Factors affecting pollinator movement and plant fitness in a specialized pollination system

The rate of pollen exchange within and among flowers may depend on pollinator attraction traits such as floral display size and flowering plant density. Variations in these traits may influence pollinator movements, pollen receipt, and seed number. To assess how floral display size and flowering plant density affect parameters of pollinator visitation rate, pollen receipt per flower, seed number per fruit and the between-plant pollinator movements, we studied the self-incompatible plant, Nierembergia linariifolia. Per-flower pollinator visitation rate and bout length increased linearly with increasing floral display size. Pollen receipt per flower increased linearly with increasing flowering plant density. For seed number per fruit, a polynomial model describing an increased seed number per fruit at low density and a decreased seed number per fruit at high density provided a significant fit. Per-flower pollinator visitation rate was not associated with pollen receipt per flower and seed number per fruit. Bees visited plants located near to the center of the population more frequently than plants located at the periphery. Increases in both floral display size and flowering plant density led to an increased chance of a plant being chosen as the center of the pollinator foraging area. These results suggest that even though large floral displays and high flowering plant density are traits that attract more pollinators, they may also reduce potential mate diversity by restricting pollen movement to conspecific mates that are closely located.     

Population Status and Reproductive Success of an Endangered Epiphytic Orchid in a Fragmented Landscape

Habitat fragmentation and disturbance are two of the most significant drivers of species extinctions in plant populations. The degree of impact of fragmentation on plant populations depends on the level of specificity of plant–animal interactions, as well as on the availability of suitable sites for seedling recruitment. In this study, we describe the population density and structure, pollen limitation and reproductive success of the endangered tropical orchid Myrmecophila christinae, an epiphytic species with a specialized pollination system. We surveyed a total of 14 populations located in a fragmented landscape. Seedling density was related to habitat disturbance and host plant density; while density of juveniles was related to density of adults. Adult and total individual densities were related to habitat affectation. We also found that fragments <1 ha had significantly fewer seedlings, as well as an over‐representation of large adults. On the other hand, fruit production was higher in fragments >10 ha, and fruit set was significantly lower in highly disturbed fragments. Hand pollination experiments showed that M. christinae was pollen limited in all the studied populations, suggesting that pollen limitation is unrelated to habitat disturbance. Overall, our results suggest that fragmentation has affected key demographic features of M. christinae, including reproduction and recruitment.     

Pollinator population size and pollination ecosystem service responses to enhancing floral and nesting resources

Modeling pollination ecosystem services requires a spatially explicit, process‐based approach because they depend on both the behavioral responses of pollinators to the amount and spatial arrangement of habitat and on the within‐ and between‐season dynamics of pollinator populations in response to land use. We describe a novel pollinator model predicting flower visitation rates by wild central‐place foragers (e.g., nesting bees) in spatially explicit landscapes. The model goes beyond existing approaches by: (1) integrating preferential use of more rewarding floral and nesting resources; (2) considering population growth over time; (3) allowing different dispersal distances for workers and reproductives; (4) providing visitation rates for use in crop pollination models. We use the model to estimate the effect of establishing grassy field margins offering nesting resources and a low quantity of flower resources, and/or late‐flowering flower strips offering no nesting resources but abundant flowers, on bumble bee populations and visitation rates to flowers in landscapes that differ in amounts of linear seminatural habitats and early mass‐flowering crops. Flower strips were three times more effective in increasing pollinator populations and visitation rates than field margins, and this effect increased over time. Late‐blooming flower strips increased early‐season visitation rates, but decreased visitation rates in other late‐season flowers. Increases in population size over time in response to flower strips and amounts of linear seminatural habitats reduced this apparent competition for pollinators. Our spatially explicit, process‐based model generates emergent patterns reflecting empirical observations, such that adding flower resources may have contrasting short‐ and long‐term effects due to apparent competition for pollinators and pollinator population size increase. It allows exploring these effects and comparing effect sizes in ways not possible with other existing models. Future applications include species comparisons, analysis of the sensitivity of predictions to life‐history traits, as well as large‐scale management intervention and policy assessment.     

Beetle pollination of Luisia teres (Orchidaceae) and implications of a geographic divergence in the pollination system

The epiphytic orchid Luisia teres on Okinawa Island, the Ryukyus, Japan, sexually attracts male Protaetia beetles for pollination through floral scent. Here, the pollination system of L. teres on the Satsuma Peninsula, Kyushu, located approximately 600 km north of Okinawa Island, is reported. The inconspicuous flowers depend on the Cetoniinae beetles Protaetia orientalis submarumorea and Pseudotorynorrhina japonica, which both usually feed on fermented sap/ripe fruits, for pollination. The flower-visiting P. orientalis submarumorea beetles included both males and females, and pseudocopulation was never observed. Pollinaria removal and pollination were achieved while beetles fed on floral nectar. Although P. orientalis submarumorea did not necessarily remove pollinaria more frequently than Ps. japonica, the pollinaria attached precisely to the central part of their clypeus, unlike on Ps. japonica. Our findings suggest that a geographic divergence in the pollination system occurs in Japanese L. teres. In the Ryukyus, the orchid adopts a specialized pollination system with only male-attractive olfactory stimuli, whereas in Kyushu it has a relatively more generalized system based on only a food reward.     

Low genetic variation reduces cross-compatibility and offspring fitness in populations of a narrow endemic plant with a self-incompatibility system

Genetic variation was shown earlier to bereduced in smaller populations of the narrowendemic putatively self-incompatible Cochlearia bavarica. To test whether thisnegatively affects plant fitness by reducedavailability of compatible mates and byinbreeding depression, we studied effects ofpopulation size and pollination treatments oncross-compatibility and offspring fitness in 16isolated populations of this plant. After openpollination, compatibility of crosses (i.e.,whether at least one fruit developed per markedflower), fruit set of compatible crosses, andcumulative fitness (number of plants permaternal ovule) after 14 months in a commongarden were lower for plants from smallerpopulations. Throughout the study, cumulativefitness was lower after hand pollination withpollen of one donor than after open pollination(finally 73.4% lower), suggesting that severalpollen donors or single pollen donors of higherquality are involved in open pollination.Moreover, cumulative fitness was lower afterhand selfing than after hand outcrossing(finally 69.4% lower), indicating bothinbreeding depression and reduced compatibilityafter selfing. High self-compatibility(40.6%), dry stigmas, and differences in thecompatibility of 11 of 33 experimentalreciprocal crosses between plant pairsconfirmed that C. bavarica has asporophytic self-incompatibility system, as iscommon in the Brassicaceae. Our studydemonstrates, that plants in smallerpopulations of species with a sporophyticself-incompatibility system can experiencetwofold fitness reductions associated withreduced genetic variability, i.e., twofoldgenetic Allee effects: via reducedcross-compatibility and via reduced offspringfitness.     

Looks matter: changes in flower form affect pollination effectiveness in a sexually deceptive orchid

Many species of the sexually deceptive genus Ophrys are characterized by insect‐like flowers. Their form has been traditionally considered to play an important role in pollinator attraction and manipulation. Yet, the evolution of the floral form remains insufficiently understood. We hypothesize that pollinator‐mediated selection is essential for driving floral form evolution in Ophrys, but that form components are being subjected to varying selection pressures depending on their role in mediating interactions with pollinators. By using the Eucera‐pollinated Ophrys leochroma as a model, our aim has been to assess whether and in what manner pollination effectiveness is altered by experimental manipulation of the flower form. Our results show that floral form plays an essential and, so far, underestimated role in ensuring effective pollination by mechanically guiding pollinators towards the reproductive structures of the flower. Pollinators are significantly less effective in interacting with flowers having forms altered to resemble those of species pollinated by different hymenopteran genera. Further, those components used by pollinators as gripping points were found to be more effective in ensuring pollinia transfer than those with which pollinators do not directly interact. Thus, mechanically active and inactive components appear to be under different selection pressures. As a consequence, mechanically active components of the flower form could reflect adaptations to the interaction with particular pollinator groups, whereas mechanically inactive components can vary more freely. Disentangling selection patterns between the functionally different components of flower form may provide valuable insights into the mechanisms driving the morphological diversification of sexually deceptive pollination systems.     

On the roles of colour and scent in a specialized floral mimicry system

Background and Aims

Sexually deceptive orchids achieve cross-pollination by mimicking the mating signals of female insects, generally hymenopterans. This pollination mechanism is often highly specific as it is based primarily on the mimicry of mating signals, especially the female sex pheromones of the targeted pollinator. Like many deceptive orchids, the Mediterranean species Ophrys arachnitiformis shows high levels of floral trait variation, especially in the colour of the perianth, which is either green or white/pinkinsh within populations. The adaptive significance of perianth colour polymorphism and its influence on pollinator visitation rates in sexually deceptive orchids remain obscure.

Methods

The relative importance of floral scent versus perianth colour in pollinator attraction in this orchid pollinator mimicry system was evaluated by performing floral scent analyses by gas chromatography-mass spectrometry (GC-MS) and behavioural bioassays with the pollinators under natural conditions were performed.

Key Results

The relative and absolute amounts of behaviourally active compounds are identical in the two colour morphs of O. arachnitiformis. Neither presence/absence nor the colour of the perianth (green versus white) influence attractiveness of the flowers to Colletes cunicularius males, the main pollinator of O. arachnitiformis.

Conclusion

Chemical signals alone can mediate the interactions in highly specialized mimicry systems. Floral colour polymorphism in O. arachnitiformis is not subjected to selection imposed by C. cunicularius males, and an interplay between different non-adaptive processes may be responsible for the maintenance of floral colour polymorphism both within and among populations.     

Flight morphology corresponds to both surrounding landscape structure and local patch conditions in a highly specialized peatland butterfly (Lycaena epixanthe)

1. Movement mediates the response of populations and communities to landscape and habitat spatial structure, yet movement capability may itself be modified by selection pressures accompanying landscape change. Insect flight morphology can be affected by both the landscape surrounding habitat patches and the distribution of resources within habitat patches. 2. This study investigated the relative influence of local habitat patch conditions and surrounding landscape structure on variation in morphological traits associated with flight in the bog copper (Lycaena epixanthe), a butterfly endemic to temperate Nearctic peatlands. 3. Eight habitat patches were sampled to assess the influence of the surrounding landscape (connectivity of potential habitat and matrix composition) and patch size (an integrated proxy of resource density and spatial distribution) on investment into flight, measured by thorax and abdomen mass, and wing area. 4. The results revealed an effect of both local habitat conditions and landscape structure on flight‐related morphological traits. Increasing forest cover in the surrounding landscape, indicative of increased habitat patch isolation, corresponded with less mobile phenotypes in both sexes. Surrounding landscapes with more water were also generally associated with less mobile phenotypes. Investment into flight was greater in smaller peatlands in which host plant density is higher and more homogeneously distributed. 5. The present study highlights that morphological traits associated with mobility may be responding to both local habitat patch characteristics and surrounding landscape structure. It also supports the hypothesis that local habitat conditions contribute to morphological variation in butterflies.     

Life history traits as predictors of plant rarity, with particular reference to hemiparasiticOrobanchaceae

We made a comparison between life history and reproductive characteristics of a group of Dutch rare (30) and common (105) plant species, all dicotyledonous and insect-visited forbs. The traits life span, clonality, breeding system, seed production, seed dispersal, and soil seed bank longevity are considered. All trait values have been ranked according to their possible effect on the vulnerability of a species for extinction, where low values indicate a low risk and high values a high risk for a species. Rare and common plants differed significantly in four traits: seed production, breeding system, seed bank longevity and clonality. The discriminant analysis showed that 79% of the 135 species was correctly classified in the group they presently belong to. Especially species that are rare now but were much more common 50–100 years ago were classified as being common species, pointing at the detrimental effect of habitat loss these species encountered at which they have not yet been able to adapted to. The data set include eight hemiparasitic plant species (familyOrobanchaceae), of which six are endangered in The Netherlands and two more common. Both rare and common hemiparasites scored high traits values, indicating that they are vulnerable for extinction. The hemiparasites had several characteristics in common: they are all annuals or biennials that have no means of clonal reproduction. They all have non-clustered, zygomorphic flowers that produce nectar and have a precise pollen presentation and are mainly visited by (bumble) bees. Mean life span was significantly shorter in the eight hemiparasiticOrobanchaceae than in the non-hemiparasiticScrophulariaceae used for comparison (4 species). Overall, we concluded that hemiparasitic plants have a special combination of life history traits which makes them vulnerable for local and regional extinction.     

The Hemipepsis wasp‐pollination system in South Africa: a comparative analysis of trait convergence in a highly specialized plant guild

Pollinator‐mediated convergence in floral traits is the fundamental basis for pollination syndromes, but it has seldom been rigorously analysed. Here we synthesize information on a guild of South African plants that are pollinated by functionally similar pompilid wasps in the genus Hemipepsis and investigate the extent of trait convergence in guild members. The guild includes members from three plant families (Apocynaceae, Orchidaceae and Asparagaceae subfamily Scilloideae) and contains remarkably high levels of functional specialization with 18 of the 23 known guild members being pollinated exclusively by Hemipepsis wasps. The distribution of the guild is centred in the moist upland grasslands of eastern South Africa. Qualitative similarities among guild members include dull greenish‐ or brownish‐white flowers, often with purple blotches, mid‐summer flowering, sweet spicy scent and exposed nectar. To assess the extent of convergent evolution within the guild, we compared floral traits of guild members with those of congeneric non‐wasp‐pollinated species. Guild members typically produce moderate volumes (> 4 µL per flower per day) of concentrated (> 50% sugar by weight) sucrose‐dominant nectar. The nectar properties of guild members did not, however, differ significantly from those of congeneric species pollinated by other vectors. Non‐metric multidimensional scaling of scent data for 15 guild members and 17 congeners (obtained through gas chromatography–mass spectrometry of headspace samples and supplemented with published data) yielded little evidence for convergent evolution in the overall scent composition of guild members. However, convergence in floral spectral reflectance was evident in the guild members; in particular, loci for colours of guild members were significantly closer to the guild centroid than loci for colours of congeners, and they formed a distinct cluster in the blue to blue–green region of the hymenopteran colour hexagon. The colours of guild members were also significantly closer to the colour of background vegetation than those of congeneric species, suggesting a role for cryptic colouring in this system. These results confirm convergence in the floral colours of plants that are pollinated by Hemipepsis spider‐hunting wasps, but also suggest that other traits, such as nectar properties, do not necessarily evolve during shifts between pollination systems. Identification of particular scent compounds and non‐sugar nectar constituents that influence wasp behaviour will be essential for illuminating the extent of biochemical convergence in the guild members. © 2012 The Linnean Society of London, Botanical Journal of the Linnean Society, 2012, 168 , 278–299.     

Pollinator functional response and plant population dynamics: Pollinators as a limiting resource

Summary During recent years, much work has focused on which factors limit the reproductive success in plants. Several studies show a strong influence of either resource limitation, pollen limitation or a combined effect of both. The theoretical arguments for resource limitation are abundant, but there has been very little work done concerning the effect of pollinator availability. In this paper we construct a model to study how the reproductive success in plants is influenced by the foraging behaviour of the pollinators. The pollinator population is assumed to have a constant population density. A functional response function for the pollinators is derived. It is similar to a Holling type II functional response. It is shown that, since the pollinators are regulated by factors not included in the model and their capability to pollinate is limited by the functional response, this is sufficient for regulating the plant population. There also exists a threshold condition for the persistence of the plant population that depended on the search rate of the pollinators and the demographic parameters of the plant population. If this threshold condition is not satisfied the plant population cannot persist and will become extinct. If the condition is satisfied the plant population grows until it is limited at the equilibrium mentioned above.     

Managing ski resorts to improve biodiversity conservation: Australian reptiles as a case study

Alpine/subalpine environments are diverse systems that support many endemic species. Worldwide, these ecosystems are under threat from ski resort disturbances – even in areas broadly designated for biodiversity conservation. The effects of ski resorts on reptiles are largely unknown, making it difficult to implement effective conservation actions. Many ski resorts do not currently address the needs of reptiles, even those listed as threatened, in their management plans. If reptiles are to continue inhabiting ski resorts in Australia, strategies must be implemented that target their conservation. To begin to address this problem, we summarise current research investigating the effects of ski resorts on reptiles. Based on this information, we recommend strategies that will enhance the conservation of reptiles in areas affected by ski‐related disturbances. Suggested strategies include (i) restricting intensive disturbances to already highly modified areas of Australian ski resorts, (ii) avoiding disturbance of remaining native vegetation and structural complexity in ski resorts and (iii) re‐establishing structural complexity at highly modified sites through revegetation programmes, or through the cessation of mowing during peak reptile activity periods. While these strategies are designed to facilitate the persistence of reptiles in ski resorts, their long‐term success can only be evaluated by monitoring their effectiveness.     

TheMoegistorhynchus longirostris (Diptera: Nemestrinidae) pollination guild: long-tubed flowers and a specialized long-proboscid fly pollination system in southern Africa

A guild of 20 late spring- and early summer-flowering species ofIridaceae, Geraniaceae andOrchidaceae is pollinated partly or exclusively by the long-proboscid flyMoegistorhynchus longirostris (Nemestrinidae). This large-bodied fly, active in late spring and early summer, has mouthparts 40–70 (90) mm long and forages for nectar from a variety of species. These plants share a suite of convergent floral features including a straight or weakly curved floral tube usually 50–70 mm long but sometimes to 90 mm, relatively short petals or tepal lobes coloured white, cream or salmon with reddish nectar guides, and often violet or red anthers and pollen. Flowers of most species with these characteristics are zygomorphic with the stamens either arcuate (mostIridaceae) or declinate (Geraniaceae and someIridaceae). The flowers are odourless and typically secrete large amounts of nectar of relatively constant sugar concentration, mostly 24–29%, with a high sucrose:hexose ratio. Guild members utilize five separate sites of pollen deposition on the body of the fly, typically utilizing different deposition sites when two or more co-occur, indicating strong selection to aviod pollen contamination.M. longirostris is restricted to the west coast of southern Africa and at least 8 species appear to depend exclusively on the insect for pollination. The remaining species in the guild are pollinated by one or both of the long-proboscid fliesPhiloliche gulosa andP. rostrata (Tabanidae) over other parts of their range. Species and races pollinated entirely byM. longirostris have longer floral tubes which makes nectar unavailable to other insects, including other species of long-proboscid fly. The only insect with mouthparts long enough to forage effectively on these long-tubed flowers isM. longirostris and this fly must be considered a keystone species in the ecosystems in which it occurs.Dedicated to emer. Univ.-Prof. DrFriedrich Ehrendorfer on the occasion of his 70th birthday     

扩散对破碎化景观上宿主-寄生种群动态的影响

景观破碎化和扩散是空间种群模型的重要因素,对生物入侵存在着深远的影响。本章将基于偶对近似模型,探讨由局部和全局宿主-寄生相互作用共同决定的扩散模式对破坏性景观上疾病入侵与传播的影响。其中,生境破坏由生境丧失量与生境破碎化程度来描述。模拟结果显示,宿主和病毒的全局扩散对疾病的入侵与种群密度产生不对称效应：病毒的全局扩散对系统产生的影响较宿主的全局扩散更为显著。不同扩散模式下,生境丧失越高或破碎化程度越低,均将越有害于寄生病毒的入侵;同时,生境的破坏程度也显著地影响了入侵阈值对扩散模式的响应机制。本文研究结果暗示,景观破碎化的空间分布格局以及病毒扩散的限制均可作为物种保护与管理中有效的疾病控制策略。该研究结果在一定意义上丰富和发展了寄生感染理论,为物种保护提供了生态学理论依据。     

Patch isolation and shape predict plant functional diversity in a naturally fragmented forest

Aims It is known that taxonomic diversity can be predicted by the spatial configuration of the habitat, in particular by its area and degree of isolation. However, taxonomic diversity is a poor predictor of ecosystem functioning. While functional diversity is strongly linked to the functionality and stability of ecosystems, little is known about how changes in the spatial configuration of the habitat affect functional diversity. In this study, we evaluated whether the spatial configuration of forest patches predicts the functional diversity of plants in a fragmented forest.Methods Five functional leaf traits (leaf dry matter content, leaf punch force, specific leaf area, leaf size and leaf thickness) were measured for 23 dominant plant species in 20 forest patches in a naturally fragmented forest on the Yucatan Peninsula. Abundance-weighted multivariate and individual trait metrics of functional diversity were calculated and correlated with size, degree of isolation and the shape of forest patches.Important findings Patch shape was negatively correlated with multivariate and individual trait (leaf dry matter content and leaf size) metrics of functional diversity. Patch isolation measures were also negatively correlated with individual trait (leaf dry matter content, leaf punch force and leaf size) metrics of functional diversity. In other words, greater patch shape irregularity and isolation degree impoverish plant functional variability. This is the first report of the negative effects of patch shape irregularity and isolation on the functional diversity of plant communities in a forest that has been fragmented for a long time.     

Pollinator‐mediated selection in a specialized pollination system: matches and mismatches across populations

Most studies on pollinator‐mediated selection have been performed in generalized rather than specialized pollination systems. This situation has impeded evaluation of the extent to which selection acts on attraction or specialized key floral traits involved in the plant‐pollinator phenotypic interphase. We studied pollinator‐mediated selection in four populations of Nierembergia linariifolia, a self‐incompatible and oil‐secreting plant pollinated exclusively by oil‐collecting bees. We evaluated whether floral traits experience variable selection among populations and whether attraction and fit traits are heterogeneously selected across populations. Populations differed in every flower trait and selection was consistently observed for corolla size and flower shape, two traits involved in the first steps of the pollination process. However, we found no selection acting on mechanical‐fit traits. The observation that selection occurred upon attraction rather than mechanical‐fit traits, suggests that plants are not currently evolving fine‐tuned morphological adaptations to local pollinators and that phenotypic matching is not necessarily an expected outcome in this specialized pollination system.