Reproductive and pollination biology of the Critically Endangered endemic Campanula vardariana in Western Anatolia (Turkey)

Background and aims – Campanula vardariana (Campanulaceae) is a critically endangered endemic chasmophyte with a single population situated in the west of Turkey. Very little is known about the reproductive biology of C. vardariana and more information is needed to develop a sound conservation strategy for this endemic species.Material and methods – Floral traits such as flower morphology, nectar, and sugar concentration, as well as pollen viability and stigma receptivity were measured in different floral phases. We observed insect visitations to the flowers and identified pollinators. Additionally, we investigated the effect of cross and self-pollination on fruit and seed production.Key results – The flowers of C. vardariana are protandrous. The length of the styles, which were 8.74 mm during the pollen loading phase, reached 11.35 mm during the pollen presentation phase. The visitor observations made on the C. vardariana flowers revealed 11 visitor species from 5 families: 5 Halictidae, 3 Apidae, and one species each from Megachilidae, Colletidae, and Bombyliidae. Lasioglossum spp. touched the anthers and stigma using several parts of their bodies and were significant pollinators of C. vardariana. Under natural conditions, the mean number of seeds per fruit was around 60 after cross pollination, while no fruits were formed when pollinators were excluded.Conclusion – Campanula vardariana is entirely dependent on pollinators for its reproductive success, and bees, especially Halictidae and to a lesser extent Apidae, play an important role. Campanula vardariana is restricted to cracks in calcareous rocks and its population is threatened by goat overgrazing and mining activities (quarry formation). Since seed production is abundant in this population, anthropogenic activities currently form the biggest threat to its existence.

Darwin's vexing contrivance: a new hypothesis for why some flowers have two kinds of anther

Heteranthery, the presence of two or more anther types in the same flower, is taxonomically widespread among bee-pollinated angiosperms, yet has puzzled botanists since Darwin. We test two competing hypotheses for its evolution: the long-standing ‘division of labour' hypothesis, which posits that some anthers are specialized as food rewards for bees whereas others are specialized for surreptitious pollination, and our new hypothesis that heteranthery is a way to gradually release pollen that maximizes pollen delivery. We examine the evolution of heteranthery and associated traits across the genus Clarkia (Onagraceae) and study plant–pollinator interactions in two heterantherous Clarkia species. Across species, heteranthery is associated with bee pollination, delayed dehiscence and colour crypsis of one anther whorl, and movement of that anther whorl upon dehiscence. Our mechanistic studies in heterantherous species show that bees notice, forage on and export pollen from each anther whorl when it is dehiscing, and that heteranthery promotes pollen export. We find no support for division of labour, but multifarious evidence that heteranthery is a mechanism for gradual pollen presentation that probably evolved through indirect male–male competition for siring success.

Pollen biodiversity – why are pollen grains different despite having the same function? A review

The main morphological, cytological and physiological characters of ripe pollen are described, compared, analysed and discussed individually, in multiple combinations and in respect to the female counterpart and the biotic and abiotic components of the environment. This is to try to understand the reasons why pollen grains have the same reproductive function, but at dispersal are morphologically and physiologically different in many respects. The considered characters are: one or more types of grain per species; shape and size; number of cells; types of pollen dispersal unit; sporoderm stratification, furrows, colpori and other kinds of apertures; pollen presentation and array; water content percentage; and mature pollen reserves and osmotics. Some of the pollen features are correlated between themselves, some with the female counterpart or male and female competition, and others with the different components of the environment where the species lives, when it flowers and when pollen presentation occurs.

Solitary bee pollinators adjust pollen foraging to the unpredictable flower opening of a species of Petunia (Solanaceae)

Details of the foraging patterns of solitary bees are much less well known than those of social species, and these patterns are often adjusted to exploit floral resources of one or only a few species. The specialized flower-visiting bees of Petunia are good models for investigating such foraging patterns. Here we analysed the floral biology and pollen presentation schedule of the endangered Petunia mantiqueirensis in mixed Araucaria forests of Serra da Mantiqueira, Brazil. Pollinators and their pollen foraging behaviour and food specialization were determined through analyses of scopa pollen loads. Flowers opened throughout the day and presented all their pollen resources within the first 30 min of anthesis, thus providing their pollen resources in an asynchronous fashion in one-flower packages throughout the day. Females of Pseudagapostemon fluminensis were the most frequent flower visitors, contacting stigmas in 96% of their visits, and were the unique effective pollinators of Petunia mantiqueirensis. These pollinators were responsible for the first three visits to 115 individually monitored flowers at any daylight hour, removing ~86% of a flower’s total pollen supply during the first visit. Although female bees harvest the majority of pollen resources of Petunia mantiqueirensis, analyses of scopa loads revealed that most of them also collect pollen from plants of other families while foraging for pollen in Petunia flowers.

The cost of fidelity: foraging oligolectic bees gather huge amounts of pollen in a highly specialized cactus–pollinator association

Plant–pollinator interactions vary along a specialization–generalization continuum. Advances in understanding the evolutionary and ecological consequences of different degrees of specialization depend on precise data on plant–pollinator interdependency. We studied the association of Parodia neohorstii (Cactaceae) and its bee pollinators focusing on pollinator foraging behaviour, flower functioning, female and male reproductive success, and pollen fate. Parodia neohorstii showed synchronized flower opening and pollen presentation but discontinuous blooming. The apparently generalized flowers partition pollen through thigmonastic stamen movements that function as a mechanical filter against generalist bees by restricting access to the major pollen reservoir to bees that show flower handling ‘know-how’, thereby favouring the oligolectic bee Arhysosage cactorum. This pollinator adjusted its pollen foraging to flower opening, removed pollen hurriedly, and promoted maximal fruit and seed set, which was minimal in its absence. Estimates of pollen fate revealed that a huge amount of pollen flows to specialized pollinators (86.5%), and only 0.9% reaches conspecific stigmas. The specialized interaction between P. neohorstii and Arhysosage cactorum, both threatened species, is efficient but fragile. Any environmental modification that causes a mismatch between the partners is likely to result in reproductive failure.

Elaborated pollen packaging and dispensing mechanism induced by petal architecture from a Papaveraceae species

Secondary pollen presentation (SPP) is a reproductive strategy that enhances the efficiency of pollen transfer, which has been explored for more than 200 years, resulting in 10 identified types of SPP. The ephemeral plant Hypecoum erectum L. (Papaveraceae) has an elaborate petal structure. The middle lobe is a key functional organ in SPP. To explore the importance of the middle lobe structure, we measured the flowering process, the curling movement and growth of the middle lobe, pollination characteristics, pollination efficiency, and the mating system in H. erectum in the field. The yellow middle lobe structure had an important role in attracting pollinators. The middle lobes on the inner petals function as a redundant cucullate structure and wrapped about 84% of the total pollen grains as soon as the anthers dehisced. These then grew upward and gradually presented pollen to pollinators via the roll out of the middle lobes. One bee species, Colletes vestitus from Colletidae, was the only effective pollinator of H. erectum. The SPP mechanism increased the efficiency of pollen transfer by C. vestitus. The middle lobes, which wrapped pollen and grew upward, contacted the stigma and provided an advantage for self-pollination and outcrossing by growing upward higher than the corolla. Hypecoum erectum L. has a mixed mating system with selfing and outcrossing. Thus, the SPP mechanism plays a key role during the pollination process and is necessary for improving pollination efficiency and promoting reproductive success.