The majority of flowering plants, including many rare and threatened species, are pollinated by animals, but little is known of pollination and breeding systems of many endangered species. Polemonium caeruleum (Polemoniaceae) is a red-listed species and is regarded as dichogamous, self-compatible and bee pollinated. However, some studies show that it is visited by a vast assemblage of anthophilous insects from many taxonomic orders and that breeding systems vary greatly between closely related taxa of this genus. Over a period of 3 years we investigated breeding system, dichogamy, nectar secretion and composition, insect visitations and pollen loads in flowers of P. caeruleum in north-eastern Poland to determine whether the reproductive biology of the plant explains its rarity. Contrary to published data, our study plants were self-incompatible and showed a high degree of outcrossing. Our experimental work confirmed the occurrence of protandry in this species, revealed that nectar is sucrose-dominant and proline-rich and, for the first time for Polemoniaceae, that nectar secretion and nectar sugar concentration in flowers of P. caeruleum is female-biased. Although flowers were visited by at least 39 species of insects from five taxonomic orders, overall the plant exhibited many characters associated with bee pollination, and analysis of insect performance showed that bumblebees and honeybees are the key pollinators; occasionally hoverflies and butterflies may also be involved. We conclude that, in terms of pollination system, P. caeruleum demonstrates high apparent generalization, but low realized generalization, and is a functional specialist, as most pollinators belong to a single functional group (guild). Its conservation status, at least in our study population, cannot be explained in terms of the biological properties of its breeding or pollination systems; rather, the present decline of the species is caused by habitat loss. However, if this process and bumblebee decline in Europe continue, P. caeruleum populations may diminish in numbers and density and, owing to the self-incompatibility of the species, quickly become severely pollen-limited, thereby accelerating further local extinctions.
Angelica sylvestris is morphologically well adapted to ecological generalization, and there is little evidence that the surveyed populations represent distinct pollination ecotypes. Most likely, the observed variation in floral characters can be interpreted as 'adaptive wandering'. Specialization in this family seems possible only under very special circumstances, for example when the pollinator community comprises insect visitor groups that clearly differ in their pollination capacity (e.g. due to differences in their functional morphology) and/or have different perceptional biases (e.g. for colour or scent). However, the barrier to the evolution of morphological adaptations resulting in the fine-tuning of the flower towards particular pollinator types may arise from the architectural constraints on the floral bauplan that make umbellifers so uniform in their floral displays and so successful in attracting large numbers of pollinators.
Most flowers are visited by a wide range of potential pollinators. However, their efficiency in pollen removal and deposition, and other behavioural factors affecting pollination effectiveness may greatly differ among taxa, and even individuals. Fritillary (Fritillaria meleagris L., Liliaceae) is a spring-flowering, critically endangered plant in the Polish flora, red-listed in most of the European countries of its range. Based on indirect evidence, that is, body pollen loads, visitation frequency and seasonal abundance, it is estimated that its key pollinators are queen bumblebees, but, as shown in the literature, the largest Fritillaria pollen loads are carried by solitary bees. To study pollinator effectiveness for floral visitors to F. meleagris, we performed a garden experiment, where we analysed pollen deposition and assessed pollen removal per single flowervisit in the plant. Similarly to field conditions reported in the literature, our experimental plants were serviced by nectarseeking bumblebee queens and two taxa of solitary bees, small pollen-collecting Andrena and large, nectar-seeking Anthophora males. When ''quality'' component was addressed, despite the character of visits, insects from all groups deposited more pollen than was found on unvisited flowers, but they did not differ significantly from each other in pollen deposition on virgin stigmas. We also found some differences in pollen removal both within-and among-visitor species and control flowers, unfortunately due to extremely high variation of the results they were all statistically insignificant. However, when ''quantity'' component of insect performance was concerned, we observed that over 81 % of visits were by bumblebees. Bombus queens stayed on flowers significantly less time than small Andrena individuals (13 % of recorded visits) and equally long as Anthophora males (only 6 % of visits). We conclude that although all the visitor groups can pollinate the flowers of F. meleagris, bumblebee queens indeed proved to be the most effective pollinators of the plant, when both quality and quantity components of pollination are concerned.
Fritillaria is a genus consisting of 130 to 140 species of bulbous plants, native to temperate regions of the northern hemisphere. Generally viewed as an insect pollinated genus with the exception of two North American species, Fritillaria gentneri and F. recurva, which are described as hummingbird-pollinated and the Asian species, F. imperialis, described as passerine-pollinated. These pollinator shifts are possibly the result of adaptive changes to the structure and morphology of the nectary, as well as a change in the nectar concentration and composition. A study was conducted in a target group of 56 Fritillaria species, based on the morphology of their nectaries and nectar composition to assess the significance of pollination mode as well as its predisposition for the evolution of bird pollination. All species studied had nectaries located at their tepal base and produced nectar, but their size, shape, color, and composition all varied. Most fritillaries had hexose-rich nectar, in easily accessible and unprotected nectaries. Scanning electron microscope (SEM) analysis revealed that the surface of the nectaries of most Fritillaria species was flat and clearly distinct from that of the surrounding tissues, which might be regarded as an adaptation for insect-pollination. Nectaries of F. imperialis were considerably larger and had dilute nectar without sucrose, which was produced profusely, thereby fulfilling the criteria characteristic of ornithophilous flowers. The copious nectar of presumed hummingbird-pollinated species was rather balanced and of medium sugar concentration. Their large lanceolate nectaries contrasted sharply with the tessellated background of their tepals. These characters might indicate a mixed pollination system that engages both birds and insects. Floral anatomy and microstructure and nectar composition for Fritillaria species in subgenera Korolkowia and Liliorhiza are studied for the first time.
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