Flowering phenology: An example of relaxation of natural selection?

Flowering phenology has normally been viewed as fundamental to a plant species' reproductive ecology. Researchers in the field have emphasized the adaptive importance of flowering at a particular time relative to other individuals in the population, or other species in the community. An alternative view, however, is that flowering phenology is a trait that may not be under strong selection, and this may have allowed some variation to appear in populations by chance.     

Finding NEMO: nestedness engendered by mutualistic organization in anemonefish and their hosts

The interaction structure of mutualistic relationships, in terms of relative specialization of the partners, is important to understanding their ecology and evolution. Analyses of the mutualistic interaction between anemonefish and their host sea anemones show that the relationship is highly nested in structure, generalist species interacting with one another and specialist species interacting mainly with generalists. This supports the hypothesis that the configuration of mutualistic interactions will tend towards nestedness. In this case, the structure of the interaction is at a much larger scale than previously hypothesized, across more than 180 degrees of longitude and some 60 degrees of latitude, probably owing to the pelagic dispersal capabilities of these species in a marine environment. Additionally, we found weak support for the hypothesis that geographically widespread species should be more generalized in their interactions than species with small ranges. This study extends understanding of the structure of mutualistic relationships into previously unexplored taxonomic and physical realms, and suggests how nestedness analysis can be applied to the conservation of obligate species interactions.     

Environmental control of reproductive phenology and the effect of pollen supplementation on resource allocation in the cleistogamous weed, Ruellia nudiflora (Acanthaceae)

• Background and Aims Mixed reproductive strategies may have evolved as a response of plants to cope with environmental variation. One example of a mixed reproductive strategy is dimorphic cleistogamy, where a single plant produces closed, obligately self-pollinated (CL) flowers and open, potentially outcrossed (CH) flowers. Frequently, optimal environmental conditions favour production of more costly CH structures whilst economical and reliable CL structures are produced under less favourable conditions. In this study we explore (1) the effect of light and water on the reproductive phenology and (2) the effect of pollen supplementation on resource allocation to seeds in the cleistogamous weed Ruellia nudiflora.• Methods Split-plot field experiments were carried out to assess the effect of shade (two levels: ambient light vs. a reduction of 50 %) and watering (two levels: non-watered vs. watered) on the onset, end and duration of the production of three reproductive structures: CH flowers, CH fruit and CL fruit. We also looked at the effect of these environmental factors on biomass allocation to seeds (seed weight) from obligately self-pollinated flowers (CL), open-pollinated CH flowers and pollen-supplemented CH flowers.• Key Results CH structures were produced for a briefer period and ended earlier under shaded conditions. These conditions also resulted in an earlier production of CL fruit. Shaded conditions also produced greater biomass allocation to CH seeds receiving extra pollen.• Conclusions Sub-optimal (shaded) conditions resulted in a briefer production period of CH structures whilst these same conditions resulted in an earlier production of CL structures. However, under sub-optimal conditions, plants also allocated more resources to seeds sired from CH flowers receiving large pollen loads. Earlier production of reproductive structures and relatively larger seed might improve subsequent success of CL and pollen-supplemented CH seeds, respectively.     

Landscape structure influences pollinator movements and directly affects plant reproductive success

Pollinators play a key role within most terrestrial communities in maintaining plant populations, as well as pollinating many agricultural crops for seed and fruit production. The mobility of pollinating animals is significant to their importance but we know little about how landscape structure influences pollinator movements. Linear landscape features such as hedgerows and embankments are conspicuous features of agricultural landscape structure and are important artificial habitats in their own right. However, there has been some debate as to the function of these landscape elements as corridors between larger expanses of semi‐natural habitat separated by urban and agricultural habitats. Few studies have specifically studied insect flight responses to linear landscape elements. By observing bumblebee flight behaviour along hedgerows and, by creating a medium‐scale experimental array of flower patches using an artificial linear feature, we examined whether such structures can elicit an oriented flight response along them and therefore facilitate insect movement through the landscape. We found that both hedgerows and artificial linear landscape features can influence the flight directions of bumblebees (Bombus spp. Hymenoptera, Apidae), one of Europe's most important groups of pollinators. A bioassay experiment in which Salvia pratensis (Lamiaceae) was planted into landscape patches with differing numbers of connecting hedgerows showed that this directional response can have a profound effect on plant reproductive success – plants had increased pollinator activity, pollen receipt and subsequent seed set in patches with more connections. The overall hedgerow connectedness of a landscape is therefore important both to bumblebee movement and to those plants which depend on bumblebees for pollination services.     

Evidence for stabilising selection acting on flowering time in Arum maculatum (Araceae): the influence of phylogeny on adaptation

The relationship between flowering time and reproductive success was investigated in the fly-pollinated, monoecious perennial herb Arum maculatum L. (Araceae). This species temporarily traps its principle pollinator, a psychodid midge. Probability of fruit set was analysed in relation to early, peak and late periods of the flowering phenology of four British populations between 1992 and 1997. In three out of five cases, plants which flowered during early and late periods were significantly less likely to set fruit. In addition, one population showed a similar relationship for percentage fruit set of individual inflorescences, and seeds from peak-flowering plants were significantly heavier. There was no variation in number of female flowers per inflorescence over the flowering season. Probability of fruit set appears to be mediated by the likelihood of trapping psychodid midges that have previously been trapped and picked up pollen, an unlikely event during early and late flowering periods when few inflorescences are open. The majority of plants in all populations produce only one inflorescence which means that timing of flowering may be crucial to reproductive success. We interpret our findings as evidence that stabilising selection may be acting on some populations and/or during some years. The ultimate cause, however, can be related to the very short (12–18 h) female phase of each inflorescence, a phylogenetically conservative trait within the Araceae. Received: 19 August 1998 / 15 February 1999     

How many flowering plants are pollinated by animals?

It is clear that the majority of flowering plants are pollinated by insects and other animals, with a minority utilising abiotic pollen vectors, mainly wind. However there is no accurate published calculation of the proportion of the ca 352 000 species of angiosperms that interact with pollinators. Widely cited figures range from 67% to 96% but these have not been based on firm data. We estimated the number and proportion of flowering plants that are pollinated by animals using published and unpublished community‐level surveys of plant pollination systems that recorded whether each species present was pollinated by animals or wind. The proportion of animal‐pollinated species rises from a mean of 78% in temperate‐zone communities to 94% in tropical communities. By correcting for the latitudinal diversity trend in flowering plants, we estimate the global number and proportion of animal pollinated angiosperms as 308 006, which is 87.5% of the estimated species‐level diversity of flowering plants. Given current concerns about the decline in pollinators and the possible resulting impacts on both natural communities and agricultural crops, such estimates are vital to both ecologists and policy makers. Further research is required to assess in detail the absolute dependency of these plants on their pollinators, and how this varies with latitude and community type, but there is no doubt that plant–pollinator interactions play a significant role in maintaining the functional integrity of most terrestrial ecosystems.     

Local and regional specialization in plant–pollinator networks

Specialization of species is often studied in ecology but its quantification and meaning is disputed. More recently, ecological network analysis has been widely used as a tool to quantify specialization, but here its true meaning is also debated. However, irrespective of the tool used, the geographic scale at which specialization is measured remains central. Consequently, we use data sets of plant–pollinator networks from Brazil and the Canary Islands to explore specialization at local and regional scales. We ask how local specialization of a species is related to its regional specialization, and whether or not species tend to interact with a non‐random set of partners in local communities. Local and regional specialization were strongly correlated around the 1:1 line, indicating that species conserve their specialization levels across spatial scales. Furthermore, most plants and pollinators also showed link conservatism repeatedly across local communities, and thus seem to be constrained in their fundamental niche. However, some species are more constrained than others, indicating true specialists. We argue that several geographically separated populations should be evaluated in order to provide a robust evaluation of species specialization.