A Continuum of Conspicuousness, Floral Signals, and Pollination Systems in Rhynchospora (Cyperaceae): Evidence of Ambophily and Entomophily in a Mostly Anemophilous Family

Floral colors and odors are evolutionary strategies used by plants to attract pollinating animals and may be absent in mostly anemophilous groups, such as Cyperaceae. However, considering that insects are floral visitors of some Rhynchospora Vahl species, the objective of this study was to characterize the floral traits and pollination systems within this genus. We analyzed 16 Rhynchospora species with regard to flower morphology, colors of floral structures, floral scents, pollen vectors, and pollination systems. We verified factors that can favor abiotic or biotic pollination in a continuum of floral traits in Rhynchospora. The flower morphology of R. dissitispicula T. Koyama, with inconspicuous brown spikelets in open panicles, is interpreted as a complete adaptation to anemophily. Conspicuous floral traits in Rhynchospora were distinguished from the background by bees. Some species also emit floral volatiles, and we made the first record of floral scent chemistry within the genus. Most of the compounds emitted by these species are known as attractants to many floral-visiting insects. Bees, beetles, and flies visited species with conspicuous floral traits and contributed to fruit set. The investigated floral traits form a continuum across the different pollination systems in Rhynchospora, from anemophilous to ambophilous and then to entomophilous representatives.

Coleopteran Pollinators of Annonaceae in the Brazilian Cerrado—A Review

The evolutionary trajectories of insects and angiosperms appear to be intimately interconnected. Increases in the diversity of phytophagous beetles and angiosperms co-occur in the Mesozoic fossil record, and there is fossil evidence of pollinivory and pollination by insects, both in flowering plants and in gymnosperms. The oldest records of angiosperm pollination indicate flies as pollen vectors. A basal group of angiosperms, the order Magnoliales, has retained plesiomorphic characters such as dozens of pistils and stamens spiraling around the receptacle. In a family of this order, Annonaceae, over 90% of species are pollinated by beetles. In many Annonaceae species, flowers display wide spaces, referred to as floral chambers, where beetles can find shelter from weather conditions and predators, food in the form of pollen and tissues, and a mating site. Two basic types of floral chambers can be distinguished: small chambers visited by small beetles (Nitidulidae, Staphylinidae, Chrysomelidae, and Curculionidae) with diurnal and/or nocturnal activity and large and thermogenic floral chambers visited by beetles of the tribe Cyclocephalini (Scarabaeoidea, Melolonthidae). In the latter case, the heat that the flowers produce may serve as a resource for the beetles that visit them, resulting in smaller endothermy costs for the scarabs. This study reviewed the literature including PhD and MSc theses on cantharophilous Annonaceae in the Cerrado. In this biome, both types of associations are found, although cantharophilous Annonaceae represent a small portion of the plant species (<5%). Cantharophilous Annonaceae in the Cerrado share attributes according to the beetles that pollinate them: species pollinated by small beetles, for instance, may flower throughout the year, whereas Annonaceae pollinated by Cyclocephalini normally flower in the beginning of the rainy season (October/November), in synchrony with the phenological patterns of their pollinators. Cantharophilous Annonaceae flowers, regardless of their size, tend to have light colors and sweet and fruity odors. In addition to the lack of studies on the attraction of beetles by these floral characters, the taxonomic composition of the beetles that pollinate Annonaceae in the Cerrado is poorly known. This review attempts to discuss, in light of what has already been published, potential fields of investigation concerning pollinating beetles’ behavior and evolution.

Disentangling the ecological basis of floral trait variation in Neotropical Piper species

In generalist pollination systems, it has been assumed that pollinators play a minor role in the diversification of floral traits, but recent studies have pointed to a different scenario. Although pollination in Piper is considered generalist, there are flower and inflorescence variations among subclades that may be associated with different pollinator functional groups. Based on this, we aimed to test whether pollinators influenced the evolution of floral traits in a clade of generalist plants, by studying 17 co-occurring Piper spp. Sixteen species were insect-pollinated (46 species: bees, beetles and flies). We found no evidence of anemophily. Eight species were dependent on pollen vectors for sexual reproduction, but no correlation between floral and pollinator traits was recorded. None of the floral traits presented phylogenetic signal, and the evolution of these traits was not correlated. Nine species were independent of pollen vectors for sexual reproduction. We did not find any evidence of pollinator-driven floral diversification of Piper spp.; we suggest a possible role of abiotic factors as agents of selection on floral diversification. As self-pollination seems to be a common feature in the genus, the flexibility of mating systems could be another important factor related to the maintenance of floral phenotypic variation.

Nocturnal pollination: an overlooked ecosystem service vulnerable to environmental change

Existing assessments of the ecosystem service of pollination have been largely restricted to diurnal insects, with a particular focus on generalist foragers such as wild and honey bees. As knowledge of how these plant-pollinator systems function, their relevance to food security and biodiversity, and the fragility of these mutually beneficial interactions increases, attention is diverting to other, less well-studied pollinator groups. One such group are those that forage at night. In this review, we document evidence that nocturnal species are providers of pollination services (including pollination of economically valuable and culturally important crops, as well as wild plants of conservation concern), but highlight how little is known about the scale of such services. We discuss the primary mechanisms involved in night-time communication between plants and insect pollen-vectors, including floral scent, visual cues (and associated specialized visual systems), and thermogenic sensitivity (associated with thermogenic flowers). We highlight that these mechanisms are vulnerable to direct and indirect disruption by a range of anthropogenic drivers of environmental change, including air and soil pollution, artificial light at night, and climate change. Lastly, we highlight a number of directions for future research that will be important if nocturnal pollination services are to be fully understood and ultimately conserved.

Adaptive response of extreme epiphyte Tillandsia species (Bromeliaceae) is demonstrated by different sexual reproduction strategies in the Brazilian Chaco

Some epiphytes are adapted to extreme environments with the ability to survive drought as a result of their morphological (xeromorphism), anatomical (foliar trichomes or scales) and physiological features. In contrast to vegetative features, they may have diverse sexual reproductive strategies. Here we compared the flowering morphology, floral biology, breeding system and pollinators of Tillandsia duratii, T. loliacea and T. recurvifolia (Bromeliaceae) adapted to an extreme environment, the Brazilian Chaco. Tillandsia duratii and T. recurvifolia flower for 5–6 months, whereas T. loliaceae flowers for 11 months, mainly in the dry season, with low to high flowering overlap between them. Although these species generally show similar flowering morphology, they differ in size, colour, odour and/or floral functionality, suggesting non-sharing of pollinators among them. Bimodal pollination occurs in T. duratii (bees, moths other than hawkmoths) and T. recurvifolia (butterflies, hummingbirds); in T. loliacea, we recorded no floral visitors. Tillandsia recurvifolia is self-incompatible, has flowers which approach herkogamy and protandry, and depends on pollen vectors for fruit set. The other two species show reverse herkogamy, autonomous self-pollination and self-compatibility (mixed breeding system); therefore, pollinators are not required. The disparity in reproductive strategies among the three species suggests the possibility of selection for diverse modes of reproduction.

Visual and olfactory floral cues related to ambophilous pollination systems in Poaceae

Usually considered anemophilous, grasses (Poaceae) have been the subject of few studies that recorded insects visiting their flowers. Such visitors, with wind, could act as pollen vectors in a possible ambophilous system. Since colour and scent are floral cues attracting pollinating insects, we aimed to provide the first insights into how these cues could be involved in the attraction of pollinators by grasses. Chloris barbata, Paspalum maritimum (with two morphotypes), P. millegrana and Urochloa plantaginea were studied. We determined the role of each pollen vector (wind and insect) and we registered the frequency of floral visitors. Also, we characterized the colour reflectance of glumes, anthers and stigmas and we determined the colour space in the bee colour hexagon and the chemical composition of floral scents. The floral structures of each species reflect colour differently and are perceived differently by pollinators. Furthermore, the volatile organic compounds detected are similar to those found in entomophilous species. In two of the grass species, we registered eight species of visitors, including bees, wasps and flies. Overall, wind was the most important vector for reproductive success. Insects, however, may induce wind pollination by creating pollen clouds when they visit inflorescences.

Diversity of nectar amino acids in the Fritillaria (Liliaceae) genus: ecological and evolutionary implications

Nectar is considered to be a primary food reward for most pollinators. It mostly contains sugars, but also has amino acids. The significance of the concentration and composition of amino acids in nectar is often less understood than that of its volume, sugar concentration and composition. However, there is a trend towards a broader approach in ecological research, which helps to understand nectar properties in an ecological context. The genus Fritillaria, exhibiting great diversity in flower morphology, nectar composition, and dominant pollinators, allows for the possibility to study some of the above. We studied the concentration and composition of amino acids in the nectar of 38 Fritillaria species attracting different groups of pollen vectors (bees, flies, passerines, and hummingbirds). The flowers of fritillaries produced nectar with a varying composition and concentration of amino acids. These differences were mostly associated with the pollinator type. The nectar of passerine bird-pollinated species was rich in amino acids, whereas humming bird-pollinated produced low amino acid nectar. Contrary to previous reports nectar of the insect-pollinated species did not contain a higher amount of proline. Two non-protein amino acids, sarcosine and norvaline, were detected in the floral nectar for the first time.