OIL FLOWERS AND OIL BEES: FURTHER EVIDENCE FOR POLLINATOR ADAPTATION

We examined foreleg length and body size variation in two species of oil-collecting bees (Rediviva; Melittidae) in southern Africa. Oil-collecting bees harvest oil from host flowers by rubbing their forelegs against oil-secreting trichomes. Significant differences in foreleg length occur among populations of both species. Rediviva “pallidula” populations vary significantly in mean foreleg length (11.34 ± 0.42 mm to 12.67 ± 0.36 mm), but not in body length (10.59 ± 0.74 to 10.80 ± 0.64), and foreleg length and body size are not significantly correlated. Instead, foreleg variation appears to be a function of host plant spur length. Ninety-two percent of the variance in foreleg length of R. “pallidula” is explained by mean Diascia spur length. Rediviva rufocincta populations vary significantly in mean foreleg length (10.12 ± 0.70 mm to 12.34 ± 0.68 mm) and in body length (9.03 ± 0.26 mm to 10.56 ± 0.24 mm). Foreleg length scales allometrically with body size in this species as 90.5% of the variance in foreleg length can be explained as a function of body length. Body size appears to be constrained by the morphology of the oil-secreting host plant. Both bees collect floral oil with specially modified setae on the tarsi of their forelegs. The length of the disti- + mediotarsus (refered to here as “tarsus”) in relation to the entire foreleg is shorter in R. rufocincta and does not increase as rapidly with increasing foreleg length as for R. “pallidula.” These differences in variation can be attributed to differences in position of oil within the flowers of the respective host plants. Rediviva “pallidula” collects oil from Diascia species that have the oil deeply situated in narrow floral spurs of varying length, while R. rufocincta collects oil from the broadly saccate flowers of Bowkeria verticillata and B. citrina.     

Floral adaptation to local pollinator guilds in a terrestrial orchid

Background and Aims

Studies of local floral adaptation in response to geographically divergent pollinators are essential for understanding floral evolution. This study investigated local pollinator adaptation and variation in floral traits in the rewarding orchid Gymnadenia odoratissima, which spans a large altitudinal gradient and thus may depend on different pollinator guilds along this gradient.

Methods

Pollinator communities were assessed and reciprocal transfer experiments were performed between lowland and mountain populations. Differences in floral traits were characterized by measuring floral morphology traits, scent composition, colour and nectar sugar content in lowland and mountain populations.

Key Results

The composition of pollinator communities differed considerably between lowland and mountain populations; flies were only found as pollinators in mountain populations. The reciprocal transfer experiments showed that when lowland plants were transferred to mountain habitats, their reproductive success did not change significantly. However, when mountain plants were moved to the lowlands, their reproductive success decreased significantly. Transfers between populations of the same altitude did not lead to significant changes in reproductive success, disproving the potential for population-specific adaptations. Flower size of lowland plants was greater than for mountain flowers. Lowland plants also had significantly higher relative amounts of aromatic floral volatiles, while the mountain plants had higher relative amounts of other floral volatiles. The floral colour of mountain flowers was significantly lighter compared with the lowland flowers.

Conclusions

Local pollinator adaptation through pollinator attraction was shown in the mountain populations, possibly due to adaptation to pollinating flies. The mountain plants were also observed to receive pollination from a greater diversity of pollinators than the lowland plants. The different floral phenotypes of the altitudinal regions are likely to be the consequence of adaptations to local pollinator guilds.     

具有泛化访花者的海芋特化传粉系统

特化传粉系统通常被认为是一种高效的植物与传粉者互作的关系, 但即使是高度特化的花朵也经常接受非传粉者的访问。当前的特化传粉系统研究主要关注于植物与特定传粉者的相互作用, 往往忽视了其他访花者的潜在影响。海芋(Alocasia odora)与芋果蝇属(Colocasiomyia)物种是典型的高度特化传粉互惠关系, 但海芋花序仍存在许多其他的访花者类群, 它们对传粉过程的影响尚不清楚。通过访花者筛除实验, 本研究证实了海芋的授粉过程必须有海芋果蝇(C. alocasia)或异海芋果蝇(C. xenalocasiae)的参与。除此以外, 海芋花序上还观察到包括露尾甲科、蜜蜂科、隐翅虫科和跗线螨科等类群的访花者, 共计10种。繁育系统实验表明, 海芋的自交不亲和性是由空间和时间上的雌雄分离造成的。因此, 尽管其他访花者未直接对海芋结实率做出贡献, 但它们对花粉和雄花不育区分泌物报酬的取食行为仍可能整体上减少了有效花粉资源, 并与传粉者竞争报酬资源, 进而间接影响有效传粉者的传粉行为。本研究为特化传粉理论提供了新视角, 未来研究应更全面考虑访花者的整体作用。     

Pollination syndromes in a specialised plant‐pollinator interaction: does floral morphology predict pollinators in Calceolaria?

Pollination syndromes are defined as suites of floral traits evolved in response to selection imposed by a particular group of pollinators (e.g., butterflies, hummingbirds, bats). Although numerous studies demonstrated their occurrence in plants pollinated by radically different pollinators, it is less known whether it is possible to identify them within species pollinated by one functional pollinator group. In such a framework, we expect floral traits to evolve also in response to pollinator subgroups (e.g., species, genera) within that unique functional group. On this, specialised pollination systems represent appropriate case studies to test such expectations. Calceolaria is a highly diversified plant genus pollinated by oil‐collecting bees in genera Centris and Chalepogenus. Variation in floral traits in Calceolaria has recently been suggested to reflect adaptations to pollinator types. However, to date no study has explicitly tested that observation. In this paper, we quantitatively test that hypothesis by evaluating the presence of pollination syndromes within the specialised pollination system formed by several Calceolaria and their insect pollinators. To do so, we use multivariate approaches and explore the structural matching between the morphology of 10 Calceolaria taxa and that of their principal pollinators. Our results identify morphological matching between floral traits related to access to the reward and insect traits involved in oil collection, confirming the presence of pollinator syndromes in Calceolaria. From a general perspective, our findings indicate that the pollination syndrome concept can be also extended to the intra‐pollinator group level.     

Pterandra pyroidea: a case of pollination shift within neotropical Malpighiaceae

Background and Aims

Most Neotropical species of Malpighiaceae produce floral fatty oils in calyx glands to attract pollinating oil-collecting bees, which depend on this resource for reproduction. This specialized type of pollination system tends to be lost in members of the family that occur outside the geographic distribution (e.g. Africa) of Neotropical oil-collecting bees. This study focused on the pollination ecology, chemical ecology and reproductive biology of an oil flower species, Pterandra pyroidea (Malpighiaceae) from the Brazilian Cerrado. Populations of this species consist of plants with oil-secreting (glandular) flowers, plants with non-oil-secreting flowers (eglandular) or a mix of both plant types. This study specifically aims to clarify the role of eglandular morphs in this species.

Methods

Data on pollinators were recorded by in situ observations. Breeding system experiments were conducted by isolating inflorescences and by enzymatic reactions. Floral resources, pollen and floral oils offered by this species were analysed by staining and a combination of various spectroscopic methods.

Key Results

Eglandular flowers of P. pyroidea do not act as mimics of their oil-producing conspecifics to attract pollinators. Instead, both oil-producing and oil-free flowers depend on pollen-collecting bees for reproduction, and their main pollinators are bumble-bees. Floral oils produced by glandular flowers are less complex than those described in closely related genera.

Conclusions

Eglandular flowers represent a shift in the pollination system in which oil is being lost and pollen is becoming the main reward of P. pyroidea flowers. Pollination shifts of this kind have hitherto not been demonstrated empirically within Neotropical Malpighiaceae and this species exhibits an unusual transition from a specialized towards a generalized pollination system in an area considered the hotspot of oil-collecting bee diversity in the Neotropics. Transitions of this type provide an opportunity to study ongoing evolutionary mechanisms that promote the persistence of species previously involved in specialized mutualistic relationships.     

Stability of floral specialization in Trollius europaeus in contrasting ecological environments

Specialization of some plants on seed‐eating pollinators is intriguing, especially when co‐pollinators exclusively feeding on nectar are also present. We examined the stability of the morphological specialization of Trollius europaeus (L.) globeflowers with respect to Chiastocheta (Pokorny) flies by artificially opening the flowers. In the montane and subalpine environments studied, other visitors contributed 2% and 28% of all the visits, respectively, and visited open flowers nearly eight times more often than closed flowers, but in both environments their contribution to pollination did not compensate for Chiastocheta aversion against open phenotypes. Net seed set (female success) was slightly higher (+4%) and pollen export (male success) was much higher (+85%) for closed than open flowers. Selection in favour of the closed phenotype was even more intense in patches where open phenotypes were most common, precluding the evolution of open flowers in the study populations.     

Geographical differentiation in floral traits across the distribution range of the Patagonian oil-secreting Calceolaria polyrhiza: do pollinators matter?

Background and Aims

The underlying evolutionary processes of pollinator-driven floral diversification are still poorly understood. According to the Grant–Stebbins model speciation begins with adaptive local differentiation in the response to spatial heterogeneity in pollinators. Although this crucial process links the micro- and macroevolution of floral adaptation, it has received little attention. In this study geographical phenotypic variation was investigated in Patagonian Calceolaria polyrhiza and its pollinators, two oil-collecting bee species that differ in body size and geographical distribution.

Methods

Patterns of phenotypic variation were examined together with their relationships with pollinators and abiotic factors. Six floral and seven vegetative traits were measured in 45 populations distributed across the entire species range. Climatic and edaphic parameters were determined for 25 selected sites, 2–16 bees per site of the most frequent pollinator species were captured, and a critical flower–bee mechanical fitting trait involved in effective pollination was measured. Geographical patterns of phenotypic and environmental variation were examined using uni- and multivariate analyses. Decoupled geographical variation between corolla area and floral traits related to the mechanical fit of pollinators was explored using a Mantel test.

Key Results

The body length of pollinators and the floral traits related to mechanical fit were strongly correlated with each other. Geographical variation of the mechanical-fit-related traits was decoupled from variation in corolla size; the latter had a geographical pattern consistent with that of the vegetative traits and was mainly affected by climatic gradients.

Conclusions

The results are consistent with pollinators playing a key role in shaping floral phenotype at a geographical scale and promoting the differentiation of two floral ecotypes. The relationship between the critical floral-fit-related trait and bee length remained significant even in models that included various environmental variables and an allometric predictor (corolla area). The abiotic environment also has an important role, mainly affecting floral size. Decoupled geographical variation between floral mechanical-fit-related traits and floral size would represent a strategy to maintain plant–pollinator phenotypic matching in this environmentally heterogeneous area.     

Reproductive biology of two Himalayan alpine gingers (Roscoea spp., Zingiberaceae) in China: pollination syndrome and compensatory floral mechanisms

According to the concept of pollination syndromes, floral traits reflect specialisation to a particular pollinator or set of pollinators. However, the reproductive biology of endemic, and often specialised, plants may require increased attention as climate change accelerates worldwide. Species of Roscoea endemic to the Himalayan region have striking orchid-like flowers with long corolla tubes, suggesting pollination by long-tongued insects. Until now, the reproductive biology of species of Roscoea has been poorly documented. We investigated the floral biology, breeding system and pollination ecology of R. cautleoides and R. humeana, from Hengduan Mountains, a global biodiversity hotspot in southwest China. We also tested whether floral longevity increases pollination success. Pollination experiments showed that the two species were self-compatible and depended on insects for fruit production. Over several flowering seasons we did not observe any potential pollinators with long tongues that matched the corolla tube visiting flowers in centres of distribution. The principal pollinators observed were pollen-collecting generalist bees, with low visitation frequencies. In general, members of the ginger family are characterised by short-lived (usually 1 day) flowers, but flowers of R. cautleoides and R. humeana last 8 and 6 days, respectively. Removing stigmas decreased fruit set in both study populations. Our results suggest that the original pollinators may have been long-tongued insects that are now absent from the Chinese Himalayas because habitats have responded to climate change. However, long-lived and self-compatible flowers, coupled with the presence of generalist pollinators, are traits that have allowed these gingers to reproduce and continue to persist in the alpine habitats.