Mani Shrestha

Flower colour and phylogeny along an altitudinal gradient in the Himalayas of Nepal

Both the phylogenetic structure and trait composition of flowering plant communities may be expected to change with altitude. In particular, floral colours are thought to vary with altitude because Hymenoptera typically decline in importance as pollinators while Diptera and Lepidoptera become more important at higher elevations. Thus, ecological filtering among elevation zones and competitive processes among co-occurring species within zones could influence the floral chromatic cues present at low and high elevations. We collected data from 107 species of native flowering plants in the Himalaya Mountains of central Nepal over an elevation range of 900-4100 m, which includes habitat ranging from subtropical to subalpine within a relatively small geographical area. Using a phylogenetic framework, we asked whether and how flower colour diversity differed between species assemblages at lower and higher elevation, between monocots and eudicots, and between our sample from central Nepal and angiosperms from other regions of the world. There was significant phylogenetic clustering in the communities as a result of monocots, particularly orchids, which were found overwhelmingly at lower elevations. Phylogenetic signal for floral colours indicated that related species had colours that were more disparate than expected under Brownian motion evolution. Floral colours were significantly more diverse in the higher elevation subalpine zone than in the subtropical zone. However, the chromatic cues at both elevations were consistent with the hue discrimination abilities of the trichromatic hymenopteran visual system. Synthesis. Flower colour is not highly differentiated between subtropical and subalpine vegetation due to differences in the available orders of insect pollinators, or by the rate or direction of colour evolution in the lineages composing the two communities.

Evaluating the spectral discrimination capabilities of different pollinators and their effect on the evolution of flower colors

Important plant pollinators like bees and birds have very different color visual systems. Previous work has attempted to relate flower syndromes to the respective visual capabilities of the most important pollinators, but has often been limited by the lack of robust means to make between-species comparisons of how flower color signals are processed. In a recent study we solved this dilemma by comparing the raw spectral signals, quantifiable by major inflection points on a wavelength scale, from different flowers whose pollinators were known from direct observation. Here we elaborate on how this method allows robust cross species comparisons that are independent of the requirement to know the complex and often inaccessible physiological data about color processing in different animals. The use of this method should thus allow for the testing of pollinator syndrome hypotheses for different animal pollinators from different regions of the world.

Flower Colours through the Lens: Quantitative Measurement with Visible and Ultraviolet Digital Photography

Background The study of the signal-receiver relationship between flowering plants and pollinators requires a capacity to accurately map both the spectral and spatial components of a signal in relation to the perceptual abilities of potential pollinators. Spectrophotometers can typically recover high resolution spectral data, but the spatial component is difficult to record simultaneously. A technique allowing for an accurate measurement of the spatial component in addition to the spectral factor of the signal is highly desirable. Methodology/Principal findings Consumer-level digital cameras potentially provide access to both colour and spatial information, but they are constrained by their non-linear response. We present a robust methodology for recovering linear values from two different camera models: one sensitive to ultraviolet (UV) radiation and another to visible wavelengths. We test responses by imaging eight different plant species varying in shape, size and in the amount of energy reflected across the UV and visible regions of the spectrum, and compare the recovery of spectral data to spectrophotometer measurements. There is often a good agreement of spectral data, although when the pattern on a flower surface is complex a spectrophotometer may underestimate the variability of the signal as would be viewed by an animal visual system. Conclusion Digital imaging presents a significant new opportunity to reliably map flower colours to understand the complexity of these signals as perceived by potential pollinators. Compared to spectrophotometer measurements, digital images can better represent the spatio-chromatic signal variability that would likely be perceived by the visual system of an animal, and should expand the possibilities for data collection in complex, natural conditions. However, and in spite of its advantages, the accuracy of the spectral information recovered from camera responses is subject to variations in the uncertainty levels, with larger uncertainties associated with low radiance levels.

Distinctive convergence in Australian floral colours seen through the eyes of Australian birds

We used a colour-space model of avian vision to assess whether a distinctive bird pollination syndrome exists for floral colour among Australian angiosperms. We also used a novel phylogenetically based method to assess whether such a syndrome represents a significant degree of convergent evolution. About half of the 80 species in our sample that attract nectarivorous birds had floral colours in a small, isolated region of colour space characterized by an emphasis on long-wavelength reflection. The distinctiveness of this ‘red arm’ region was much greater when colours were modelled for violet-sensitive (VS) avian vision than for the ultraviolet-sensitive visual system. Honeyeaters (Meliphagidae) are the dominant avian nectarivores in Australia and have VS vision. Ancestral state reconstructions suggest that 31 lineages evolved into the red arm region, whereas simulations indicate that an average of five or six lineages and a maximum of 22 are likely to have entered in the absence of selection. Thus, significant evolutionary convergence on a distinctive floral colour syndrome for bird pollination has occurred in Australia, although only a subset of bird-pollinated taxa belongs to this syndrome. The visual system of honeyeaters has been the apparent driver of this convergence.

Innate colour preferences of the Australian native stingless bee Tetragonula carbonaria Sm.

Innate preferences promote the capacity of pollinators to find flowers. Honeybees and bumblebees have strong preferences for ‘blue’ stimuli, and flowers of this colour typically present higher nectar rewards. Interestingly, flowers from multiple different locations around the world independently have the same distribution in bee colour space. Currently, however, there is a paucity of data on the innate colour preferences of stingless bees that are often implicated as being key pollinators in many parts of the world. In Australia, the endemic stingless bee Tetragonula carbonaria is widely distributed and known to be an efficient pollinator of both native plants and agricultural crops. In controlled laboratory conditions, we tested the innate colour responses of naïve bees using standard broadband reflectance stimuli representative of common flower colours. Colorimetric analyses considering hymenopteran vision and a hexagon colour space revealed a difference between test colonies, and a significant effect of green contrast and an interaction effect of green contrast with spectral purity on bee choices. We also observed colour preferences for stimuli from the blue and blue–green categorical regions of colour space. Our results are discussed in relation to the similar distribution of flower colours observed from bee pollination around the world.

Distribution Patterns of Medicinal Plants along an Elevational Gradient in Central Himalaya, Nepal

This study aimed to compare the distribution patterns and trends of plant parts used among different groups of medicinal plants, geographical regions, and between medicinal plants and all vascular plants. We used the published sources for elevation records of 2,331 medicinal plant species to interpolate presence between minimum and maximum elevations and estimated medicinal plant richness for each 100-m elevational band. Monte Carlo simulations were used to test whether differences in elevational distribution between different groups of medicinal plants were significant. Total number of medicinal plants as well as different groups showed unimodal relationship with elevation. The elevational distributions of medicinal plants significantly differ between regions and between medicinal plant groups. When comparing the richness of all medicinal plants to all vascular plants, Monte Carlo simulations indicated that the numbers of medicinal plants are higher than expected at low elevations. The highest richness of medicinal plants at low elevation could be possibly due to favorable environmental factors such as high temperature, rainfall, sunlight or due to higher density of human population and thus higher pressure on use of any plants in lower elevations.

Floral colours in a world without birds and bees: the plants of Macquarie Island.

We studied biotically pollinated angiosperms on Macquarie Island, a remote site in the Southern Ocean with a predominately or exclusively dipteran pollinator fauna, in an effort to understand how flower colour affects community assembly. We compared a distinctive group of cream-green Macquarie Island flowers to the flora of likely source pools of immigrants and to a continental flora from a high latitude in the northern hemisphere. We used both dipteran and hymenopteran colour models and phylogenetically informed analyses to explore the chromatic component of community assembly. The species with cream-green flowers are very restricted in colour space models of both fly vision and bee vision and represent a distinct group that plays a very minor role in other communities. It is unlikely that such a community could form through random immigration from continental source pools. Our findings suggest that fly pollination has imposed a strong ecological filter on Macquarie Island, favouring floral colours that are rare in continental floras. This is one of the strongest demonstrations that plant-pollinator interactions play an important role in plant community assembly. Future work exploring colour choices by dipteran flower visitors would be valuable.

Why background colour matters to bees and flowers

Flowers are often viewed by bee pollinators against a variety of different backgrounds. On the Australian continent, backgrounds are very diverse and include surface examples of all major geological stages of the Earth’s history, which have been present during the entire evolutionary period of Angiosperms. Flower signals in Australia are also representative of typical worldwide evolutionary spectral adaptations that enable successful pollination. We measured the spectral properties of 581 natural surfaces, including rocks, sand, green leaves, and dry plant materials, sampled from tropical Cairns through to the southern tip of mainland Australia. We modelled in a hexagon colour space, how interactions between background spectra and flower-like colour stimuli affect reliable discrimination and detection in bee pollinators. We calculated the extent to which a given locus would be conflated with the loci of a different flower-colour stimulus using empirically determined colour discrimination regions for bee vision. Our results reveal that whilst colour signals are robust in homogeneous background viewing conditions, there could be significant pressure on plant flowers to evolve saliently-different colours to overcome background spectral noise. We thus show that perceptual noise has a large influence on how colour information can be used in natural conditions.

Out of Africa: evidence of the obligate mutualism between long corolla tubed plant and long-tongued fly in the Himalayas

Summary Mutualism between long corolla tubed plants and their potential pollinators, long‐tongued flies, is a classic example of coevolution, but to date, has only been reported from the regions of southern Africa. Many plant species from the Himalayas also show botanical characteristics that could be consistent with pollination by long‐tongued flies. Here, we seek the evidence of the “long‐tongued‐long tubed fly/flower” mutualism out of Africa, in a different continent and climatic region, the Himalayas. Floral traits of Himalayan region endemic alpine genus, Roscoea, indicate possible mutualism with long‐tongued flies for pollination success; however, effective pollinators of this genus are yet unknown. This study investigates whether long‐tongued flies and Roscoea purpurea in Nepal Himalayas show exclusive mutualism for their survival/reproduction. We made extensive observations of floral visitors of R. purpurea and food source of Philoliche longirostris across their wide ranges of populations in Nepal Himalayas for three consecutive years (2012–2014). To confirm the obligate reliance of R. purpurea upon P. longirostris for pollination success, manipulated pollination experiments were conducted at two populations for 2 years. Similarly foraging behavior, visitation frequency, and pollination efficiency of P. longirostris were assessed at two populations for 2 years, and its contribution for the reproductive success of R. purpurea was evaluated. Our results indicate that R. purpurea is self‐compatible but lacks autonomous selfing and obligatorily relies on P. longirostris for reproductive success. Across all populations, P. longirostris was observed as an exclusive and highly efficient pollinator of R. purpurea, while P. longirostris exclusively depends up on R. purpurea for food source. Out of Africa, this study provides the first evidence of long‐tongued fly pollination system and indicates the possibility of additional instances of such a rare phenomenon in the Himalayas. Finding of specialized pollinator of Roscoea only at its evolutionary center indicates that Roscoea species are originally pollinated by long‐tongued flies. Spatial mismatch with specialized pollinators may have induced the evolution of autonomous selfing in North Indochinese clades of Roscoea. This finding thus substantiates how geographic disjunction causes the shifting of pollination mechanism in closely related plant species.

The colorful language of Australian flowers

The enormous increase in phylogenetic information in recent years has allowed many old questions to be reexamined from a macroevolutionary perspective. We have recently considered evolutionary convergence in floral colors within pollination syndromes, using bird-pollinated species in Australia. We combined quantitative measurements of floral reflectance spectra, models of avian color vision, and a phylogenetic tree of 234 Australian species to show that bird-pollinated flowers as a group do not have colors that are significantly different from the colors of insect-pollinated flowers. However, about half the bird-pollinated flowers have convergently evolved a narrow range of colors with dominant long-wavelength reflection far more often than would be expected by chance. These convergent colors would be seen as distinctly different from other floral colors in our sample when viewed by honeyeaters (family Meliphagidae), birds with a phylogenetically ancestral type of color vision and the dominant avian pollinators in Australia. Our analysis shows how qualitative ideas in natural history, like the concept of pollination syndromes, can be given more precise definition and rigorous statistical testing that takes into account phylogenetic information.