Alpine dwarf shrubs show high proportions of nonfunctional xylem: Visualization and quantification of species-specific patterns.

Xylem conductive capacity is a key determinant of plant hydraulic function and intimately linked to photosynthesis and productivity, but can be impeded by temporary or permanent conduit dysfunctions. Here we show that persistent xylem dysfunctions in unstressed plants are frequent in Alpine dwarf shrubs and occur in various but species-specific cross-sectional patterns. Combined synchrotron micro-computed tomography (micro-CT) imaging, xylem staining, and flow measurements in saturated samples of six widespread Ericaceae species evidence a high proportion (19%-50%) of hydraulically nonfunctional xylem areas in the absence of drought stress, with regular distribution of dysfunctions between or within growth rings. Dysfunctions were only partly reversible and reduced the specific hydraulic conductivity to 1.38 to 3.57 ×10-4 m2 s-1 MPa-1 . Decommission of inner growth rings was clearly related to stem age and a higher vulnerability to cavitation of older rings, while the high proportion of nonfunctional conduits in each annual ring needs further investigations. The lower the xylem fraction contributing to the transport function, the higher was the hydraulic efficiency of conducting xylem areas. Improved understanding of the functional lifespan of xylem elements and the prevalence and nature of dysfunctions is critical to correctly assess structure-function relationships and whole-plant hydraulic strategies.

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.

In vitro axenic germination and cultivation of mixotrophic Pyroloideae (Ericaceae) and their post-germination ontogenetic development.

BACKGROUND AND AIMS: Pyroloids, forest sub-shrubs of the Ericaceae family, are an important model for their mixotrophic nutrition, which mixes carbon from photosynthesis and from their mycorrhizal fungi. They have medical uses but are difficult to cultivate ex situ; in particular, their dust seeds contain undifferentiated, few-celled embryos, whose germination is normally fully supported by fungal partners. Their germination and early ontogenesis thus remain elusive. METHODS: An optimized in vitro cultivation system of five representatives from the subfamily Pyroloideae was developed to study the strength of seed dormancy and the effect of different media and conditions (including light, gibberellins and soluble saccharides) on germination. The obtained plants were analysed for morphological, anatomical and histochemical development. KEY RESULTS: Thanks to this novel cultivation method, which breaks dormancy and achieved up to 100 % germination, leafy shoots were obtained in vitro for representatives of all pyroloid genera (Moneses, Orthilia, Pyrola and Chimaphila). In all cases, the first post-germination stage is an undifferentiated structure, from which a root meristem later emerges, well before formation of an adventive shoot. CONCLUSIONS: This cultivation method can be used for further research or for ex situ conservation of pyroloid species. After strong seed dormancy is broken, the tiny globular embryo of pyroloids germinates into an intermediary zone, which is functionally convergent with the protocorm of other plants with dust seeds such as orchids. Like the orchid protocorm, this intermediary zone produces a single meristem: however, unlike orchids, which produce a shoot meristem, pyroloids first generate a root meristem.

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.

The light response of mesophyll conductance is controlled by structure across leaf profiles.

Mesophyll conductance to CO2 (gm ) may respond to light either through regulated dynamic mechanisms or due to anatomical and structural factors. At low light, some layers of cells in the leaf cross-section approach photocompensation and contribute minimally to bulk leaf photosynthesis and little to whole leaf gm (gm,leaf ). Thus, the bulk gm,leaf will appear to respond to light despite being based upon cells having an anatomically fixed mesophyll conductance. Such behaviour was observed in species with contrasting leaf structure using the variable J or stable isotope method of measuring gm,leaf . A species with bifacial structure, Arbutus × 'Marina', and an isobilateral species, Triticum durum L., had contrasting responses of gm,leaf upon varying adaxial or abaxial illumination. Anatomical observations, when coupled with the proposed model of gm,leaf to photosynthetic photon flux density (PPFD) response, successfully represented the observed gas exchange data. The theoretical and observed evidence that gm,leaf apparently responds to light has large implications for how gm,leaf values are interpreted, particularly limitation analyses, and indicates the importance of measuring gm under full light saturation. Responses of gm,leaf to the environment should be treated as an emergent property of a distributed 3D structure, and not solely a leaf area-based phenomenon.

Leaf anatomy, BVOC emission and CO2 exchange of arctic plants following snow addition and summer warming.

BACKGROUND AND AIMS: Climate change in the Arctic is projected to increase temperature, precipitation and snowfall. This may alter leaf anatomy and gas exchange either directly or indirectly. Our aim was to assess whether increased snow depth and warming modify leaf anatomy and affect biogenic volatile organic compound (BVOC) emissions and CO2 exchange of the widespread arctic shrubs Betula nana and Empetrum nigrum ssp. hermaphroditum METHODS: Measurements were conducted in a full-factorial field experiment in Central West Greenland, with passive summer warming by open-top chambers and snow addition using snow fences. Leaf anatomy was assessed using light microscopy and scanning electron microscopy. BVOC emissions were measured using a dynamic enclosure system and collection of BVOCs into adsorbent cartridges analysed by gas chromatography-mass spectrometry. Carbon dioxide exchange was measured using an infrared gas analyser. KEY RESULTS: Despite a later snowmelt and reduced photosynthesis for B. nana especially, no apparent delays in the BVOC emissions were observed in response to snow addition. Only a few effects of the treatments were seen for the BVOC emissions, with sesquiterpenes being the most responsive compound group. Snow addition affected leaf anatomy by increasing the glandular trichome density in B. nana and modifying the mesophyll of E. hermaphroditum The open-top chambers thickened the epidermis of B. nana, while increasing the glandular trichome density and reducing the palisade:spongy mesophyll ratio in E. hermaphroditum CONCLUSIONS: Leaf anatomy was modified by both treatments already after the first winter and we suggest links between leaf anatomy, CO2 exchange and BVOC emissions. While warming is likely to reduce soil moisture, melt water from a deeper snow pack alleviates water stress in the early growing season. The study emphasizes the ecological importance of changes in winter precipitation in the Arctic, which can interact with climate-warming effects.

Low among-provenance differences in structural and functional plasticity in response to nutrients in saplings of the circum-Mediterranean tree Arbutus unedo L.

The Mediterranean region is an area of special interest for conservation where the incidence of multiple drivers of global change is expected to increase. One of the factors predicted to change is soil-nutrient availability, an essential factor for plant growth. Thus, study of the effects of variation in this parameter is especially relevant in species with a circum-Mediterranean distribution, such as Arbutus unedo L., in which the different provenances grow in different habitats, which must differ in nutritional conditions. We aimed to determine the effect of provenance on plasticity, to establish whether structural and morphological traits differ in the level of plasticity and to assess how nutrients affect the photosynthetic light response. In a common garden experiment, we studied seven provenances from the circum-Mediterranean range of A. unedo and established two nutrient treatments (low and high nutrient availability). We measured physiological and structural traits in 1-year-old sapling and determined a phenotypic plasticity index (PPI) to quantify the level of plasticity, whereas the radiation effects were tested by construction and analysis of light response curves. Interestingly, provenance did not explain a significant amount of variance, but the plasticity was four times higher for the structural traits than for the physiological traits. Therefore, the plasticity to nutrient availability will not favour or prevent the expansion or contraction of the range of any of these provenances of A. unedo. Furthermore, the structural plasticity demonstrated the ability of the strawberry tree to optimize resource allocation, whereas the physiology remained stable, thus avoiding extra expenditure. The study findings also suggest that increased availability of nutrients would improve the performance of the species during the Mediterranean summer, characterized by high irradiance. These abilities will be key to the survival of saplings of the species under the future scenario of changes in nutrient availability.

Seasonal variability of foliar photosynthetic and morphological traits and drought impacts in a Mediterranean mixed forest.

The Mediterranean region is a hot spot of climate change vulnerable to increased droughts and heat waves. Scaling carbon fluxes from leaf to landscape levels is particularly challenging under drought conditions. We aimed to improve the mechanistic understanding of the seasonal acclimation of photosynthesis and morphology in sunlit and shaded leaves of four Mediterranean trees (Quercus ilex L., Pinus halepensis Mill., Arbutus unedo L. and Quercus pubescens Willd.) under natural conditions. Vc,max and Jmax were not constant, and mesophyll conductance was not infinite, as assumed in most terrestrial biosphere models, but varied significantly between seasons, tree species and leaf position. Favourable conditions in winter led to photosynthetic recovery and growth in the evergreens. Under moderate drought, adjustments in the photo/biochemistry and stomatal/mesophyllic diffusion behaviour effectively protected the photosynthetic machineries. Severe drought, however, induced early leaf senescence mostly in A. unedo and Q. pubescens, and significantly increased leaf mass per area in Q. ilex and P. halepensis. Shaded leaves had lower photosynthetic potentials but cushioned negative effects during stress periods. Species-specificity, seasonal variations and leaf position are key factors to explain vegetation responses to abiotic stress and hold great potential to reduce uncertainties in terrestrial biosphere models especially under drought conditions.

Climate change alters leaf anatomy, but has no effects on volatile emissions from Arctic plants.

Biogenic volatile organic compound (BVOC) emissions are expected to change substantially because of the rapid advancement of climate change in the Arctic. BVOC emission changes can feed back both positively and negatively on climate warming. We investigated the effects of elevated temperature and shading on BVOC emissions from arctic plant species Empetrum hermaphroditum, Cassiope tetragona, Betula nana and Salix arctica. Measurements were performed in situ in long-term field experiments in subarctic and high Arctic using a dynamic enclosure system and collection of BVOCs into adsorbent cartridges analysed by gas chromatography-mass spectrometry. In order to assess whether the treatments had resulted in anatomical adaptations, we additionally examined leaf anatomy using light microscopy and scanning electron microscopy. Against expectations based on the known temperature and light-dependency of BVOC emissions, the emissions were barely affected by the treatments. In contrast, leaf anatomy of the studied plants was significantly altered in response to the treatments, and these responses appear to differ from species found at lower latitudes. We suggest that leaf anatomical acclimation may partially explain the lacking treatment effects on BVOC emissions at plant shoot-level. However, more studies are needed to unravel why BVOC emission responses in arctic plants differ from temperate species.

Do pollinator distributions underlie the evolution of pollination ecotypes in the Cape shrub Erica plukenetii?

BACKGROUND AND AIMS: According to the Grant-Stebbins model of pollinator-driven divergence, plants that disperse beyond the range of their specialized pollinator may adapt to a new pollination system. Although this model provides a compelling explanation for pollination ecotype formation, few studies have directly tested its validity in nature. Here we investigate the distribution and pollination biology of several subspecies of the shrub Erica plukenetii from the Cape Floristic Region in South Africa. We analyse these data in a phylogenetic context and combine these results with information on pollinator ranges to test whether the evolution of pollination ecotypes is consistent with the Grant-Stebbins model. METHODS AND KEY RESULTS: Pollinator observations showed that the most common form of E. plukenetii with intermediate corolla length is pollinated by short-billed Orange-breasted sunbirds. Populations at the northern fringe of the distribution are characterized by long corollas, and are mainly pollinated by long-billed Malachite sunbirds. A population with short corollas in the centre of the range was mainly pollinated by insects, particularly short-tongued noctuid moths. Bird exclusion in this population did not have an effect on fruit set, while insect exclusion reduced fruit set. An analysis of floral scent across the range, using coupled gas chromatography-mass spectrometry, showed that the scent bouquets of flowers from moth-pollinated populations are characterized by a larger number of scent compounds and higher emission rates than those in bird-pollinated populations. This was also reflected in clear separation of moth- and bird-pollinated populations in a two-dimensional phenotype space based on non-metric multidimensional scaling analysis of scent data. Phylogenetic analyses of chloroplast and nuclear DNA sequences strongly supported monophyly of E. plukenetii, but not of all the subspecies. Reconstruction of ancestral character states suggests two shifts from traits associated with short-billed Orange-breasted sunbird pollination: one towards traits associated with moth pollination, and one towards traits associated with pollination by long-billed Malachite sunbirds. The latter shift coincided with the colonization of Namaqualand in which Orange-breasted sunbirds are absent. CONCLUSIONS: Erica plukenetii is characterized by three pollination ecotypes, but only the evolutionary transition from short- to long-billed sunbird pollination can be clearly explained by the Grant-Stebbins model. Corolla length is a key character for both ecotype transitions, while floral scent emission was important for the transition from bird to moth pollination.

Growth and community responses of alpine dwarf shrubs to in situ CO2 enrichment and soil warming.

• Rising CO2 concentrations and the associated global warming are expected to have large impacts on high-elevation ecosystems, yet long-term multifactor experiments in these environments are rare. • We investigated how growth of dominant dwarf shrub species (Vaccinium myrtillus, Vaccinium gaultherioides and Empetrum hermaphroditum) and community composition in the understorey of larch and pine trees responded to 9 yr of CO2 enrichment and 3 yr of soil warming at the treeline in the Swiss Alps. • Vaccinium myrtillus was the only species that showed a clear positive effect of CO2 on growth, with no decline over time in the annual shoot growth response. Soil warming stimulated V. myrtillus growth even more than elevated CO2 and was accompanied by increased plant-available soil nitrogen (N) and leaf N concentrations. Growth of Vaccinium gaultherioides and E. hermaphroditum was not influenced by warming. Vascular plant species richness declined in elevated CO2 plots with larch, while the number of moss and lichen species decreased under warming. • Ongoing environmental change could lead to less diverse plant communities and increased dominance of the particularly responsive V. myrtillus in the studied alpine treeline. These changes are the consequence of independent CO2 and soil warming effects, a result that should facilitate predictive modelling approaches.

Effects of nutrient addition on leaf chemistry, morphology, and photosynthetic capacity of three bog shrubs.

Plants in nutrient-poor environments typically have low foliar nitrogen (N) concentrations, long-lived tissues with leaf traits designed to use nutrients efficiently, and low rates of photosynthesis. We postulated that increasing N availability due to atmospheric deposition would increase photosynthetic capacity, foliar N, and specific leaf area (SLA) of bog shrubs. We measured photosynthesis, foliar chemistry and leaf morphology in three ericaceous shrubs (Vaccinium myrtilloides, Ledum groenlandicum and Chamaedaphne calyculata) in a long-term fertilization experiment at Mer Bleue bog, Ontario, Canada, with a background deposition of 0.8 g N m(-2) a(-1). While biomass and chlorophyll concentrations increased in the highest nutrient treatment for C. calyculata, we found no change in the rates of light-saturated photosynthesis (A(max)), carboxylation (V(cmax)), or SLA with nutrient (N with and without PK) addition, with the exception of a weak positive correlation between foliar N and A(max) for C. calyculata, and higher V(cmax) in L. groenlandicum with low nutrient addition. We found negative correlations between photosynthetic N use efficiency (PNUE) and foliar N, accompanied by a species-specific increase in one or more amino acids, which may be a sign of excess N availability and/or a mechanism to reduce ammonium (NH(4)) toxicity. We also observed a decrease in foliar soluble Ca and Mg concentrations, essential minerals for plant growth, but no change in polyamines, indicators of physiological stress under conditions of high N accumulation. These results suggest that plants adapted to low-nutrient environments do not shift their resource allocation to photosynthetic processes, even after reaching N sufficiency, but instead store the excess N in organic compounds for future use. In the long term, bog species may not be able to take advantage of elevated nutrients, resulting in them being replaced by species that are better adapted to a higher nutrient environment.

Selective uptake of major and trace elements in Erica andevalensis, an endemic species to extreme habitats in the Iberian Pyrite Belt.

To assess the ecophysiological traits and the phytoremediation potential of the endemic heather Erica andevalensis, we determined the concentrations of major and trace elements in different plant parts and in rizosphere soils from Riotinto mining district (Huelva, Spain). The results showed that E. andevalensis may grow on substrates with very high As, Cu, Fe and Pb concentrations (up to 4114, 1050, 71900 and 15614 microg/g dry weight, respectively), very low availability of macro- and micronutrients and with pH values ranging from 3.3 to 4.9. In these harsh edaphic conditions E. andevalensis selectively absorbed and translocated essential nutrients and excludes potentially phytotoxic elements, which were accumulated in the root epidermis. The concentrations of major and trace elements in E. andevalensis aerial parts from the Riotinto mining district were in the normal range for plants; likewise other Erica species it accumulated Mn and only in a very polluted site we measured leaf concentrations of As and Pb within the excessive or toxic limits for plants. Differently from previous studies, which emphasized the soil pH and bioavailability of phytotoxic elements as the main stress factors, this study showed that in the Riotinto region, E. andevalensis can tolerate wide range of pH and toxic element concentrations; the harshest environments colonized by monospecific patches of this species were characterized above all by very low availability of nutrients. The extraordinary capability to adapt to these extreme habitats made E. andevalensis a priority species to promote the phytostabilization and the development of a self-sustaining vegetative cover on Riotinto mine tailings.

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.

Effects of nursery preconditioning through mycorrhizal inoculation and drought in Arbutus unedo L. plants.

The influence of a water deficit treatment and mycorrhizal inoculation with Pisolithus tinctorius (Pers.) Coker and Couch on the water relations, gas exchange, and plant growth in Arbutus unedo L. plants was studied in order to evaluate the hardening process during the nursery period. The ability to withstand the adverse conditions after transplantation was also studied. Mycorrhizal and non-mycorrhizal seedlings of A. unedo were pot-grown for 4 months in a greenhouse (nursery period), during which time two irrigation treatments, well watered (100% water holding capacity, leaching 20% of the applied water) and deficit irrigation (50% of the well watered), were applied. Subsequently, the plants were transplanted to the field and well irrigated (transplanting period), after which and until the end of the experiment they received no water (establishment period). At the end of the nursery period, both water deficit and mycorrhizae were seen to have altered the plant morphology. Mycorrhizal plants had lower leaf area and improved leaf color parameters, while the water deficit increased root dry weight and the root/shoot ratio. Mycorrhizal plants had higher leaf water potential values than non-inoculated plants. Mycorrhizae increased stomatal conductance and photosynthesis values, especially in stressed plants. Drought led to an osmotic adjustment and a decrease in the leaf water potential values at turgor loss point in the mycorrhizal plants. Cell wall rigidity, measured as increased bulk modulus of elasticity, was decreased by the mycorrhizae effect. After transplanting, no differences were found in the water relations or gas exchange values between treatments. During the establishment period, the plants that had been exposed to both drought and mycorrhizae showed a better water status (higher leaf water and turgor potential values) and higher gas exchange values. In conclusion, water deficit and mycorrhizal inoculation of A. unedo plants in nursery produced changes in tissue water relations, gas exchange, and growth, related with the acclimation process in the seedlings, which could provide better resistance to drought and stress conditions following planting.

Variations of vessel diameter and δ13C in false rings of Arbutus unedo L. reflect different environmental conditions.

Woody species in Mediterranean ecosystems form intra-annual density fluctuations (IADFs) in tree rings in response to changes in environmental conditions, especially water availability. Dendrochronology, quantitative wood anatomy and high-resolution isotopic analysis (using a laser ablation technique) were used to characterize IADFs in Arbutus unedo shrubs grown on two sites with different water availability on the island of Elba (Italy). Our findings show that IADF characterization can provide information about the relationship between environmental factors and tree growth at the seasonal level. At the more xeric site, IADFs mainly located in the early and middle parts of the annual ring, showed a decrease in vessel size and an increase in δ(13) C as a result of drought deficit. Opposite trends were found at the more mesic site, with IADFs located at the end of the ring and associated with a lower δ(13) C. Moreover, at the first site, IADFs are induced by drought deficit, while at the second site IADFs are linked with the regrowth in the last part of the growing season triggered by favourable wet conditions. This combined approach is a promising way for dating problematic wood samples and interpreting the phenomena that trigger the formation of IADFs in the Mediterranean environment.

Pollination ecology of Disterigma stereophyllum (Ericaceae) in south-western Colombia.

Several authors have recently expressed doubts that the 'pollination syndromes' as usually expressed are an adequate description of correlated suites of floral characters, or that they adequately describe evolutionary or ecological associations of plants with pollinators. Disterigma stereophyllum is a neotropical Ericaceae with floral characteristics intermediate between the 'entomophilous' syndrome and the 'ornithophilous' syndrome: the corolla is short, white and urceolate, but flowers produce large amounts of dilute nectar. We studied the pollination ecology of this species in south-western Colombia, and found it to be pollinated almost exclusively by hummingbirds at our study site. Two hummingbird species were responsible for about 75 of visits. Despite the fact that nectar standing crop remained more or less constant throughout the day, visit frequencies were highest in the morning and declined throughout the day. Pollinator efficiency, measured as the number of pollen grains deposited on a virgin stigma by each visitor after one visit, did not differ among the species of hummingbirds, but was lower for a nectar-robbing bird, Diglossa albilatera. This species does not contact the surface of the stigma during nectar robbing, but can produce some self-pollination indirectly because it shakes branches vigorously while piercing the flower. These findings indicate a need for further studies of neotropical Ericaceae in order to elucidate whether floral visitors of species like D. stereophyllum fluctuate through time or space, and whether floral characteristics reflect a compromise between such different visitors, or a transitional stage between pollination syndromes, or some other possibility.

A hybrid zone dominated by fertile F1s of two alpine shrub species, Phyllodoce caerulea and Phyllodoce aleutica, along a snowmelt gradient.

In alpine ecosystems, the steep environmental gradients produced by the difference in snowmelt timing create a dynamic selective regime for alpine plants. As these gradients directly alter flowering phenology, they can affect pollen-mediated gene flow among populations of single and related species. In northern Japan, we found a hybrid zone dominated by fertile F(1)s of two alpine shrub species, Phyllodoce caerulea and P. aleutica, along a snowmelt gradient. Seed germination confirmed the fertility of F(1) hybrid, making the rarity and absence of backcross and F(2) plants puzzling. The long-term clonal perpetuation of F(1) hybrids (at least a few thousand years ago) contributes the maintenance of this unique hybrid zone. The distribution patterns of chloroplast DNA haplotypes suggest that F(1) formation might be caused by directional pollen flow between parental species along the snowmelt gradient. Based on these results, we discuss the ecological and evolutionary significance of this unique hybrid zone.

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.

Dormancy and the fire-centric focus: germination of three Leucopogon species (Ericaceae) from South-eastern Australia.

BACKGROUND AND AIMS: Germination studies of species from fire-prone habitats are often focused on the role that fire plays in breaking dormancy. However, for some plant groups in these habitats, such as the genus Leucopogon (Ericaceae), dormancy of fresh seeds is not broken by fire cues. In the field, these same species display a flush of seedling emergence post-fire. Dormancy and germination mechanisms therefore appear complex and mostly unknown. This study aimed to identify these mechanisms by establishing dormancy class and testing the effects of a set of typical germination cues, including those directly related to fire and entirely independent of fire. METHODS: To classify dormancy, we assessed seed permeability and embryo morphology, and conducted germination experiments at seasonal temperatures in incubators. To test the effects of fire cues on germination, factorial combinations of smoke, heat and dark treatments were applied. Ageing treatments, using burial and seasonal incubation, were also tested. Germination phenology was established. KEY RESULTS: Seeds were dormant at release and had underdeveloped embryos. Primary dormancy of the study species was classified as morphophysiological. Seasonal temperature changes overcame primary dormancy and controlled timing of germination. Fire cues did not break primary dormancy, but there was a trend for smoke to enhance germination once this dormancy was overcome. CONCLUSIONS: Despite the fact that fire is a predominant disturbance and that many species display a flush of emergence post-fire, seasonal temperatures broke the primary physiological dormancy of the study species. It is important to distinguish between fire being responsible for breaking dormancy and solely having a role in enhancing levels of post-fire germination for seeds in which dormancy has been overcome by other factors. Biogeographical evidence suggests that morphological and physiological factors, and therefore seasonal temperatures, are likely to be important in controlling the dormancy and patterns of post-fire germination of many species in fire-prone regions.