Flower Colour Polymorphism, Pollination Modes, Breeding System and Gene Flow in Anemone coronaria.

The flower colour of Anemone coronaria (Ranunculaceae) is a genetically inherited trait. Such intra-specific flower colour polymorphism might be driven by pollinators, other non-pollinating agents, or by abiotic factors. We investigated the genetic relations among red, white and purple-blue flower colour morphs growing in 10 populations of A. coronaria in Israel, in relation to their breeding system, pollination modes, differential perception by bees and visitors' behaviour. Flowers of these three morphs differed in their reflectance that could be perceived by bees. Honeybees, solitary bees and flies demonstrated only partial preferences for the different colour morphs. No spontaneous self-pollination was found; however, fruit set under nets, excluding insects but allowing wind pollination, was not significantly lower than that of natural free pollinated flowers, indicating a potential role of wind pollination. Anemone coronaria flowers were visited by various insects, honeybees and Andrena sp. preferred the white and purple-blue morphs, while the syrphid flies preferred the white flowers. Thus, visitor behaviour can only partially explain the evolution or maintenance of the colour polymorphism. No significant genetic differences were found among the populations or colour morphs. Wind pollination, causing random gene flow, may explain why no significant genetic divergence was found among all studied populations and their colour morphs. The existence of monomorphic red populations, along other polymorphic populations, might be explained by linked resistance to aridity and/or grazing.

Genetic Structure of a Naturally Regenerating Post-Fire Seedling Population: Pinus halepensis As a Case Study.

To study the effects of wildfire on population genetics of a wind pollinated and wind dispersed tree, we have analyzed the genetic structure of a post-fire, naturally regenerating seedling population of Pinus halepensis Miller, on Mt. Carmel, Israel. We tested the existence of spatial genetic structure, which is expected due to the special spatial demographic structure of the post-fire seedling and sapling populations of this species. Explicitly, we asked whether or not seedlings that germinated under large, burned, dead pine trees are also their offspring. The results revealed that the post-fire seedling population is polymorphic, diverse, and reflects the pre-fire random mating system. In contrast to our prediction, we found no division of the post-fire seedling population to distinct sub-populations. Furthermore, as a result of post-fire seed dispersal to longer range than the average pre-fire inter-tree distance, seedlings found under individual burned trees were not necessarily their sole offspring. Although the population as a whole showed a Hardy-Weinberg equilibrium, significant excess of heterozygotes was found within each tallest seedlings group growing under single, large, burned pine trees. Our finding indicates the possible existence of intense natural selection for the most vigorous heterozygous genotypes that are best adapted to the special post-fire regeneration niche, which is the thick ash bed under large, dead, pine trees.

Relationships between tree size, crown shape, gender segregation and sex allocation in Pinus halepensis, a Mediterranean pine tree.

BACKGROUND AND AIMS: Sex allocation has been studied mainly in small herbaceous plants but much less in monoecious wind-pollinated trees. The aim of this study was to explore changes in gender segregation and sex allocation by Pinus halepensis, a Mediterranean lowland pine tree, within tree crowns and between trees differing in their size or crown shape. METHODS: The production of new male and female cones and sex allocation of biomass, nitrogen and phosphorus were studied. The relationship between branch location, its reproductive status and proxies of branch vigour was also studied. KEY RESULTS: Small trees produced only female cones, but, as trees grew, they produced both male and female cones. Female cones were produced mainly in the upper part of the crown, and male cones in its middle and lower parts. Lateral branch density was correlated with the number of male but not female cones; lateral branches were more dense in large than in small trees and even denser in hemispherical trees. Apical branches grew faster, were thicker and their phosphorus concentration was higher than in lateral shoots. Nitrogen concentration was higher in cone-bearing apical branches than in apical vegetative branches and in lateral branches with or without cones. Allocation to male relative to female function increased with tree size as predicted by sex allocation theory. CONCLUSIONS: The adaptive values of sex allocation and gender segregation patterns in P. halepensis, in relation to its unique life history, are demonstrated and discussed. Small trees produce only female cones that have a higher probability of being pollinated than the probability of male cones pollinating; the female-first strategy enhances population spread. Hemispherical old trees are loaded with serotinous cones that supply enough seeds for post-fire germination; thus, allocation to males is more beneficial than to females.

A framework for comparing pollinator performance: effectiveness and efficiency.

Measuring pollinator performance has become increasingly important with emerging needs for risk assessment in conservation and sustainable agriculture that require multi-year and multi-site comparisons across studies. However, comparing pollinator performance across studies is difficult because of the diversity of concepts and disparate methods in use. Our review of the literature shows many unresolved ambiguities. Two different assessment concepts predominate: the first estimates stigmatic pollen deposition and the underlying pollinator behaviour parameters, while the second estimates the pollinator's contribution to plant reproductive success, for example in terms of seed set. Both concepts include a number of parameters combined in diverse ways and named under a diversity of synonyms and homonyms. However, these concepts are overlapping because pollen deposition success is the most frequently used proxy for assessing the pollinator's contribution to plant reproductive success. We analyse the diverse concepts and methods in the context of a new proposed conceptual framework with a modular approach based on pollen deposition, visit frequency, and contribution to seed set relative to the plant's maximum female reproductive potential. A system of equations is proposed to optimize the balance between idealised theoretical concepts and practical operational methods. Our framework permits comparisons over a range of floral phenotypes, and spatial and temporal scales, because scaling up is based on the same fundamental unit of analysis, the single visit.

Unripe red fruits may be aposematic.

The unripe fruits of certain species are red. Some of these species disperse their seeds by wind (Nerium oleander, Anabasis articulata), others by adhering to animals with their spines (Emex spinosa) or prickles (Hedysarum spinosissimum). Certainly neither type uses red coloration as advertisement to attract the seed dispersing agents. Fleshy-fruited species (Rhamnus alaternus, Rubus sanguineus and Pistacia sp.), which disperse their seeds via frugivores, change fruit color from green to red while still unripe and then to black or dark blue upon ripening. The red color does not seem to function primarily in dispersal (unless red fruits form advertisement flags when there are already black ripe fruits on the plant) because the red unripe fruits of these species are poisonous, spiny, or unpalatable. The unripe red fruits of Nerium oleander are very poisonous, those of Rhamnus alaternus and Anabasis articulata are moderately poisonous, those of Rubus sanguineus are very sour, those of Pistacia sp. contain unpalatable resin and those of Emex spinosa and Hedysarum spinosissimum are prickly. We propose that these unripe red fruits are aposematic, protecting them from herbivory before seed maturation.

A sheep in wolf's clothing: do carrion and dung odours of flowers not only attract pollinators but also deter herbivores?

Carrion and dung odours of various flowers have traditionally been considered an adaptation for attracting the flies and beetles that pollinate them. While we accept the role of such odours in pollinator attraction, we propose that they may also have another, overlooked, anti-herbivore defensive function. We suggest that such odours may deter mammalian herbivores, especially during the critical period of flowering. Carrion odour is a good predictor for two potential dangers to mammalian herbivores: (1) pathogenic microbes, (2) proximity of carnivores. Similarly, dung odour predicts faeces-contaminated habitats that present high risks of parasitism. These are two new types of repulsive olfactory aposematic mimicry by plants: (1) olfactory feigning of carcass (thanatosis), a well-known behavioural defensive strategy in animals, (2) olfactory mimicry of faeces, which also has a defensive visual parallel in animals.

Does post-fire plant regeneration mode affect the germination response to fire-related cues?

Vegetative resprouting, soil or canopy-stored seed banks, post-fire seed dispersal and germination are the major strategies by which plants regenerate after fires. Post-fire regeneration modes of plants are commonly based on the presence or absence of post-fire recruitment as well as the presence or absence of post-fire resprouting. High temperatures, smoke and ash are characteristics of fire and the post-fire environment. We hypothesized that heat, smoke, ash and pH will have differential effects on seed germination depending on species' post-fire regeneration strategies: serotinous vs. nonserotinous (which may have soil seed banks) and resprouters vs. nonresprouters (which may be obligate seeders). Here we examined the effects of these factors on the germination of 27 common east Australian species. Most serotinous species supported our hypothesis by showing no effect or reduced germination in response to heat. However, contrary to our prediction, all nonserotinous nonresprouting species also showed no effect or reduced germination in response to heat. Smoke, contrary to our hypothesis, had a negative or no effect on all serotinous and nonresprouting species, but no clear directional effect on serotinous and resprouting species. Supporting our hypotheses, ash and high pH showed positive or nonsignificant effects on the germination of all serotinous resprouting species, and a negative or no effect on nonserotinous resprouting species. However, contrary to our prediction, it had a negative or no effect on the serotinous nonresprouting species and no clear effect on nonserotinous nonresprouting species. We also discovered large differences in germination responses between conspecific populations that varied in their degree of resprouting. Although our data confirmed several of our predictions, the overall conclusion is that the responses of seeds to heat, smoke, ash and pH are not tightly associated with post-fire regeneration functional types.

Rheum palaestinum (desert rhubarb), a self-irrigating desert plant.

The rare plant Rheum palaestinum (Polygonaceae) is a perennial hemicryptophyte that grows during the rainy winter in desert mountainous areas in Israel and Jordan that receive an average annual rainfall of ca. 75 mm. It produces between one and four large round leaves that are tightly attached to the ground and form large rosettes of up to 1 m(2). These leaves differ markedly from the typical small leaves of most desert plants. Moreover, they have a unique 3D morphology resembling a scaled-down mountainous area with well-developed steep drainage systems, raising the question which selective agents were involved in their evolution. We propose that the large leaves collect rainwater that then infiltrates the soil surrounding the root. We measured the seasonal course of leaf growth, examined the area of wet soil surrounding the root after actual and simulated rain, and modeled the water harvesting capacity using the plant leaf area and the weekly precipitation. We show that even in the slightest rains, water flows above the veins to the leaf's base where it irrigates the vertical root. A typical plant harvests more than 4,100 cm(3) of water per year, and enjoys a water regime of about 427 mm/year, equivalent to the water supply in a Mediterranean climate. This is the first example of self-irrigation by large leaves in a desert plant, creating a leaf-made mini oasis.

Comment on "Early domesticated fig in the Jordan Valley".

Kislev et al. (Reports, 2 June 2006, p. 1372) described Neolithic parthenocarpic fig fruits and proposed that they derive from trees propagated only by cuttings and thus represent the first domesticated plant of the Neolithic Revolution. Because parthenocarpic fig trees naturally produce both seeded and seedless fruits and are capable of spontaneous reproduction, we argue that the finds do not necessarily indicate cultivation, nor horticulture predating grain crops.

The effects of nectar-nicotine on colony fitness of caged honeybees.

Nectar of many bee flowers contains secondary compounds, which are considered toxic for honeybees on repeated exposure. Although many anecdotal reports indicate the toxicity of secondary compounds to bees, only a few studies have tested the extent of toxicity at different honeybee ages, especially at the larval stages. Honeybees encounter nicotine at trace concentrations (between 0.1 and 5 ppm) in floral nectar of a few Nicotiana spp. and in Tilia cordata. Adult honeybee workers tolerate these nicotine concentrations. In controlled nonchoice feeding experiments with caged bees, we investigated the effect of nicotine on hatching success and larval and forager survival. Naturally occurring concentrations of nectar-nicotine did not affect hatching success of larvae or their survival, but the latter was negatively affected by higher concentrations of nicotine (50 ppm). Concentrations of nicotine in fresh honey samples from the hives were 90% lower than the concentrations in the offered experimental sucrose solutions. Our results indicate that honeybees can cope with naturally occurring concentrations of nicotine, without notable mortality, even when consumed in large quantities for more than 3 weeks.

Morning floral heat as a reward to the pollinators of the Oncocyclus irises.

Relationships between flowering plants and their pollinators are usually affected by the amount of reward, mainly pollen or nectar, offered to pollinators by flowers, with these amounts usually positively correlated with floral display. The large Oncocyclus iris flowers, despite being the largest flowers in the East Mediterranean flora, are nectarless and have hidden pollen. No pollinators visit the flowers during daytime, and these flowers are pollinated only by night-sheltering solitary male bees. These iris flowers are partially or fully dark-colored, suggesting that they gather heat by absorbing solar radiation. Here we test the hypothesis that the dark-colored flowers of the Oncocyclus irises offer heat reward to their male solitary bee pollinators. Floral temperature was higher by 2.5 degrees C than ambient air after sunrise. Solitary male bees emerged earlier after sheltering in Oncocyclus flowers than from other experimental shelter types. Pollination tunnels facing east towards the rising sun hosted more male bees than other aspects. We suggest that floral heat reward can explain the evolution of dark floral colors in Oncocyclus irises, mediated by the pollinators' behavior.

Limits to reproductive success of Sarracenia purpurea (Sarraceniaceae).

Plant biologists have an enduring interest in assessing components of plant fitness and determining limits to seed set. Consequently, the relative contributions of resource and pollinator availability have been documented for a large number of plant species. We experimentally examined the roles of resource and pollen availability on seed set by the northern pitcher plant Sarracenia purpurea. We were able to distinguish the relative contributions of carbon (photosynthate) and mineral nutrients (nitrogen) to reproductive success. We also determined potential pollinators of this species. The bees Bombus affinis and Augochlorella aurata and the fly Fletcherimyia fletcheri were the only floral visitors to S. purpurea that collected pollen. Supplemental pollination increased seed set by <10%, a much lower percentage than would be expected, given data from noncarnivorous, animal-pollinated taxa. Seed set was reduced by 14% in plants that could not capture prey and by another 23% in plants whose pitcher-shaped leaves were also prevented from photosynthesizing. We conclude that resources are more important than pollen availability in determining seed set by this pitcher plant and that reproductive output may be another "cost" of the carnivorous habit.

Plant coloration undermines herbivorous insect camouflage.

The main point of our hypothesis "coloration undermines camouflage" is that many color patterns in plants undermine the camouflage of invertebrate herbivores, especially insects, thus exposing them to predation and causing them to avoid plant organs with unsuitable coloration, to the benefit of the plants. This is a common case of "the enemy of my enemy is my friend" and a visual parallel of the chemical signals that plants emit to call wasps when attacked by caterpillars. Moreover, this is also a common natural version of the well-known case of industrial melanism, which illustrates the great importance of plant-based camouflage for herbivorous insects and can serve as an independent test for our hypothesis. We claim that the enormous variations in coloration of leaves, petioles and stems as well as of flowers and fruits undermine the camouflage of invertebrate herbivores, especially insects. We assume that the same principle might operate in certain animal-parasite interactions. Our hypothesis, however, does not contrast or exclude other previous or future explanations of specific types of plant coloration. Traits such as coloration that have more than one type of benefit may be selected for by several agents and evolve more rapidly than ones with a single type of advantage.