Amparo Lázaro

How does climate warming affect plant‐pollinator interactions?

Climate warming affects the phenology, local abundance and large-scale distribution of plants and pollinators. Despite this, there is still limited knowledge of how elevated temperatures affect plant-pollinator mutualisms and how changed availability of mutualistic partners influences the persistence of interacting species. Here we review the evidence of climate warming effects on plants and pollinators and discuss how their interactions may be affected by increased temperatures. The onset of flowering in plants and first appearance dates of pollinators in several cases appear to advance linearly in response to recent temperature increases. Phenological responses to climate warming may therefore occur at parallel magnitudes in plants and pollinators, although considerable variation in responses across species should be expected. Despite the overall similarities in responses, a few studies have shown that climate warming may generate temporal mismatches among the mutualistic partners. Mismatches in pollination interactions are still rarely explored and their demographic consequences are largely unknown. Studies on multi-species plant-pollinator assemblages indicate that the overall structure of pollination networks probably are robust against perturbations caused by climate warming. We suggest potential ways of studying warming-caused mismatches and their consequences for plant-pollinator interactions, and highlight the strengths and limitations of such approaches.

The relationships between floral traits and specificity of pollination systems in three Scandinavian plant communities

The pollination syndrome hypothesis has provided a major conceptual framework for how plants and pollinators interact. However, the assumption of specialization in pollination systems and the reliability of floral traits in predicting the main pollinators have been questioned recently. In addition, the relationship between ecological and evolutionary specialization in pollination interactions is still poorly understood. We used data of 62 plant species from three communities across southern Norway to test: (1) the relationships between floral traits and the identity of pollinators, (2) the association between floral traits (evolutionary specialization) and ecological generalization, and (3) the consistency of both relationships across communities. Floral traits significantly affected the identity of pollinators in the three communities in a way consistent with the predictions derived from the pollination syndrome concept. However, hover flies and butterflies visited flowers with different shapes in different communities, which we mainly attribute to among-community variation in pollinator assemblages. Interestingly, ecological generalization depended more on the community-context (i.e. the plant and pollinator assemblages in the communities) than on specific floral traits. While open yellow and white flowers were the most generalist in two communities, they were the most specialist in the alpine community. Our results warn against the use of single measures of ecological generalization to question the pollination syndrome concept, and highlight the importance of community comparisons to assess the pollination syndromes, and to understand the relationships between ecological and evolutionary specialization in plant–pollinator interactions.

A Phylogeographic Split in Buxus balearica (Buxaceae) as Evidenced by Nuclear Ribosomal Markers: When ITS Paralogues Are Welcome

Sequences from the ribosomal nuclear internal transcribed spacers (ITS) have been widely used to infer evolutionary hypotheses across a broad range of living organisms. Intraspecific sequence variation is assumed to be absent or negliable in most species, but few detailed studies have been conducted to assess the apportionment of ITS sequence variation within and between plant populations. Buxus balearica was chosen as a model species to assess the levels of infraspecific and intragenomic ITS variation in rare and endangered species occurring in disjunct populations around the Mediterranean basin. Intragenomic polymorphic sites were detected for western and eastern accessions of B. balearica and in two accessions of the sister species B. sempervirens. Overall, 19 different ribotypes were found in B. balearica after sequencing 48 clones, whereas 15 ribotypes were detected in 19 clones of B. sempervirens. The integrity and secondary structure stability of the ribosomal sequences suggest that they are not pseudogenes. The high number of ribotypes recovered through cloning suggested that some sequences could be chimeric or generated in vivo by partial homogenization through gene conversion or unequal crossing-over. Average sequence divergence among B. balearica clones was 0.768%, and the most divergent sequences differed by 1.62%. Available evidence does not suggest that B. balearica paralogues have been obtained from other extant Buxus species through interspecific hybridization. The presence of several ribosomal sequences in box implies that the molecular forces driving the concerted evolution of this multigene family are not fully operational in this genus. Phylogenetic analyses of cloned ITS sequences from B. balearica displayed very poor resolution and only two clades received moderate bootstrap support. Despite the marked intragenomic sequence divergence found, ribosomal data suggest a clear phylogeographic split in B. balearica between western and eastern accessions. The distinct, nonchimeric sequences that are postulated as being present in each biogeographic group suggest that box populations from Anatolia (eastern Mediterranean) are relict.

Reproductive success of the endangered shrub Buxus balearica Lam. (Buxaceae): pollen limitation, and inbreeding and outbreeding depression

The increasingly common phenomenon of habitat fragmentation raises the probability of pollination failure in a number of species, as both pollen quantity and quality often decrease as populations become isolated. We experimentally investigated whether pollen was limiting reproductive success of the endangered shrub Buxus balearica in five populations, two continental and three insular, during 2002 and 2003. Pollen limitation varied among populations and years, but such variation was not related to density or degree of isolation. All populations showed inbreeding depression at different phases of the reproductive cycle, although its effects differed greatly among sites. Between-population outcrossing did not have a consistent effect on several components of fitness. The highest levels of inbreeding depression – detected at the level of fruit and seed set- occurred at the smallest and least fecund populations from each region. This indicates that further fragmentation of the populations of this already endangered species could certainly threaten their survival.

Local floral composition and the behaviour of pollinators: attraction to and foraging within experimental patches.

1. Understanding how foraging decisions take place at the local scale is relevant because they may directly affect the fitness of individual plants. However, little is known about how local diversity and density affect the foraging behaviour of most pollinator groups.

Does the spatial variation in selective pressures explain among-site differences in seed mass? A test with Buxus balearica

Positive effects of seed size on germination and survival can be offset by a greater probability of predation or a poorer dispersal of larger seeds. We hypothesized that spatial variation in local selective pressures acting on seed mass may lead to differences in both optimal and observed seed mass among discrete populations. We first examined the variation in seed mass across a total of 14 populations of Buxus balearica; in six of such populations, we further examined the temporal variation by measuring seed mass during three consecutive years. Second, we evaluated seed mass effects on different recruitment phases (seed rain to seedling establishment) in three populations. Lastly, for these three populations, we estimated the resulting overall phenotypic selection on seed mass during recruitment and compared the observed and the predicted optimal seed masses. Most variation (c. 70%) in seed mass occurred among populations, and although we found inter-annual variation in seed mass, the differences in seed mass among populations were consistent over time. Conflicting selective pressures on seed mass appeared during recruitment, and their direction and strength varied among populations, depending on the relative local importance of seed predation vs. germination and establishment. Observed seed mass matched predicted optimal seed mass in two of the three examined populations, suggesting local adaptative responses to the spatial mosaic of selective pressures.

How do pollinator visitation rate and seed set relate to species’ floral traits and community context?

Differences among plant species in visitation rate and seed set within a community may be explained both by the species’ floral traits and the community context. Additionally, the importance of species’ floral traits vs. community context on visitation rate and seed set may vary among communities. In communities where the pollinator-to-flower ratio is low, floral traits may be more important than community context, as pollinators may have the opportunity to be choosier when visiting plant species. In this study we investigated whether species’ floral traits (flower shape, size and number, and flowering duration) and community context (conspecific and heterospecific flower density, and pollinator abundance) could explain among-species variation in visitation rate and seed set. For this, we used data on 47 plant species from two Norwegian plant communities differing in pollinator-to-flower ratio. Differences among species in visitation rate and seed set within a community could be explained by similar variables as those explaining visitation rate and seed set within species. As expected, we found floral traits to be more important than community context in the community with a lower pollinator-to-flower ratio; whereas in the community with a higher pollinator-to-flower ratio, community context played a bigger role. Our study gives significant insights into the relative importance of floral traits on species’ visitation rate and seed set, and contributes to our understanding of the role of the community context on the fitness of plant species.

Moderation is best: Effects of grazing intensity on plant-flower visitor networks in Mediterranean communities

The structure of pollination networks is an important indicator of ecosystem stability and functioning. Livestock grazing is a frequent land use practice that directly affects the abundance and diversity of flowers and pollinators and, therefore, may indirectly affect the structure of pollination networks. We studied how grazing intensity affected the structure of plant-flower visitor networks along a wide range of grazing intensities by sheep and goats, using data from 11 Mediterranean plant-flower visitor communities from Lesvos Island, Greece. We hypothesized that intermediate grazing might result in higher diversity as predicted by the Intermediate Disturbance Hypothesis, which could in turn confer more stability to the networks. Indeed, we found that networks at intermediate grazing intensities were larger, more generalized, more modular, and contained more diverse and even interactions. Despite general responses at the network level, the number of interactions and selectiveness of particular flower visitor and plant taxa in the networks responded differently to grazing intensity, presumably as a consequence of variation in the abundance of different taxa with grazing. Our results highlight the benefit of maintaining moderate levels of livestock grazing by sheep and goats to preserve the complexity and biodiversity of the rich Mediterranean communities, which have a long history of grazing by these domestic animals.

Effects of grazing intensity on pollinator abundance and diversity, and on pollination services

1. Pollinating insects provide important ecosystem services and are influenced by the intensity of grazing. Based on the Intermediate Disturbance Hypothesis (IDH), pollinator diversity is expected to peak at intermediate grazing intensities. However, this hump‐shaped relationship is rarely found.

Effects of experimentally simulated pollinator decline on recruitment in two European herbs

Summary1. Pollinator decline causes great concern because reduced pollinator availability may negativelyaffect plants’ seed production and in turn influence seedling recruitment and densities of plant popu-lations.2. We used a novel experimental approach that simulates pollinator decline to assess the effects of a4-year reduction in pollinator availability on reproduction and recruitment in two common Europeanherbs, Centaurea jacea and Leucanthemum vulgare.3. The experiment strongly reduced visitation rates (by 59%) and seed set (by 23%) in Leucanthe-mum, but this did not cause reduced seedling recruitment, suggesting that other factors, such aswater availability or the existence of a seed bank, are important for recruitment in this population ofLeucanthemum.4. The experiment also significantly reduced visitation rates to Centaurea (by 33%), but did notaffect seed set or seedling recruitment. In this species, however, seed set increased with visitationrates and local recruitment increased with seed production, indicating that a consistent pollinatordecline may negatively affect Centaurea population densities.5. Synthesis. Our experiment reduced pollinator availability without artificially affecting qualitycomponents of the pollination process. Therefore, this approach may represent a realistic simulationof a pollinator decline. To our knowledge, this is the first study that uses an experimental reductionin pollinator visitation to assess its effects on population recruitment in plants. This study shows thata large pollinator decline may affect population densities, but only in populations where seedlingrecruitment is limited by seed production. Our results also highlight the importance of going beyondvisitation rates and seed production to study the impact of pollinator decline on plant populations.Key-words: Centaurea jacea, Leucanthemum vulgare, pollinator loss, reproductive ecology, seedset, seed weight, seedling density, visitation rateIntroduction