Effects of floral diversity and pollinator behaviour on the persistence of hybrid zones between plants sharing pollinators

Background: Plant hybrid zones often display a large diversity in floral traits due to segregation and recombination occurring in the second and later hybrid generations. Such diversity can have important effects on pollinator behaviour.

Aims: We hypothesised that the number of floral types and their relative densities may influence pollinator preferences and level of flower constancy.

Methods: To test the influence of floral diversity on the number of visits to inflorescences of the most common type and on the level of flower constancy, we monitored bumblebees’ behaviour in experimental arrays of artificial inflorescences with a common (‘parental’) floral type mixed with either one or three uncommon (‘hybrid’) floral types.

Results: We found that pollinators preferentially visited the most common flower colour and showed slightly greater constancy when the number of uncommon floral types was increased.

Conclusions: We conclude that diversity in flower colour could promote moderate pollinator constancy in some types of hybrid zones. Segregation of floral phenotypes due to hybridisation may result in small levels of pollinator-mediated assortative mating and contribute, to a small extent, and most probably in combination with other factors, to the persistence of some hybrid zones.     

Permanent coexistence in general models of three interacting species

We address the question of the long term coexistence of three interacting species whose dynamics are governed by the ordinary differential equations x _i = X _i f _i (i = 1, 2, 3). In order for any theory in this area to be useful in practice, it must utilize as little information as possible concerning the forms of the f _i , in view of the great difficulty of determining these experimentally. Here we obtain, under rather general conditions on the equations, a criterion for judging whether the species will coexist in a biologically realistic manner. This criterion depends only on the behaviour near the one or two species equilibria of the two dimensional subsystems, the behaviour there being relatively easy to examine experimentally. We show that with the exception of one class of cases, which is a generalization of a classical example of May and Leonard [21], invasibility at each such equilibrium suitably interpreted is both necessary and sufficient for a strong form of coexistence to hold. In the exceptional case, a single additional condition at the equilibria is enough to ensure coexistence.     

鸡嗉子榕内传粉小蜂与非传粉小蜂的繁殖稳定共存机制研究

【目的】榕-蜂系统是自然界协同进化和互利共生的典型模型,榕-蜂之间如何实现繁殖分配从而实现稳定共存一直是倍受研究者关注的问题,但对传粉小蜂和非传粉小蜂的繁殖稳定共存研究较少。【方法】本文选取分布在西双版纳地区的雌雄异株榕树鸡嗉子榕Ficus semicordata的雄果,通过对榕果内唯一的传粉小蜂Ceratosolen gravelyi Grandi以及4种非传粉小蜂Philotrypesis dunia Joseph,Apocrypta sp.,Platyneura sp.,Sycoscapter trifemmensis Joseph的产卵时序进行了观察,并对5种小蜂的种群数量进行统计,以及对每种小蜂产卵小花的空间分布、瘿花体积等特征进行了研究。【结果】发现:5种小蜂在产卵时间上存在分离,且传粉小蜂的种群数量远大于其他4种非传粉小蜂的种群数量之和。5种小蜂的瘿花特征也存在差异:瘿花的子房大小不同,且花梗长度即分布空间也有差异,传粉小蜂最接近果腔而非传粉小蜂则更多地利用中层和内层的小花。【结论】结果说明在鸡嗉子榕有限的空间内,5种小蜂通过产卵时间和生长空间的分离,实现了减少竞争以及长期共存的特征,维持了榕-蜂系统的稳定性。     

Honey bee orientation behaviour and the influence of flower distribution on foraging movements

ABSTRACT.

• 1 Directional movement by foraging honey bees (Apis mellifera L.) was studied on several flower arrays. The most frequent move among equidistant flower stalks was straight ahead from stalk to stalk with frequencies decreasing for increasing turn angles. Turns to the left were about equal in frequency to turns to the right.
• 2 Bees maintained directionality when moving from flower stalks that had been rotated 90° counterclockwise while the bee was on the stalk (no difference between moves from rotated stalks and unrotated controls). Thus, directionality is maintained by the bee and is not an artefact of flower distribution.
• 3 Bees also maintained directionality when the entire array was rotated around the flower stalk the bee was on. Thus, bees use an external cue to orientate in a given direction rather than fixing on an inflorescence within the flower array.
• 4 Bees foraging on very different flower arrays differed in patterns of directionality and in distances flown between flower stalks. Therefore, even though bees maintain directionality using external cues, flower distribution can nevertheless influence flight patterns.

     

Interpopulation variation in predator foraging behaviour promotes the evolutionary divergence of prey

Despite intensive investigation of the role of predation on evolutionary processes, few studies have questioned the possibility of the evolutionary divergence of prey populations in response to interpopulation variation in predator foraging behaviour. In an interaction between a seed-predatory insect, the camellia weevil (Curculio camelliae), and its host plant, the Japanese camellia (Camellia japonica), I tested whether the evolutionary differentiation of the plant's defensive trait, pericarp thickness, was related to the interpopulation variation in the foraging behaviour of female weevils. I found that the preference of weevils for the plant fruit based on pericarp thickness varied across 13 populations in Japan. Importantly, variation in weevil behaviour explained interpopulation variation in pericarp thickness and the direction/strength of natural selection on the trait. Overall, I show that adaptive foraging of predators can result in the evolutionary divergence of predator-prey interactions.     

Adaptive omnivory and species coexistence in tri-trophic food webs

The commonness of omnivory in natural communities is puzzling, because simple dynamic models of tri-trophic systems with omnivory are prone to species extinction. In particular, the intermediate consumer is frequently excluded by the omnivore at high levels of enrichment. It has been suggested that adaptive foraging by the omnivore may facilitate coexistence, because the intermediate consumer should persist more easily if it is occasionally dropped from the omnivore's diet. We explore theoretically how species permanence in tri-trophic systems is affected if the omnivore forages adaptively according to the "diet rule", i.e., feeds on the less profitable of its two prey species only if the more profitable one is sufficiently rare. We show that, compared to systems where omnivory is fixed, adaptive omnivory may indeed facilitate 3-species persistence. Counter to intuition, however, facilitation of 3-species coexistence requires that the intermediate consumer is a more profitable prey than the basal resource. Consequently, adaptive omnivory does not facilitate persistence of the intermediate consumer but enlarges the persistence region of the omnivore towards parameter space where a fixed omnivore would be excluded by the intermediate consumer. Overall, the positive effect of adaptive omnivory on 3-species persistence is, however, small. Generally, whether omnivory is fixed or adaptive, 3-species permanence is most likely when profitability (=conversion efficiency into omnivores) is low for basal resources and high for intermediate consumers.     

Within-patch foraging behaviour of the aphid parasitoidAphidius funebris: plant architecture, host behaviour, and individual variation

The influence of plant architecture, host colony size, and host colony structure on the foraging behaviour of the aphid parasitoidAphidius funebris Mackauer (Hymenoptera: Aphidiidae) was investigated using a factorial experimental design. The factorial design involved releasing individual parasitoid females in aphid colonies consisting of either 10 or 20 individuals ofUroleucon jaceae L. (Homoptera: Aphididae) of either only larval instar L3 or a mixture of host instars, both on unmanipulated plants and on plants that had all leaves adjacent to the colony removed. Interactions between the parasitoid and its host were recorded until the parasitoid had left the plant. The time females spent on the host plant and the number of eggs laid varied greatly among females. Host colony size significantly affected patch residence time and the number of contacts between parasitoids and aphids. Plant architecture influenced the time-budget of the parasitoids which used leaves adjacent to the aphid colony for attacking aphids. Female oviposition rate was higher on unmanipulated plants than on manipulated plants. No further significant treatment effects on patch residence time, the number of contacts, attacks or ovipositions were found. Oviposition success ofA. funebris was influenced by instar-specific host behaviour. Several rules-of-thumb proposed by foraging theory did not account for parasitoid patch-leaving behaviour.     

Thermoregulation of foraging honeybees on flowering plants: seasonal variability and influence of radiative heat gain

1. During nectar and pollen foraging in a temperate climate, honeybees are exposed to a broad range of ambient temperatures, challenging their thermoregulatory ability. The body temperature that the bees exhibit results from endothermic heat production, exogenous heat gain from solar radiation, and heat loss. In addition to profitability of foraging, season was suggested to have a considerable influence on thermoregulation. To assess the relative importance of these factors, the thermoregulatory behaviour of foragers on 33 flowering plants in dependence on season and environmental factors was investigated.2. The bees (Apis mellifera carnica Pollman) were always endothermic. On average, the thorax surface temperature (T(th)) was regulated at a high and rather constant level over a broad range of ambient temperatures (T(th) = 33.7-35.7°C, T(a) = 10-27°C). However, at a certain T(a), T(th) showed a strong variation, depending on the plants from which the bees were foraging. At warmer conditions (T(a) = 27-32°C) the T(th) increased nearly linearly with T(a) to a maximal average level of 42.6 °C. The thorax temperature excess decreased strongly with increasing T(a) (T(th)-T(a) = 21.6 - 3.6°C).3. The bees used the heat gain from solar radiation to elevate the temperature excess of thorax, head, and abdomen. Seasonal dependance was reflected in a 2.7 °C higher mean T(th) in the spring than in the summer. An anova revealed that season had the greatest effect on T(th), followed by T(a) and radiation.4. It was presumed the foragers' motivational status to be the main factor responsible for the variation of T(th) between seasons and different plants.     

Floral preferences of hoverflies (Diptera: Syrphidae) in response to the abundance and species richness of flowering plants

For understanding the relationship between floral resources and the abundance of hoverflies, five meadows were selected at different elevations in a forested region. At each site the number of flowering herbaceous plants was assessed along a transect at 10-m intervals. To measure floral density, we counted the number of open flowers per species in each plot. Hoverflies visiting flowering plants were captured in each plot during 10 minutes. Path analysis showed that there was a negative correlation between plant species richness and syrphid abundance. Flower abundance had a strong positive direct effect on syrphid abundance but had indirect effects on the number of syrphid species. Only altitude had a strong positive direct effect on the number of syrphid species and this may be explained by the different climatic conditions. Some plant species attracted more syrphids than others. There were significantly fewer species of hoverfly per plot with increasing numbers of flowers. We interpret this to mean that plant species with a high density of flowers attracted more syrphids of the same species. Analysis of pollen grains extracted from two species of hoverflies showed that flower constancy occurred in three examined sites.     

Adaptive foraging by predators as a cause of predator-prey cycles

Summary When foraging has costs, it is generally adaptive for foragers to adjust their foraging effort in response to changes in the population density of their food. If effort decreases in response to increased food density, this can result in a type-2 functional response; intake rate increases in a negatively accelerated manner as prey density increases. Unlike other mechanisms for type-2 responses, adaptive foraging usually involves a timelag, because foraging behaviours do not often change instantaneously with changes in food density or risks. This paper investigates predator-prey models in which there are explicit dynamics for the rate of adaptive change. Models appropriate to both behavioural and evolutionary change are considered. Both types of change can produce cycles under similar circumstances, but under some evolutionary models there is not sufficient genetic variability for evolutionary change to produce cycles. If there is sufficient variability, the remaining conditions required for cycles are surprisingly insensitive to the nature of the adaptive process. A predator population that approaches the optimum foraging strategy very slowly usually produces cycles under similar conditions as does a very rapidly adapting population.     

Adaptive feeding across environmental gradients and its effect on population dynamics

This paper analyzes a consumer's adaptive feeding response to environmental gradients. We consider a consumer-resource system where resources are distributed among many discrete resource patches. Each consumer exhibits a feeding morphology allowing it to remove resources from a patch down to some threshold density (or level) before having to seek resources elsewhere. Assuming consumers trade off resource extraction with patch access and predation, we show that for a given environment there often exists a single evolutionarily stable feeding threshold and it is an evolutionary attractor. We then investigate how the population dynamics of the resource and the consumer change as the environment changes. Two cases are considered: (i) all consumers exhibit a fixed feeding threshold that is adaptive for an intermediate environment; and (ii) the consumer population adapts and adopts the evolutionarily stable feeding threshold associated with the current environment. In less harsh environments (i.e., environments where consumers experience a lower risk of predation, or environments where resource patches are more abundant) the adaptive consumer population is predicted to evolve so that resources within a patch are depleted to lower densities. We show that the change in consumer density due to environmental change can be rather different depending on whether or not the population can adapt. In some situations we observe that when the consumer's environment becomes harsher, the consumer population may increase in density before a rapid crash to extinction. This result has implications for monitoring and managing a population.     

The correlation of learning speed and natural foraging success in bumble-bees

Despite the widespread assumption that the learning abilities of animals are adapted to the particular environments in which they operate, the quantitative effects of learning performance on fitness remain virtually unknown. Here, we evaluate the learning performance of bumble-bees (Bombus terrestris) from multiple colonies in an ecologically relevant associative learning task under laboratory conditions, before testing the foraging performance of the same colonies under the field conditions. We demonstrate that variation in learning speed among bumble-bee colonies is directly correlated with the foraging performance, a robust fitness measure, under natural conditions. Colonies vary in learning speed by a factor of nearly five, with the slowest learning colonies collecting 40% less nectar than the fastest learning colonies. Such a steep fitness function is suggestive of strong selection for higher learning speed. Partial correlation analysis reveals that other factors such as forager body size or colour preference appear to be negligible in our study. Although our study does not directly prove causality of learning on foraging success, our approach of correlating natural within-species variation in these two factors represents a major advance over traditional between-species correlative analyses where comparability can be compromised by the fact that species vary along multiple dimensions.     

匍匐茎草本植物形态可塑性、整合作用与觅食行为研究进展

综述了匍匐茎型克隆植物在形态可塑性、整合作用及觅食行为方面的研究进展。资源斑块性分布是生境异质性的特征之一,适应于异质性生境,匍匐茎植物对环境资源表现了一系列可塑性反应。本文着重从匍匐茎植物对光、水、肥的可塑性反应及其整合作用以及觅食行为等方面的研究进行总结分析,以期对匍匐茎型克隆植物进行更广泛深入的研究。     

Concurrent niche and neutral processes in the competition-colonization model of species coexistence

The importance of neutral dynamics is contentiously debated in the ecological literature. This debate focuses on neutral theory's assumption of fitness equivalency among individuals, which conflicts with stabilizing fitness that promotes coexistence through niche differentiation. I take advantage of competition-colonization trade-offs between species of aquatic micro-organisms (protozoans and rotifers) to show that equalizing and stabilizing mechanisms can operate simultaneously. Competition trials between species with similar colonization abilities were less likely to result in competitive exclusion than for species further apart. While the stabilizing mechanism (colonization differences) facilitates coexistence at large spatial scales, species with similar colonization abilities also exhibited local coexistence probably due to fitness similarities allowing weak stabilizing mechanisms to operate. These results suggest that neutral- and niche-based mechanisms of coexistence can simultaneously operate at differing temporal and spatial scales, and such a spatially explicit view of coexistence may be one way to reconcile niche and neutral dynamics.     

Floral volatiles, pollinator sharing and diversification in the fig-wasp mutualism: insights from Ficus natalensis, and its two wasp pollinators (South Africa)

Combining biogeographic, ecological, morphological, molecular and chemical data, we document departure from strict specialization in the fig-pollinating wasp mutualism. We show that the pollinating wasps Elisabethiella stuckenbergi and Elisabethiella socotrensis form a species complex of five lineages in East and Southern Africa. Up to two morphologically distinct lineages were found to co-occur locally in the southern African region. Wasps belonging to a single lineage were frequently the main regional pollinators of several Ficus species. In South Africa, two sister lineages, E. stuckenbergi and E. socotrensis, pollinate Ficus natalensis but only E. stuckenbergi also regularly pollinates Ficus burkei. The two wasp species co-occur in individual trees of F. natalensis throughout KwaZulu-Natal. Floral volatile blends emitted by F. natalensis in KwaZulu-Natal were similar to those emitted by F. burkei and different from those produced by other African Ficus species. The fig odour similarity suggests evolutionary convergence to attract particular wasp species. The observed pattern may result from selection for pollinator sharing among Ficus species. Such a process, with one wasp species regionally pollinating several hosts, but several wasp species pollinating a given Ficus species across its geographical range could play an important role in the evolutionary dynamics of the Ficus-pollinating wasp association.     

Learning in the nectar foraging behaviour of Helicoverpa armigera

Abstract .1. Learning may enable insects to obtain nectar from flowers more efficiently. Learning in nectar foraging has been shown primarily in studies of bees and butterflies. Here, learning is demonstrated in the nectar foraging behaviour of a noctuid moth, Helicoverpa armigera .
2. The present studies show that: (1) previous experience with a flowering host species increases the probability of that species being selected for nectar foraging, and (2) previous experience of a particular flower type (food source at bottom or top of the corolla tube) increases the likelihood of the food source being found when that flower type is being searched.
3. The implications of these findings for understanding the pattern of oviposition observed in wild populations of this important pest species are discussed.     

The presence of co-flowering species facilitates reproductive success of Pedicularis monbeigiana (Orobanchaceae) through variation in bumble-bee foraging behaviour

Background and Aims The presence of co-flowering species can alter pollinator foraging behaviour and, in turn, positively or negatively affect the reproductive success of the focal species. Such interactions were investigated between a focal species, Pedicularis monbeigiana, and a co-flowering species, Vicia dichroantha, which was mediated by behaviour alteration of the shared bumble-bee pollinator. Methods Floral display size and floral colour change of P. monbeigiana were compared between pure (P. monbeigiana only) and mixed (P. monbeigiana and V. dichroantha) plots in two populations. Pollinator visitation rates, interspecific floral switching and successive within-plant pollinator visits were recorded. In addition, supplemental pollination at plant level was performed, and the fruit set and seed set were analysed in pure and mixed plots with different densities of P. monbeigiana. Key Results Pollinator visitation rates were dramatically higher in mixed plots than in pure plots. The higher pollinator visitation rates were recorded in both low- and high-density plots. In particular, successive flower visits within an individual plant were significantly lower in mixed plots. Supplemental pollination significantly increased fruit set and seed set of individuals in pure plots, while it only marginally increased seed set per fruit of plants in mixed plots. Conclusions The presence of V. dichroantha can facilitate pollination and increase female reproductive success of P. monbeigiana via both quantity (mitigating pollinator limitation) and quality (reducing geitonogamy) effects. This study suggests that successive pollinator movements among flowers within a plant, as well as pollinator visitation rates and interspecific flower switching, may be important determinants of the direction and mechanisms of interaction between species.