Evolutionary constraints on disparity of ericaceous pollen grains.

BACKGROUND AND AIMS: Flowering plants show a high diversity of pollen morphology, assumed to reflect not only variations in the underlying design, but also stress imposed by ecological conditions related to pollen survival and germination. Both components are expected to constrain the accumulation of pollen disparity. However, this assumption has rarely been tested using empirical data. METHODS: This study is designed to test this hypothesis by inferring the accumulation of pollen disparity in Ericaceae, a large eudicot family with recent, ongoing radiations, with focus on three functionally significant pollen characters using a dated phylogeny. KEY RESULTS: Multiple lines of evidence supported the hypothesis that pollen disparity in Ericaceae did not evolve steadily but rather pulsed over time, clearly decoupling from the relative constant rate pattern of species diversification inferred. In a 3-D pollen morphospace, most major clades appear to occupy distinct neighbouring regions, whereas the subfamily Epacridoideae overlaps extensively with other subfamilies. No evidence for correlations was found between dimension of pollen disparity and species diversity at either the subfamily or generic level. Furthermore, the distribution of species in present pollen morphospace showed a strong central tendency, with the core compartment containing a large number of species from species-rich genera. CONCLUSIONS: The recovered evidence fits well with the expectations of limitations on available pollen morphological disparity, and suggests that innovation of pollen germination traits may have little effect on species diversification.

Systematic importance of morphological features of pollen grains of species from Erica (Ericaceae) genus.

The Erica genus has as yet not been investigated satisfactorily in terms of palynology. Its complicated taxonomic system, large number of species, as well as its extensive but disrupted range of occurrence, all contribute to the fact that few researchers have undertaken investigations of this species. It was assumed that the research results would be representative thanks to a complex comparative analysis of all diagnostical, morphological pollen features performed on properly selected plant material, representing the most important distinguished intrageneric taxons at the present time (45 species from all five subgenera and 22 sections), both discriminated pollen dispersal units (tetrads and monads) as well as the main centres of genus occurrence and diversification (species from Europe, the Republic of South Africa (RSA) and Madagascar). The study revealed that the diagnostic features of the pollen grains studied were: pollen dispersal unit, exine ornamentation, P/E ratio, tetrad diameter (D) and length of polar axis (P). On the basis of these traits, 14 Erica species (six creating monads and eight-tetrads) were distinguished which, in the case of pollen features, constitutes a significant number. Other heaths created small groups, usually containing two or three species, but up to seven species. The present study, based on the highest number of Erica species (45) analysed so far, corroborated the view that an examination of palynological features may assist in clarifying classification systems for the large and taxonomically very difficult Erica genus, in particular, at the level of the subgenus and section, but also at species level. The results obtained indicate the need to continue palynological investigations on the Erica genus.

Phylogeny of Pyroleae (Ericaceae): implications for character evolution.

Pyroleae (Ericaceae) consist of four genera, all of which are distributed widely in temperate coniferous or sometimes deciduous forests of the Northern Hemisphere. To investigate the phylogenetic relationships among these genera and to explore the evolution of the characteristics of the subfamily, we conducted maximum parsimony and Bayesian analyses with nrDNA ITS and three cpDNA intergenic spacers (atpB-rbcL, trnS-trnG and trnL-trnF). The results from cpDNA and combined cpDNA + ITS data sets strongly support the monophyly of Pyroleae as well as a sister relationship between Pyrola and Moneses-Chimaphila, with Orthilia as the basal lineage. The sister-group relationship between Moneses and Chimaphila is supported by a set of synapomorphies, e.g., single flower, colpate pollen, five bundles in the style, straight fruiting pedicel orientation, complete capsule dehiscence, and the basic chromosome number, x = 13. The Moneses-Chimaphila-Pyrola clade is supported by at least one homologous character of pollen in tetrads. Conflicts associated with the phylogenetic position of Orthilia may imply a hybrid origin for it, and therefore further study is needed.

Pollen of Ceratostema (Ericaceae, Vaccinieae): tetrads without septa.

The pollen morphology of two species of the Neotropical genus Ceratostema (Ericaceae) was examined by light, scanning and transmission electron microscopy. The Ceratostema species examined have 3-colporate pollen grains united in permanent tetrahedral tetrads that show a common condition encountered in the Ericaceae. But the septal exine was absent between two neighboring grains in each pollen tetrad of Ceratostema. The pollen tetrads without septa are the first report for the Ericaceae as well as other angiosperm families.

[Mechanisms that limit pollinator range in Ericaceae].

Studies were conducted in 2001-2003 at Valdai National Park (Novgorod region) and at the Zvenigorod biological station of Moscow State University. The morphology of flowers, flowering dynamics and composition of insect visiting flowers of Ericaceae species: Andromeda polifolia, Chamaedaphne calyculata, Ledum palustre, Oxycoccus palustris, Vaccinium myrtillus, V. uliginosum, and V. vitis-idaea L. were studied. Some species of insects visiting flowers were excluded from the list of pollinators on the basis of observation on their behavior. L. palustre was visited mainly by flies where as other investigated species were visited mainly by bumblebees. In some cases bumblebees were the only visitors of the investigated plants. Mechanisms that protect flowers from flies and short-tongued solitary bees visits and ensure a best pollination by bumblebees are various among different species of Ericaceae. Efficiency of nectary protection also differs among different plant species and is defined by particularities of their habitats and flowering phenology. As far as all species of this family during the flowering are dominants in typical habitats, a competition for the pollination with species of other families in most cases is megligible. Flowering periods of V. vitis-idaea and V. myrtillus in forest ecosystems overlapped weakly. Moreover, V. myrtillus is pollinated mainly by bumblebee queens where as pollinators of V. vitis-idaea are bumblebee workers, solitary bees and horse flies. The other investigated plant species inhabit only oligotrophic peat bogs. Thery are pollinated by bumblebees but periods of flowering are not overlapped and consequently follow one after another. L. palustre and V. uliginosum flower simultaneosly but they are pollinated by different pollinators.