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1.  Functional implications of the staminal lever mechanism in Salvia cyclostegia (Lamiaceae) 
Annals of Botany  2011;107(4):621-628.
Background and Aims
Flower morphology and inflorescence architecture affect pollinator foraging behaviour and thereby influence the process of pollination and the reproductive success of plants. This study explored possible ecological functions of the lever-like stamens and the floral design in Salvia cyclostegia.
Methods
Flower construction was experimentally manipulated by removing either the lower lever arms or the upper fertile thecae of the two stamens from a flower. The two types of manipulated individuals were intermixed with the control ones and randomly distributed in the population.
Key Results
Removing the sterile lower lever arms significantly reduced handling time per flower of the main pollinator, Bombus personatus. Interestingly, this manipulation did not increase the number of flowers probed per plant visit, but instead reduced it, i.e. shortened the visit sequence of the bumble-bees. Both loss of staminal lever function by removing lower lever arms and exclusion of self pollen by removing upper fertile thecae significantly reduced seed set per flower and seed set per plant. Both the manipulations interacted significantly with inflorescence size for the effect on female reproductive output.
Conclusions
Though the intact flowers demand a long handling time for pollinators, the reversible staminal lever is of advantage by promoting dispersal of pollen and thus the male function. The particular floral design in S. cyclostegia contributes to the floral constancy of B. personatus bumble-bees, with the lower lever arms acting as an optical cue for foraging cognition.
doi:10.1093/aob/mcr011
PMCID: PMC3064543  PMID: 21292677
Adaptation; Bombus personatus; experimental flower manipulation; floral constancy; floral design; foraging behaviour; geitonogamy; Salvia
2.  Pollination syndromes in African Marantaceae 
Annals of Botany  2009;104(1):41-56.
Background and Aims
The Marantaceae (550 spp.) is the most derived family in the order Zingiberales and exhibits a complex explosive pollination mechanism. To understand the evolutionary significance of this unique process of pollen transfer, comparative morphological and ecological studies were conducted in Gabon.
Methods
During a total stay of 11 months, 31 species of Marantaceae were investigated at different sites in Gabon. The study included analyses of floral diversity, observations on the pollinator spectrum as well as ecological measurements (e.g. nectar sugar concentration and volume).
Key Results
Analyses reveal five flower types based on flower size and pigmentation, spatial arrangement of the floral tube and presence/absence of nectar guides and conspicuous outer staminodes. Each type is associated with a specific functional pollinator group leading to the description of distinct pollination syndromes. The ‘small (horizontal)’ flowers are predominantly pollinated by small bees (Thrinchostoma spp., Allodapula ornaticeps), the ‘large (horizontal)’ and ‘medium-sized (horizontal)’ flowers by medium-sized bees (Amegilla vivida, Thrinchostoma bicometes), the ‘locked (horizontal)’ flowers by large bees (Xylocopa nigrita, X. varipes) and the ‘(large) vertical’ flowers by sunbirds.
Conclusions
The longevity of Marantaceae individuals and the omnipresence of their pollinators allowed the specialization to a given functional pollinator group. Intermediate ecological values, however, make occasional pollinator overlaps possible, indicating potential pathways of pollinator shifts. Similar radiation tendencies observed on other continents hint at similar selective pressures and evolutionary constraints.
doi:10.1093/aob/mcp106
PMCID: PMC2706729  PMID: 19443460
Africa; floral diversity; functional pollinator groups; Gabon; Marantaceae; nectar; plant–animal interaction; pollination syndromes
4.  Bilabiate Flowers: The Ultimate Response to Bees? 
Annals of Botany  2007;100(2):361-374.
Background and Aims
Bilabiate flowers have evolved in many lineages of the angiosperms, thus representing a convincing example of parallel evolution. Similar to keel blossoms, they have obviously evolved in order to protect pollen against pollen-collecting bees. Although many examples are known, a comprehensive survey on floral diversity and functional constraints of bilabiate flowers is lacking. Here, the concept is widened and described as a general pattern.
Methods
The present paper is a conceptional review including personal observations of the authors. To form a survey on the diversity of bilabiate blossoms, a search was made for examples across the angiosperms and these were combined with personal observations collected during the last 25 years, coupled with knowledge from the literature. New functional terms are introduced that are independent of morphological and taxonomic associations.
Key Results
Bilabiate constructions occur in at least 38 angiosperm families. They are characterized by dorsiventral organization and dorsal pollen transfer. They are most often realised on the level of a single flower, but may also be present in an inflorescence or as part of a so-called ‘walk-around flower’. Interestingly, in functional terms all nototribic blossoms represent bilabiate constructions. The great majority of specialized bee-flowers can thus be included under bilabiate and keel blossoms. The syndrome introduced here, however, also paves the way for the inclusion of larger animals such as birds and bats. The most important evolutionary trends appear to be in the saving of pollen and the precision of its transfer. With special reference to the Lamiales, selected examples of bilabiate flowers are presented and their functional significance is discussed.
Conclusions
Bilabiate blossoms protect their pollen against pollen-collecting bees and at the same time render their pollination more precisely. The huge diversity of realised forms indicate the high selection pressure towards the bilabiate syndrome. As bees are very inventive, however, bilabiate constructions will not represent the ultimate response to bees.
doi:10.1093/aob/mcm123
PMCID: PMC2735325  PMID: 17652341
Bilabiate flowers; nototribic (dorsal) pollination; floral diversity; bee blossoms; functional morphology
5.  Floral Diversity and Pollen Transfer Mechanisms in Bird-pollinated Salvia Species 
Annals of Botany  2007;100(2):401-421.
Background and Aims
Bird-pollinated (ornithophilous) Salvia species (sages) transfer pollen either by means of a staminal lever mechanism or by immovable stamens. As the distribution of the two modes within the genus is not known, we present a survey of all ornithophilous sages. The main focus is given to floral diversity especially with respect to functional lever morphology. Thereby the hypothesis is tested that, due to a pollinator shift from bees to birds, the lever mechanism became unnecessary.
Methods
To get a general idea about the diversity of pollen transfer mechanisms, 186 ornithophilous Salvia species were classified according to the functional morphology of the stamen and the need for a lever movement. To test the functionality of the staminal levers and the fitting between flowers and birds the process of pollen transfer was examined by pollinator observations and tested by inserting museum skins and metal rods into fresh flowers.
Key Results
The diversity of pollen transfer mechanisms is represented by eight case studies illustrating three main groups. In group I (approx. 50 %) the staminal lever mechanism is necessary to open access to nectar and to enable the transfer of pollen that is hidden in the upper lip. In group II (approx. 34 %) pollen is freely accessible and the lever mechanism is reduced in different ways and to different degrees. In group III (approx. 4 %) the lever works as in group I, but pollen is freely accessible as in II. The remaining approx. 13 % are not clearly classified.
Conclusions
It is considered that the driving force behind the diverse modes of reduction is the necessity to increase the distance between nectar and pollen, thereby ensuring pollen deposition on the bird's feathered head. This is achieved several times in parallel by corolla elongation and/or exposure of the pollen-sacs. As soon as pollen is freely accessible, the lever movement loses its significance for pollination.
doi:10.1093/aob/mcm036
PMCID: PMC2735312  PMID: 17522077
6.  New Insights into the Functional Morphology of the Lever Mechanism of Salvia pratensis (Lamiaceae) 
Annals of Botany  2007;100(2):393-400.
Background and Aims
The functional morphology of Salvia pratensis flowers was re-investigated, after new insights revealed that pollen dispensing is one of the main functions of the staminal lever. In particular, no detailed information was available regarding the process of pollen transfer and the forces arising between the pollen-bearing thecae and the pollinating bee's body. The assumption was made that these forces play a significant role in pollen dispensing.
Methods
The functional morphology of S. pratensis flowers and the interaction between flowers and bees (Apis mellifera) were studied by reconstructing stress and strains by using qualitative and semi-quantitative theoretical analysis. Flowers were manipulated to study the spatial arrangement of the filament and lever, and of the head and proboscis of the visiting bee inside the tube. Photographs and films of bee visits on flowers were used to analyse the interaction of pollinator and staminal lever.
Key Results
The spoon-shaped lower lever of S. pratensis has a small hole through which a bee introduces its proboscis into the corolla tube. Although mentioned for the first time by Kerner von Marilaun in 1891, presented here is the first drawing and the first photograph showing this interaction in detail. The analysis of the interaction of flower visitor and the lever mechanism revealed that the position of bees on different flowers is spatially very similar. Flower morphology constrains postures of legitimately nectar-probing bees within narrow bounds. A theoretical discussion on structural elements and force progression in the flower allows the principles of lightweight architecture in flower morphology to be recognized.
Conclusions
The staminal lever of S. pratensis is a pollen-dispensing device. It seems to influence the amount of pollen deposited on pollinators by determining the forces arising between the pollinator and the pollen. The relevant forces occur either during the first, dynamic phase or during the second, almost static phase of a flower visit.
doi:10.1093/aob/mcm031
PMCID: PMC2735310  PMID: 17416914
Flower–pollinator interaction; bee; Apis mellifera; pollination; pollen uptake; see-saw mechanism; biomechanics; pollen dispensing

Results 1-6 (6)