Replicated host-race formation in bogus yucca moths: genetic and ecological divergence of Prodoxus quinquepunctellus on yucca hosts

Goal: Assess host-race formation in certain moths by examining their genetic and ecological differentiation. Organisms: Stalk-feeding moths, Prodoxus quinquepunctellus, collected from sympatric populations of Yucca elata and Y rostrata in west Texas, USA. Results: Moths on the two yuccas differed significantly in mtDNA haplotype frequencies, emergence time, wing dot number, body size, and ovipositor size and shape. Conclusion: Host-race formation has probably occurred in this yucca moth although genetic divergence was low

Asymmetrical nature of the Trollius-Chiastocheta interaction: insights into the evolution of nursery pollination systems

The mutualistic versus antagonistic nature of an interaction is defined by costs and benefits of each partner, which may vary depending on the environment. Contrasting with this dynamic view, several pollination interactions are considered as strictly obligate and mutualistic. Here, we focus on the interaction between Trollius europaeus and Chiastocheta flies, considered as a specialized and obligate nursery pollination system - the flies are thought to be exclusive pollinators of the plant and their larvae develop only in T.europaeus fruits. In this system, features such as the globelike flower shape are claimed to have evolved in a coevolutionary context. We examine the specificity of this pollination system and measure traits related to offspring fitness in isolated T.europaeus populations, in some of which Chiastocheta flies have gone extinct. We hypothesize that if this interaction is specific and obligate, the plant should experience dramatic drop in its relative fitness in the absence of Chiastocheta. Contrasting with this hypothesis, T.europaeus populations without flies demonstrate a similar relative fitness to those with the flies present, contradicting the putative obligatory nature of this pollination system. It also agrees with our observation that many other insects also visit and carry pollen among T.europaeus flowers. We propose that the interaction could have evolved through maximization of by-product benefits of the Chiastocheta visits, through the male flower function, and selection on floral traits by the most effective pollinator. We argue this mechanism is also central in the evolution of other nursery pollination systems

Comparative Phylogeography of a Coevolved Community: Concerted Population Expansions in Joshua Trees and Four Yucca Moths

Comparative phylogeographic studies have had mixed success in identifying common phylogeographic patterns among co-distributed organisms. Whereas some have found broadly similar patterns across a diverse array of taxa, others have found that the histories of different species are more idiosyncratic than congruent. The variation in the results of comparative phylogeographic studies could indicate that the extent to which sympatrically-distributed organisms share common biogeographic histories varies depending on the strength and specificity of ecological interactions between them. To test this hypothesis, we examined demographic and phylogeographic patterns in a highly specialized, coevolved community – Joshua trees (Yucca brevifolia) and their associated yucca moths. This tightly-integrated, mutually interdependent community is known to have experienced significant range changes at the end of the last glacial period, so there is a strong a priori expectation that these organisms will show common signatures of demographic and distributional changes over time. Using a database of >5000 GPS records for Joshua trees, and multi-locus DNA sequence data from the Joshua tree and four species of yucca moth, we combined paleaodistribution modeling with coalescent-based analyses of demographic and phylgeographic history. We extensively evaluated the power of our methods to infer past population size and distributional changes by evaluating the effect of different inference procedures on our results, comparing our palaeodistribution models to Pleistocene-aged packrat midden records, and simulating DNA sequence data under a variety of alternative demographic histories. Together the results indicate that these organisms have shared a common history of population expansion, and that these expansions were broadly coincident in time. However, contrary to our expectations, none of our analyses indicated significant range or population size reductions at the end of the last glacial period, and the inferred demographic changes substantially predate Holocene climate changes

Floral temperature and optimal foraging: is heat a feasible floral reward for pollinators?

As well as nutritional rewards, some plants also reward ectothermic pollinators with warmth. Bumble bees have some control over their temperature, but have been shown to forage at warmer flowers when given a choice, suggesting that there is some advantage to them of foraging at warm flowers (such as reducing the energy required to raise their body to flight temperature before leaving the flower). We describe a model that considers how a heat reward affects the foraging behaviour in a thermogenic central-place forager (such as a bumble bee). We show that although the pollinator should spend a longer time on individual flowers if they are warm, the increase in total visit time is likely to be small. The pollinator's net rate of energy gain will be increased by landing on warmer flowers. Therefore, if a plant provides a heat reward, it could reduce the amount of nectar it produces, whilst still providing its pollinator with the same net rate of gain. We suggest how heat rewards may link with plant life history strategies

Gelechiidae Moths Are Capable of Chemically Dissolving the Pollen of Their Host Plants: First Documented Sporopollenin Breakdown by an Animal

Background: Many insects feed on pollen surface lipids and contents accessible through the germination pores. Pollen walls, however, are not broken down because they consist of sporopollenin and are highly resistant to physical and enzymatic damage. Here we report that certain Microlepidoptera chemically dissolve pollen grains with exudates from their mouthparts. Methodology/Principal Findings: Field observations and experiments in tropical China revealed that two species of Deltophora (Gelechioidea) are the exclusive pollinators of two species of Phyllanthus (Phyllanthaceae) on which their larvae develop and from which the adults take pollen and nectar. DNA sequences placed the moths and plants phylogenetically and confirmed that larvae were those of the pollinating moths; molecular clock dating suggests that the moth clade is younger than the plant clade. Captive moths with pollen on their mouthparts after 2-3 days of starvation no longer carried intact grains, and SEM photographs showed exine fragments on their proboscises. GC-MS revealed cis-b-ocimene as the dominant volatile in leaves and flowers, but GC-MS analyses of proboscis extracts failed to reveal an obvious sporopollenindissolving compound. A candidate is ethanolamine, which occurs in insect hemolymphs and is used to dissolve sporopollenin by palynologists. Conclusions/Significance: This is the first report of any insect and indeed any animal chemically dissolving pollen

Ability to gall: The ultimate basis of host specificity in fig wasps?

1. Fig trees (Ficus spp.) and their host-specific pollinator fig wasps (Agaonidae) are partners in an obligate mutualism. Receptive phase figs release specific volatiles to attract their pollinators, and this is generally effective in preventing pollinator species from entering figs of the wrong hosts. 2. If entry is attempted into atypical host figs, then ostiole size and shape and style length may also prevent reproduction. In spite of these barriers, there is increasing evidence that fig wasps enter atypical hosts, and that this can result in hybrid seed and fig wasp offspring. 3. This study examines the basis of pollinator specificity in two dioecious fig species from different geographical areas. Kradibia tentacularis pollinates Ficus montana in Asia. Ficus asperifolia from East Africa is closely related but is pollinated by a different species of Kradibia. 4. In glasshouses, K. tentacularis was attracted to its normal host, F1s and backcrosses, but only rarely entered figs of F. asperifolia. Foundresses were able to lay eggs in hybrids, backcrosses, and F. asperifolia, although flower occupancy was lowest in F. asperifolia figs and intermediate in hybrids. 5. The fig wasp failed to reproduce in female F. montana, male F. asperifolia, and male F1s, and most but not all backcrosses to F. montana. This was a result of the failure to initiate gall production. 6. Host specificity in this fig wasp is strongly influenced by host volatiles, but the ability to gall may be the ultimate determinant of whether it can reproduce

Diversity of floral visitors to sympatric Lithophragma species differing in floral morphology

Most coevolving relationships between pairs of species are embedded in a broader multispecific interaction network. The mutualistic interaction between Lithophragma parviflorum (Saxifragaceae) and its pollinating floral parasite Greya politella (Lepidoptera, Prodoxidae) occurs in some communities as a pairwise set apart from most other interactions in those communities. In other communities, however, this pair of species occurs with congeners and with other floral visitors to Lithophragma. We analyzed local and geographic differences in the network formed by interactions between Lithophragma plants and Greya moths in communities containing two Lithophragma species, two Greya species, and floral visitors other than Greya that visit Lithophragma flowers. Our goal was to evaluate if non-Greya visitors were common, if visitor assembly differs between Lithophragma species and populations and if these visitors act as effective pollinators. Sympatric populations of L. heterophyllum and L. parviflorum differ in floral traits that may affect assemblies of floral visitors. Visitation rates by non-Greya floral visitors were low, and the asymptotic number of visitor species was less than 20 species in all populations. Lithophragma species shared some of the visitors, with visitor assemblages differing between sites more for L. heterophyllum than for L. parviflorum. Pollination efficacy experiments showed that most visitors were poor pollinators. Single visits to flowers by this assemblage of species resulted in significantly higher seed set in Lithophragma heterophyllum (30.6 ± 3.9 SE) than in L. parviflorum (4.7 ± 3.4 SE). This difference was consistent between sites, suggesting that these visitors provide a better fit to the floral morphology of L. heterophyllum. Overall, none of the non-Greya visitors appears to be either sufficiently common or efficient as a pollinator to impose strong selection on any of these four Lithophragma populations in comparison with Greya, which occurs within almost all populations of these species throughout their geographic ranges