differently on pollinators with differing foraging strategies. This was done both by creating field structures with a randomised geography, and by using landscape data based upon the British agricultural landscape. These models demonstrate that enhancing field margins should lead to an increase in the availability of forage to pollinators that nest within the landscape. With the exception of species that only forage within a very short range of their nest (less than 125 m), a given amount of field margin manipulation should enhance the proportion of land available to a pollinator for foraging regardless of the distance over which it normally travels to find food. A fixed amount of field edge manipulation should therefore be equally beneficial for both longer-distance nesting foragers such as honeybees, and short-distance foragers such as solitary bees.     

Yearlong association of insect pollinator,Pseudapis oxybeloides with flowering plants: Planted forest vs. agricultural landscape

The yearlong association of a native bee, Pseudapis oxybeloides (Halictidae: Hymenoptera) was studied with 72 plant species in a sub-tropical planted forest and some adjacent agricultural landscapes at Multan, Pakistan. The study resulted in 66 interactions of P. oxybeloides with only 24 plant species in 15 families while other 48 plant species were not visited by this bee. The maximum abundance of P. oxybeloides (7–9 individuals) was recorded on Achyranthes aspera and Launaea procumbens followed by Ageratum conyzoides, Trianthema portulacastrum and Cleome viscosa (5–6 individuals). Majority (19) of plant species were visited by only 1–4 individuals. The bee activity was started in the month of March which attained its peak in May followed by a gradual decline until September. No bees were observed during the months of January and February. There was a significant positive relationship between bee abundance and number of flowering plant species. Bee abundance had a strong positive relationship with temperature while it had a strong negative relationship with relative humidity (%). Floral abundance increased with the number of flowering plant species while it was not influenced by floral span of plant species. Besides giving the floral host plants of P. oxybeloides, the current study also gives a better understanding of its seasonality along with its relationships with different biotic and abiotic factors under local conditions. These findings can help in maintaining and managing P. oxybeloides population particularly and other native bees in general at local scale.     

Variations of behavioural patterns during development in the water stick insect

Variations of associations between different behavioural patterns were analysed during post-embryonic development in the water stick insect,Ranatra linearis L. (Heteroptera: Nepidae). Behavioural patterns recorded were: quiescence, swimming, crawling, leg waving, body swaying, respiratory ascents breathing, respiratory descents, grooming and feeding activities (predatory attempts, foreleg movements, ingestion). A factorial analysis of correspondance stressed the contrast between the organization of larval behaviour and that of adult behaviour. No behavioural patterns were dropped from the repertoire during post-embryonic development ofRanatra linearis and no new patterns developed in adults outside the reproductive period. Larval behaviour was characterized by more frequent respiratory activities and grooming and adult behaviour was characterized by more frequent locomotory activities. At all larval stages grooming was associated with respiratory activities and crawling, but not with feeding activities.     

The influence of climatic seasonality on the diversity of different tropical pollinator groups

Tropical South America is rich in different groups of pollinators, but the biotic and abiotic factors determining the geographical distribution of their species richness are poorly understood. We analyzed the species richness of three groups of pollinators (bees and wasps, butterflies, hummingbirds) in six tropical forests in the Bolivian lowlands along a gradient of climatic seasonality and precipitation ranging from 410 mm to 6250 mm. At each site, we sampled the three pollinator groups and their food plants twice for 16 days in both the dry and rainy seasons. The richness of the pollinator groups was related to climatic factors by linear regressions. Differences in species numbers between pollinator groups were analyzed by Wilcoxon tests for matched pairs and the proportion in species numbers between pollinator groups by correlation analyses. Species richness of hummingbirds was most closely correlated to the continuous availability of food, that of bees and wasps to the number of food plant species and flowers, and that of butterflies to air temperature. Only the species number of butterflies differed significantly between seasons. We were not able to find shifts in the proportion of species numbers of the different groups of pollinators along the study gradient. Thus, we conclude that the diversity of pollinator guilds is determined by group-specific factors and that the constant proportions in species numbers of the different pollinator groups constitute a general pattern.     

Diversity and abundance patterns of phytophagous insect communities on alien and native host plants in the Brassicaceae

The herbivore load (abundance and species richness of herbivores) on alien plants is supposed to be one of the keys to understand the invasiveness of species. We investigate the phytophagous insect communities on cabbage plants (Brassicaceae) in Europe. We compare the communities of endophagous and ectophagous insects as well as of Coleoptera and Lepidoptera on native and alien cabbage plant species. Contrary to many other reports, we found no differences in the herbivore load between native and alien hosts. The majority of insect species attacked alien as well as native hosts. Across insect species, there was no difference in the patterns of host range on native and on alien hosts. Likewise the similarity of insect communities across pairs of host species was not different between natives and aliens. We conclude that the general similarity in the community patterns between native and alien cabbage plant species are due to the chemical characteristics of this plant family. All cabbage plants share glucosinolates. This may facilitate host switches from natives to aliens. Hence the presence of native congeners may influence invasiveness of alien plants.     

Wound induced defences in plants and their consequences for patterns of insect grazing

Summary Three scales of wound-induced chemical responses in plants are identified: (1) highly localised chemical changes associated with disruption of cell compartmentation; (2) changes induced in cells surrounding the damaged area, forming a kind of halo around the damage, and (3) more widely-dispersed changes which may affect an entire organ, branch or plant. A brief review of the literature reveals that such chemical responses are very widespread in plants, and many of the substances formed are known to affect adversely the growth, development, or reproduction of insects. It is argued that wound-induced changes in plant chemistry represent for insects a powerful selective pressure for the dispersal of grazing. Levels and patterns of invertebrate grazing in a range of herbaceous and deciduous woody plants sampled at the end of the growing seasons were examined. Leaves of many species exhibited a strikingly evident over-dispersion of grazing initiations, and in some cases the arrangement of holes appeared close to regularity. The pattern of damage between leaves was, in most cases, heavily biased towards a large proportion of leaves receiving a low level of grazing. These highly dispersed patterns of grazing damage are consistent with the hypothesis that wound-induced responses play an important role in determining patterns of insect feeding. They have important implications for the expected levels of insect exploitation of host plants and for the advantages to the plant of distributing grazing damage evenly through the canopy.     

Voracity of the main stored-product insect pests on wheat grain

The results of an experimental study of voracity of adults and larvae of seven main stored-product insect pests are described. One adult of Sitophilus oryzae, S. granarius, Rhizopertha dominica, Tribolium castaneum, T. confusum, and Oryzaephilus surinamensis consumed 0.38, 0.48, 0.89, 0.15, 0.16, and 0.11 mg of wheat grain per day, respectively. During their development, larvae of these insects consumed 7.60, 13.77, 7.92, 3.32, 3.60, and 2.32 mg of wheat grain, respectively. One larva of Sitotroga cerealella consumed 17.46 mg of wheat grain. The coefficients of harmfulness of these species are calculated, and a method for estimating the “total infestation density” of grain is proposed.     

Movement patterns of a clear-wing hawkmoth,Hemaris fuciformis,foraging at red catchfly,Viscaria vulgaris

Summary The bee hawkmoth, Hemaris fuciformis, tended to fly in the same direction between successive visits to inflorescences of Viscaria vulgaris. Upon leaving an inflorescence it did not fly to the first inflorescence in its path, but to the second. At inflorescences, the number of animals that started probing decreased from the bottom to the top. These movement patterns probably serve to reduce the risk of revisiting flowers. The movements of the moths at inflorescences with 2 flowers per tier were investigated and related to the nectar distribution. The nectar is distributed according to a bonanza-blank pattern, flowers being either empty or full, probably because previous visitors have emptied some, but not all, flowers. It is shown that the two flowers of a tier more often than random belong to the category: blank-bonanza. Thus, if a blank is found, the animal should move sideways to the second flower at the same tier in order to find a bonanza. The flowers of the inflorescences tend to be arranged in vertical rows, and it is shown that two flowers, one above the other, more often than random belong to the two categories: blank-blank or bonanza-bonanza. Thus, if a bonanza is found, the animal should move up to the flower above, in order to find another bonanza. Rules of movement at inflorescences and a rule of departure are proposed based on the probability distribution of the nectar. The observed movements agree quite well with the predicted ones.     

Parallels between the foraging strategies of ants and plants

Animal and plant ecologists generally follow separate paths. This often leads to disjointed approaches to solving similar ecological problems. In the past 20 years, two related, but unconnected, research fields have undergone rapid development: modular demography, with its morphological and functional analysis of resource capture, until now basically the domain of plant ecology; and foraging theory, traditionally applied and developed in animal ecology. The results of recent research on the foraging strategies of ants and clonal plants, however, outline a general framework of functional parallels between both types of organisms that could link important aspects of animal and plant foraging ecology.     

The colour of flowers in spectrally variable illumination and insect pollinator vision

The spectral reflectance of differently coloured Australian native plant flowers and foliage was measured and plotted in a colour triangle to represent the colour space of the honeybee. Spectral variations in illumination are shown to significantly change plant colours for bee vision without colour constancy. A model of chromatic adaptation based upon the von Kries coefficient law shows a reduction in plant colour shift, with the degree of correction depending upon position in colour space. A set of artificial reflectances is used to map relative colour shift caused by spectrally variable illumination for the entire colour space of the honeybee. The rarity of some flower colours in nature shows a correlation to a larger colour shift for these colours when illuminated by spectrally variable radiation. The model of chromatic adaptation is applied to illuminations used in a behavioural study on honeybee colour constancy by Neumeyer 1981. Surface colours used by Neumeyer are plotted in colour space for the various illuminations. The results show that an illumination-dependent colour shift correlates to a decrease in the frequency of bees correctly choosing a colour to which it was trained. Accepted: 23 February 1998     

Movement patterns of selected insect groups between natural forest,open land and rubber plantation in a tropical landscape (southern Yunnan,SW China)

In many regions of tropical Asia, the expansion of rubber monoculture plantations is conducted by replacement of natural forest areas and strongly affects biodiversity and movement patterns of wild species, including insects. Against this background, we conducted a study on selected insect groups (longhorn beetles, bark beetles, wild bees and hoverflies) along transects between rainforest patches, open uncultivated land and rubber plantation habitats in a region of Xishuangbanna (southern Yunnan, China), with the objectives to identify (a) movement directions and patterns of selected insect groups based on their abundances in modified Malaise traps in the different habitats, and (b) the role of remaining natural rainforest patches and rubber plantations, respectively, for insect diversity maintenance and conservation. The highest total numbers of species and individuals of bark beetles, longhorn beetles and wild bees were recorded from the natural forest edge compared to open land and rubber plantation edge. This result clearly indicates that the natural forest plays an important role in maintenance of these three insect groups. However, the highest number of hoverfly species and individuals was recorded from the open land sites, indicating the most relevant habitat type for this group of species. Overall, the lowest species and individual numbers were recorded from the rubber plantation edge, indicating the unsuitability of this habitat type for all insect groups considered. The distribution of species and individuals in the opposite trap sides along the transect indicates that longhorn beetles, bark beetles and wild bees show not only movements from the forest to the surrounding habitats, but also return back after encountering the unsuitable rubber plantation habitat. Bark beetle composition showed the relatively highest similarity between all trap sites and opposite trap sides among the insect groups considered, indicating a higher movement activity than the other groups. The four insect groups considered in this study show different movement modes between the forest, open land and rubber plantation, which are not the same for all taxa. Except for hoverflies, the natural forest was found to be the most important habitat for the maintenance of species diversity in the different land use types of the study area.     

Frequency-dependent pollinator discrimination acts against female plants in the gynodioecious Geranium maculatum

MethodsTo test the severity and consequences of this type of pollinator discrimination in Geranium maculatum, experimental populations with the range of sex ratios observed in nature were created, ranging from 13 % to 42 % females. Pollinators were observed in order to measure the strength of discrimination, and pollen deposition and seed production of both sexes were measured to determine the fitness consequences of this discrimination. Additionally a comparison was made across the sex ratios to determine whether discrimination was frequency-dependent.ConclusionsThe results suggest that pollinator discrimination negatively affects females'' relative fitness when they are rare. Thus, the initial spread of females in a population, the first step in the evolution of gynodioecy, may be made more difficult due to pollinator discrimination.