Sexual dichromatism and color diversity in the spiny lava lizard Tropidurus spinulosus using lizard visual modelling.

Colors are important vehicles for social signals in many taxa. In Squamata, previous studies have linked color characteristics and chromatic diversity to sexual selection and, particularly, species showing male-biased body size dimorphism also showed male-biased dichromatism and color diversity. Sexual dichromatism may occur in body regions used for conspecific communication and it may be expressed at wavelengths, such as ultraviolet, easily perceivable by conspecifics. We tested this prediction in a social lizard model, Tropidurus spinulosus, using spectrophotometry and visual modelling which enable colors to be interpreted as the individuals of the same taxon see them. Our results indicate that sexual dichromatism occurs in the ventral regions and the flanks, which are the body regions involved in sexual displays. Males show greater color diversity, having larger color volumes and more contrasting colors. These findings reinforce the idea that sexual selection towards males is coupled with the evolution of male-biased, diverse, coloration which could act as a signal in social reproductive contexts.

Breakdown of species boundaries in Mandevilla: floral morphological intermediacy, novel fragrances and asymmetric pollen flow.

Phenotypic intermediacy is an indicator of putative hybrid origin and has provided the main clues to discovering hybrid plants in nature. Mandevilla pentlandiana and M. laxa (Apocynaceae) are sister species with clear differences in floral phenotype and associated pollinator guilds: diurnal Hymenoptera and nocturnal hawkmoths, respectively. The presence of individuals with intermediate phenotypes in a wild population raises questions about the roles of visual and olfactory signals (i.e. corolla morphology and floral fragrances) as barriers to interbreeding, and how the breakdown of floral isolation occurs. We examined phenotypic variation in a mixed Mandevilla population, analysing the chemical composition of floral fragrances, characterising floral shape through geometric morphometrics and assessing individual grouping through taxonomically relevant traits and an unsupervised learning algorithm. We quantified the visitation frequencies of floral visitors and tracked their foraging movements using pollen analogues. The presence of morphologically intermediate individuals and pollen analogue movement suggested extensive hybridisation between M. laxa and M. pentlandiana, along with asymmetrical rates of backcrossing between these putative hybrids and M. laxa. Floral volatiles from putative hybrid individuals showed a transgressive phenotype, with additional compounds not emitted by either parental species. Our results suggest the presence of a hybrid swarm between sympatric M. pentlandiana and M. laxa and indicate that initial hybridisation events between these parental species are rare, but once they occur, visits between putative hybrids and M. laxa are common and facilitate continued introgression.

Drift effects on the multivariate floral phenotype of Calceolaria polyrhiza during a post-glacial expansion in Patagonia.

Quaternary environmental changes substantially impacted the landscape and promoted rapid evolutionary changes in many species; however, analyses of adaptive phenotypic variation in plants have usually neglected the underlying historical context. Here, we associate phylogeography and phenotypic evolution by analysing the divergence of Calceolaria polyrhiza multivariate floral phenotype after a Pleistocene post-glacial expansion in Patagonia. Phenotypic matrix (P) properties (size, shape, orientation and phenotypic integration) of six refugium and six recent populations from two different phylogroups were compared following different approaches. We found that P-matrix shape and orientation remained stable despite the strong phylogeographic footprint of post-glacial expansion. However, average proportional reductions in matrix size supported the expectation that drift had a significant effect on the floral phenotype in the northern phylogroup. When phylogeographic history was not included in the analyses, the results overestimated phenotypic differences, whereas under explicit phylogeographic control, drift appeared as the best explanation for matrix differences. In general, recent populations showed a larger phenotypic divergence among them, but a lower overall phenotypic variation than refugium populations. Random Skewers analyses indicated a lower potential response to selection in recently colonized populations than in refugium populations. We discuss that the combination of phylogeographic analyses with geographical distribution of functional phenotypic (genotypic) variation is critical not only to understand how historical effects influence adaptive evolution, but also to improve field comparisons in evolutionary ecology studies.

Selection on signal­reward correlation: limits and opportunities to the evolution of deceit in Turnera ulmifolia L.

Because pollinators are unable to directly assess the amount of rewards offered by flowers, they rely on the information provided by advertising floral traits. Thus, having a lower intra-individual correlation between signal and reward (signal accuracy) than other plants in the population provides the opportunity to reduce investment in rewards and cheat pollinators. However, pollinators' cognitive capacities can impose a limit to the evolution of this plant cheating strategy if they can punish those plants with low signal accuracy. In this study, we examined the opportunity for cheating in the perennial weed Turnera ulmifolia L. evaluating the selective value of signal accuracy, floral display and reward production in a natural population. We found that plant reproductive success was positively related to signal accuracy and floral display, but not to nectar production. The intensity of selection on floral display was more than three times higher than on signal accuracy. The pattern of selection indicated that pollinators can select for signal accuracy provided by plants and suggests that learning abilities of pollinators can limit the evolution of deceptive strategies in T. ulmifolia.

A simple floral fragrance and unusual osmophore structure in Cyclopogon elatus (Orchidaceae).

We studied gland morphology, anatomy and the chemical composition of the floral fragrance in the sweat bee-pollinated orchid Cyclopogon elatus. This is apparently the first such analysis for any Cyclopogon species, and one of very few studies in which both odour and osmophore are characterised in a nectar-rewarding orchid. Structures responsible for floral scent production were localised with neutral red staining and histochemical assays for lipids and starch. Their morphology and anatomy were studied with scanning electron microscopy and light microscopy thin sections, respectively. Fragrance samples were collected using SPME fibres and analysed with GC-MS. Anatomical evidence suggests that two parallel oval-shaped patches of unicellular trichomes on the abaxial surface of the labellum are osmophores. These are rich in stored lipids, while the parenchyma surrounding the vascular bundles contains starch. Only freshly opened flowers produced odours, while buds and withered flowers lacked scent. The chemical composition of the odour was dominated (>99.8%) by a single compound, trans-4,8-dimethyl-nona-1,3,7-triene (DMNT). Gland anatomy and position on the outside of the perianth are unusual for scent glands in general. The presence of DMNT, a nearly ubiquitous compound in herbivore-induced vegetative emissions and one of the major floral volatiles of Yucca, is not surprising in view of hypotheses on the evolutionary origin of flower scents, suggesting that wound volatiles are utilised as kairomonal attractants by florivores whose activities result in pollination.

Variable selection patterns on the labellum shape of Geoblasta pennicillata, a sexually deceptive orchid.

By mimicking shape and female mating pheromones, flowers of sexually deceptive orchids attract sexually excited males which pollinate them while trying to copulate. Although many studies have demonstrated the crucial importance of odour signals in these systems, most flowers pollinated by pseudocopulation resemble, at least superficially, an insect body and these visual cues may be important to cheat pollinators. In this 2-year study, we show that the shape of the labellum of Geoblasta pennicillata is a target of pollinator-mediated natural selection. Contrary to our expectations, plants with a labellum shape more similar to female wasps were not favoured. The strength and pattern of phenotypic selection varied between study years and sexual functions. Although selection through female success was probably associated to the fine-tuning of the mechanical fit between flower form and male wasp, shape was the target of natural selection through male success in both study years indicating that male wasps use this trait when choosing flowers. The imperfect mimicry and patterns of selection observed indicated that an exact imitation is not needed to attract and deceive the pollinators and they suggested a receiver perceptual bias towards uncommon phenotypes.