Floral evolution and pollinator diversification in Hedychium J.Koenig (Zingiberaceae): one of Mr. Darwin’s tropical fantasies

ABSTRACTPREMISEHedychium J.Koenig (ginger lilies: Zingiberaceae) is endemic to the Indo-Malayan Realm (IMR) and is known for its fragrant flowers. Two different pollination syndromes characterize the genus: diurnal or bird pollination and nocturnal or moth pollination systems. To date, no attempt has been undertaken to understand the evolution of floral traits in this genus.METHODSWe estimated ancestral character-states, phylogenetic signals, and character correlations for thirteen discrete and eight continuous floral traits representing 75% species diversity of Hedychium. Diversification rate estimation analyses were also employed to understand trait-dependent diversification in the genus.RESULTSInflorescence structure, cincinnus capacity, and curvature of floral tubes revealed strong phylogenetic dependence, whereas number of open flowers per inflorescence per day, color of the labellum, and exertion of the stigma characterized higher ecological effects. Diversification rate estimations suggested that the labellum width, floral tube length, and labellum color played a major role in the evolutionary diversification of Hedychium.CONCLUSIONSWe identified bract type and cincinnus capacity as synapomorphies for Hedychium, while the island-specific clade III was characterized by slender cylindrical inflorescence, coiling of floral tubes, and longer bract to calyx ratio. The circum-Himalayan clade IV is the most speciose, derived, and with most variable floral traits. Although floral color and size lacked any association with pollinator-specific traits (moth and bird pollination), pale colored flowers were most common in the early diverging clades (clade I, II-el., and II-de.), indicating their ancestral nature, when compared to brightly colored flowers.

Three options are better than two: compensatory nature of different pollination modes in Salix caprea L.

Mixed pollination systems have been reported for many angiosperms. Here, we report on a dioecious willow, Salix caprea L., which employs three pollination modes, namely pollination by wind, nocturnal moths, and diurnal bees. In four field populations of S. caprea in Germany, we addressed the following questions: (a) How are the pollination modes affected by local mate availability? (b) What is the contribution of each pollination mode to seed set? (c) How does flower opening time affect pollination? We compared seed set among seven pollination treatments (wind pollination, diurnal pollination, nocturnal pollination, natural pollination, apomictic reproduction, daytime/night-time artificial pollination). Pollen limitation was observed across populations. On the other hand, limitations in mate availability affected the pollination modes differently. Insect pollination outperformed wind pollination in situations where nearby males were few or all positioned in the same direction from the females. The contribution of nocturnal moths was lower than that of the other pollen vectors. However, moth pollination worked complementarily with bee pollination in compensating for the lack of wind pollination. Furthermore, we found that the onset of flower anthesis peaked at sunset, and that cold night temperatures had no negative effect on seed set or pollen viability. Thus, nocturnal anthesis and prior pollination by moths may minimize male fitness loss due to pollen collection and grooming by bees. We suggest that S. caprea maximizes its fitness by combining multiple pollination modes that work in different environmental conditions, while optimizing the anthesis so that more pollen would be delivered to stigmas.

Testing Pollination Syndromes in Oenothera (Onagraceae)

Pollinators are considered a major selective force in shaping the diversification of angiosperms. It has been hypothesized that convergent evolution of floral form has resulted in “pollination syndromes” - i.e. suites of floral traits that correspond to attraction of particular pollinator functional groups. Across the literature, the pollination syndrome concept has received mixed support. This may be due to studies using different methods to describe floral traits and/or the pollination syndrome concept being supported more often in species highly reliant on pollinators for reproduction. Here, we assess the predictive ability of pollination syndromes in Oenothera, a species rich clade with pollination systems existing on a gradient of specialization, and in which species are either self-compatible or self-incompatible. We ask the following questions: Do Oenothera species follow the pollination syndrome concept using traditional, categorical floral trait descriptions and/or quantitative floral trait measurements? And, are floral traits more predictive of primary pollinators in species with specialized pollination systems and/or species that are self-incompatible? Mapping floral traits of 54 Oenothera species into morphospace, we do not find support for the pollination syndrome concept using either categorical or quantitative floral trait descriptions. We do not find support for specialization or breeding system influencing the prediction of primary pollinators. However, we find pollination syndromes were more predictive in Oenothera species with moth pollination systems. Collectively, these results suggest that the pollination syndrome concept cannot be generally applied across taxa and that evolutionary history is important to consider when evaluating the relationship between floral form and contemporary pollinators.

Diurnal hawk moth pollination of Melampyrum koreanum (Orobanchaceae) and the origin of this endemic Korean species

Melampyrum koreanum (Orobanchaceae) was described recently as an endemic species in Korea. This study examined the distribution range, relationship with the host plant, flower developmental pattern, pollinator, pollination mechanism, and seed development of M. koreanum (Orobanchaceae) on the island of Somaemul-do, Korea. M. koreanum is hemiparasitic and a strong candidate for its host plant is Pinus thunbergii, with which it is distributed sympatrically. The flower of M. koreanum represents a typical hawk moth pollination syndrome. Macroglossum pyrrhostictum Butler (Sphingidae, Lepidoptera), a diurnal hawk moth, is the only efficient pollinator. The proboscis length of M. pyrrhostictum and the floral tube length of M. koreanum are well coordinated, and the positions of the anther and stigma of M. koreanum are mechanically conformed to hawk moth pollination. The results suggest that adaptation to the pollinator is the main factor that has affected the evolution of the longer floral tube of M. koreanum compare to other species. Pollinator conservation in the native range of M. koreanum is an important issue for the conservation of endemic plant species, and it is necessary to establish a conservation plan for the host plant, Pinus thunbergii, as well as M. koreanum.

The trichotomosulcate asparagoids: pollen morphology of Hemerocallidaceae in relation to systematics and pollination biology

We examined pollen of 19 genera of Hemerocallidaceae by using scanning electron microscopy (SEM), and one genus (Dianella) by using transmission electron microscopy (TEM). Pollen was generally small in size, with a rounded triangular outline when hydrated, and a characteristic three-armed aperture, a distal trichotomosulcus. The pollen surface was finely sculptured and the exine was thin. Microreticulate pollen is a potential synapomorphy for several species of the ‘crown phormioid’ subclade recognised in molecular analyses. Perforate and fossulate pollen supports a relationship between several species of Dianella. Microrugulate pollen is more frequent in the johnsonioids than in the phormioids. Hemerocallis is distinguished by elongated monosulcate pollen, a relatively thick exine with a pronounced reticulate surface, and large globules of attached pollenkitt. We hypothesise that Hemerocallidaceae are ancestrally buzz-pollinated, and their pollen morphology is an adaptation to this pollination type. A reversal to butterfly or moth pollination occurred in Hemerocallis, with associated changes in pollen morphology.