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Representative members of Empetreae (Ericoideae). 54. Ceratiola ericoides, close-up of male (left) and female (right) flowers (image by S. Zona). 55. Ceratiola ericoides, in fruit (image by A.A.A.). 56. Empetrum hermaphroditum, close-up of fruit (image by A.A.A.).

Representative members of Empetreae (Ericoideae). 54. Ceratiola ericoides, close-up of male (left) and female (right) flowers (image by S. Zona). 55. Ceratiola ericoides, in fruit (image by A.A.A.). 56. Empetrum hermaphroditum, close-up of fruit (image by A.A.A.).

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A new classification of Ericaceae is presented based on phylogenetic analyses of nuclear and chloroplast DNA sequence data, morphology, anatomy, and embryology. Eight subfamilies and 20 tribes are recognized. In this classification Epacridaceae are included as Styphelioideae and Empetraceae as tribe Empetreae within the Ericoideae. The herbaceous t...

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... sensu stricto (tribe Pyroleae, family Ericaceae). This tribe is related to the fully mycoheterotrophic tribes Monotropeae and Pterosporeae but contains plant species with varying levels of mycoheterotrophy (Hynson et al., 2009;Kron et al., 2002;Lallemand et al., 2019). P. japonica and P. subaphylla are suitable taxa to investigate the changes associated with mycoheterotrophic evolution because they differ significantly in their levels of mycoheterotrophy and because recent systematic research suggests that they are sister species (e.g. ...
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Land plants are typically photosynthetic, but some species have lost the ability to photosynthesize, instead relying on mycorrhizal fungi to obtain carbon. Increasing levels of partial mycoheterotrophy, in which seemingly autotrophic plants receive fungal carbon from their fungal partners, and reduced sprouting in concert with greater reproduction when sprouting, may be intermediate steps in the evolution of this trait. We studied the microevolutionary demography of Pyrola japonica and the closely related species P. subaphylla, which are currently considered sister species. While these species are both partially mycoheterotrophic, P. subaphylla is more strongly so, indicating that it may be closer to evolving full mycoheterotrophy. We tracked individuals in two nearby populations in Fukushima, Japan from 2015 to 2020. We analysed vital rates, population trends and long‐run population structure with historical function‐based matrix projection models. We assessed how shifts in P. subaphylla demography relate to fitness using a stochastic life table response experiment (SLTRE). P. subaphylla exhibited strong costs of growth to survival and of reproduction to sprouting, size and fecundity. It sprouted more than P. japonica but to smaller stages. P. subaphylla's flowering frequency was approximately four times that of P. japonica, but after flowering it was smaller and more likely to die. These life history costs appeared to drive differences in fitness, measured as stochastic population growth rate. Relative to P. japonica, shifts in fitness in P. subaphylla were due more to shifts in means than standard deviations of matrix elements. Shifts in growth transitions had the strongest negative impacts on fitness but also had the strongest positive impacts. Increased shifts to small non‐flowering stages were associated with increased fitness, while transitions involving dormancy were associated more strongly with drops in fitness. Synthesis. As far as we are aware, we found the first evidence that costs of growth and reproduction drive the evolution of increased sprouting and smaller size as a species becomes more intensely mycoheterotrophic. Surprisingly, vegetative dormancy did not evolve with mycoheterotrophy. We suggest further studies assessing whether these seemingly maladaptive trends result from genetic drift, genetic linkage or another mechanism.
... The Ericaceae family is highly diverse in terms of its mycorrhizal associations (Lallemand et al. 2016): while the earliest-diverging genus Enkianthus retained the ancestral AM (Obase et al. 2013), the subfamilies Monotropoideae and Arbutoideae form ectendomycorrhizas (a type of EM with intracellular colonization), and the subfamilies Ericoideae, Cassiopoideae, Harrimanelloideae, Styphelioideae, and Vaccinioideae, a monophyletic clade known as the "early anther inversion clade" (EAI clade; Kron et al. 2002), form ErM (Selosse et al. 2007). ...
... 4500 species globally ranging from sea level to high mountains over 5000 m (POWO 2023). The most recent classification system of Ericaceae comprises eight sub-families, and 20 tribes with 121 genera (POWO 2023;Kron et al. 2002). The family is distributed worldwide, especially in tropical and temperate regions, except for Antarctica (Kron and Luteyn 2005). ...
... The areas of higher species richness are localized in the montane tropics of South America, Asia, New Guinea, Australia, and Africa (Kron and Luteyn 2005). The members of this family grow in a wide range of habitats and with diverse life history strategies ranging from shrubs and trees to various often ignored kinds of epiphytes: climbing epiphyte, climbing shrub, epiphytic shrub, scrambling epiphyte, scrambling shrub, and true lianas (POWO 2023;Kron et al. 2002). It also includes achlorophyllous, mycoheterotrophic species relying on their mycorrhizal fungi for carbon nutrition (Merckx 2013). ...
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Mycorrhizal symbioses (mycorrhizas) of Ericaceae, including ericoid mycorrhiza (ErM), have been mainly studied in the Northern Hemisphere, although the highest diversity of ericaceous plants is located in the Southern Hemisphere, where several regions remain largely unexplored. One of them is South America, which harbors a remarkably high diversity of Ericaceae (691 species and 33 genera) in a wide range of environmental conditions, and a specific mycorrhizal type called cavendishioid. In this review, we compile all available information on mycorrhizas of Ericaceae in South America. We report data on the mycorrhizal type and fungal diversity in 17 and 11 ericaceous genera, respectively. We show that South American Ericaceae exhibit a high diversity of habitats and life forms and that some species from typical ErM subfamilies may also host arbuscular mycorrhiza. Also, a possible geographical pattern in South American ErM fungal communities is suggested, with Sebacinales being the dominant mycorrhizal partners of the Andean clade species from tropical mountains, while archetypal ErM fungi are common partners in southern South America species. The gathered information challenges some common assumptions about ErM and suggests that focusing on understudied regions would improve our understanding of the evolution of mycorrhizal associations in this intriguing family.
... The genera Arbutus and Arctostaphylos are closely related and grouped in the subfamily Arbutoideae (Kron et al. 2002), and many insect species share these genera as hosts. This is also the case for several leafminers: Coptodisca arbutiella, Marmara arbutiella, Coleophora glaucella Walsingham, 1882, Epinotia nigralbana (Walsingham, 1879), Epinotia terracoctana (Walsingham, 1879), and an undescribed Stigmella species (Eiseman 2022). ...
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Etainia thoraceleuca van Nieukerken, Epstein & Davis, sp. nov. is the second native American species of Etainia Beirne, 1945, and the second known Etainia species feeding on Ericaceae. The species is known from light-collected adults in the USA (California, Arizona) and Canada (Ontario). These were linked via DNA barcodes to larvae that make short leafmines on Arbutus and Arctostaphylos species, then continue feeding in stems and branches, causing damage in nurseries and planted trees in Sonoma and Marin Counties, California. The holotype was accidentally reared from Arbutus arizonica , without observing the damage. Life history and damage are described in detail. Damage in Arctostaphylos uva-ursi found in Washington State probably belongs to E. thoraceleuca , which is a sister species to the European E. albibimaculella (Larsen, 1927).
... To generate quantitative climate estimates, we mapped observed fossil pollen types onto botanical taxa, yielding lists of species and/or genera that could plausibly have been the source of each fossil pollen type (as detailed in the Supplementary Information, Supplementary Table 4 and using refs. [80][81][82][83][84][85][86][87][88]. We estimated the climate tolerances of these potential source species from species occurrence data from the Atlas of Living Australia (https://www.ala.org.au/) ...
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Late Pleistocene ice-age climates are routinely characterized as having imposed moisture stress on low- to mid-latitude ecosystems1–5. This idea is largely based on fossil pollen evidence for widespread, low-biomass glacial vegetation, interpreted as indicating climatic dryness⁶. However, woody plant growth is inhibited under low atmospheric CO2 (refs. 7,8), so understanding glacial environments requires the development of new palaeoclimate indicators that are independent of vegetation⁹. Here we show that, contrary to expectations, during the past 350 kyr, peaks in southern Australian climatic moisture availability were largely confined to glacial periods, including the Last Glacial Maximum, whereas warm interglacials were relatively dry. By measuring the timing of speleothem growth in the Southern Hemisphere subtropics, which today has a predominantly negative annual moisture balance, we developed a record of climatic moisture availability that is independent of vegetation and extends through multiple glacial–interglacial cycles. Our results demonstrate that a cool-moist response is consistent across the austral subtropics and, in part, may result from reduced evaporation under cool glacial temperatures. Insofar as cold glacial environments in the Southern Hemisphere subtropics have been portrayed as uniformly arid3,10,11, our findings suggest that their characterization as evolutionary or physiological obstacles to movement and expansion of animal, plant and, potentially, human populations¹⁰ should be reconsidered.
... O gênero Agarista inclui 33 espécies (Sampaio et al. 2023) e está inserido na tribo Lyonieae, juntamente com Lyonia Nutt., Pieris D.Don, Craibiodendron W.W.Sm. A tribo possui como sinapomorfias a presença de tecido fibroso no mesofilo, filetes geniculados e células epidérmicas lignificadas (Kron et al. 2002). O gênero é dividido em duas seções morfologicamente distintas e com distribuições disjuntas (Stevens 1971;Judd 1984Judd , 1995. ...
Article
É apresentado aqui a Flora de Agarista (Ericaceae) para o estado da Bahia, Brasil. São reconhecidas dez espécies, sendo duas novas ocorrências para a Bahia, A. glaberrima e A. virgata, e uma espécie endêmica de Morro do Chapéu, A. revolutissima. O presente trabalho inclui chave de identificação para os gêneros de Ericaceae representados na Bahia e espécies de Agarista, descrições, comentários, imagens das plantas e mapas de distribuição das espécies no estado.
... The combined analysis of both morphological and molecular data has recently shown that there are 20 tribes belonging to eight subfamilies of Ericaceae s.l.: Enkianthoideae, Monotropoideae, Arbutoideae, Cassiopoideae, Ericoideae, Harrimanelloideae, Styphelioideae, Vaccinioideae (Kron et al. 2002). Later it was revealed that the subfamily Monotropoideae sensu Kron et al. (2002) is paraphyletic and the recognition of one more subfamily of Ericaceae s.l., Pyroloidaeae, was proposed (Liu et al. 2014, Schwery et al. 2014, Freudenstein et al. 2016. ...
... The combined analysis of both morphological and molecular data has recently shown that there are 20 tribes belonging to eight subfamilies of Ericaceae s.l.: Enkianthoideae, Monotropoideae, Arbutoideae, Cassiopoideae, Ericoideae, Harrimanelloideae, Styphelioideae, Vaccinioideae (Kron et al. 2002). Later it was revealed that the subfamily Monotropoideae sensu Kron et al. (2002) is paraphyletic and the recognition of one more subfamily of Ericaceae s.l., Pyroloidaeae, was proposed (Liu et al. 2014, Schwery et al. 2014, Freudenstein et al. 2016. ...
... The fruits of Ericaceae are coenocarpous with up to 12 locules and with different types of placentation: usually axile to intruded parietal and, rarely, apical or basal. There are one to hundreds of ovules per locule (Kron et al. 2002, Stevens et al. 2004. ...
Article
According to modern phylogenetic data, the monotypic subfamily Enkianthoideae is sister to all other subfamilies of the family Ericaceae s.l. Aiming to determine the peculiarities of development and structure of Enkianthus pericarp and to reveal the principal characters of its structure, which can be original for the Ericaceae, we have studied fruit anatomical structure at different stages of development in four species of the genus Enkianthus belonging to three sections: Meisteria, Enkiantella, Enkianthus. The pericarp of superior loculicidal capsule of Enkianthus is differentiated into three histogenetic zones: (i) one-layered exocarp, (ii) multilayered mesocarp differentiated into two histological zones: the peripheral parenchymatous one, and the inner sclerenchymatous one, and (iii) one-layered endocarp composed of tangentially elongated sclereids. It has been shown that the lignification of cells of the endocarp proceeds after the lignification of the inner zone of the mesocarp. Our investigation demonstrates that the superior loculicidal capsules of Enkianthus are capsules of Hamamelis type, which are recognized as the probable ancestral (plesiomorphic) fruit type for other subfamilies of Ericaceae. This fruit type is recognized as the original in several modes of morphogenetic transformations of fruit in Ericaceae family, from which other types of capsules, pyrenariums, and berries originated.
... The most important Vaccinium crop species are found in the sections Cyanococcus (blueberries), Oxycoccus (cranberry), Vitis-Idaea (lingonberry), and Myrtillus (bilberry) 5 . However, species and section delimitations have been extensively discussed in the literature, as they do not form monophyletic groups 6,7 . The taxonomic classification has been difficult to resolve because of considerable phenotypic variability with overlapping morphologies, complex ploidy series (ranging from diploids to hexaploids), and general lack of crossing barriers leading to numerous hybridization events 5 . ...
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Vaccinium is a large genus of shrubs that includes a handful of economically important berry crops. Given the numerous hybridizations and polyploidization events, the taxonomy of this genus has remained the subject of long debate. In addition, berries and berry-based products are liable to adulteration, either fraudulent or unintentional due to misidentification of species. The availability of more genomic information could help achieve higher phylogenetic resolution for the genus, provide molecular markers for berry crops identification, and a framework for efficient genetic engineering of chloroplasts. Therefore, in this study we assembled five Vaccinium chloroplast sequences representing the economically relevant berry types: northern highbush blueberry (V. corymbosum), southern highbush blueberry (V. corymbosum hybrids), rabbiteye blueberry (V. virgatum), lowbush blueberry (V. angustifolium), and bilberry (V. myrtillus). Comparative analyses showed that the Vaccinium chloroplast genomes exhibited an overall highly conserved synteny and sequence identity among them. Polymorphic regions included the expansion/contraction of inverted repeats, gene copy number variation, simple sequence repeats, indels, and single nucleotide polymorphisms. Based on their in silico discrimination power, we suggested variants that could be developed into molecular markers for berry crops identification. Phylogenetic analysis revealed multiple origins of highbush blueberry plastomes, likely due to the hybridization events that occurred during northern and southern highbush blueberry domestication.
... The subfamily Monotropoideae (Ericaceae), is distributed throughout the Northern Hemisphere (Bidartondo and Bruns 2001;Kron et al. 2002). It is characterized by its achlorophyllous and fully mycoheterotrophic habit, with scale-like leaves, bisexual and actinomorphic flowers, free sepals and petals, and a superior 1-6 locule ovary (Wallace 1987;Wu et al. 2016). ...
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Due to their reduced morphology, non-photosynthetic plants have been one of the most challenging groups to delimit to species level. The mycoheterotrophic genus Monotropastrum , with the monotypic species M. humile , has been a particularly taxonomically challenging group, owing to its highly reduced vegetative and root morphology. Using integrative species delimitation, we have focused on Japanese Monotropastrum , with a special focus on an unknown taxon with rosy pink petals and sepals. We investigated its flowering phenology, morphology, molecular identity, and associated fungi. Detailed morphological investigation has indicated that it can be distinguished from M. humile by its rosy pink tepals and sepals that are generally more numerous, elliptic, and constantly appressed to the petals throughout its flowering period, and by its obscure root balls that are unified with the surrounding soil, with root tips that hardly protrude. Based on genome-wide single-nucleotide polymorphisms, molecular data has provided clear genetic differentiation between this unknown taxon and M. humile . Monotropastrum humile and this taxon are associated with different Russula lineages, even when they are sympatric. Based on this multifaceted evidence, we describe this unknown taxon as the new species M. kirishimense . Assortative mating resulting from phenological differences has likely contributed to the persistent sympatry between these two species, with distinct mycorrhizal specificity.
... The morphological characters available suggest affinities with Ericaceae subfam. Vaccinioideae (one of the superior-ovaried groups; see Kron et al. 2002). Similar fruits of Ericaceae are well documented by anatomically preserved fossils in the middle Miocene Fasterholt flora of Denmark, including those assigned to Lyonia, Eubotrys and Zenobia, by Friis (1985). ...
Article
Selected fruits and seeds preserved as molds and casts in sediments from the Anjou ora of Maine-et-Loire have been reexamined with the aid of X-ray tomography. Virtual casts and surface renderings from micro-CT scanning data reveal external and internal morphological characters that were not visible by standard re ected light microscopy. Application of this methodology leads to a revision of the fruit formerly treated as Juglandicarya. It is a 5-valved capsule of likely sapindalean a nity, and is placed in Vaudoisia gruetii (Vaudois-Miéja) gen. et. comb. nov. A seed with the characteristic rumination of Anonaspermum is also recognized for the rst time, as are capsular fruits of Ericaceae. We note that many disseminules are hidden from optical viewing because they are buried within the hand samples. A more extensive micro-CT scan investigation of more samples, including those showing only fossil leaf impressions at the surface, may be expected to yield a wealth of new information on this classic ora.
... 1). This result is supported by the phylogenetic affinity of Erica and Rhododendron, which both pertain to subfamily Ericoideae (Kron et al. 2002). ...
... Vaccinioideae), and Gaultheria shallon (subfam. Vaccinioideae, tribe Gaulterieae) clade (Kron et al. 2002) all lie within the E. elevata/vaccinii complex (FIG. 1). ...
Article
The genus Rhododendron comprises over 1000 evergreen and deciduous species. In the Pacific Northwest Coast region of North America (PNWC), powdery mildews infecting deciduous Rhododendron spp. are well documented but less so on evergreen Rhododendron spp. Infections of both groups of hosts historically have been attributed to Erysiphe azaleae or E. vaccinii. No formal characterizations of powdery mildew fungi infecting either deciduous or evergreen Rhododendron spp. in the PNWC have been completed. The objectives of this study were to identify the powdery mildew pathogens infecting evergreen Rhododendron spp. in the PNWC and to assess the phylo-genetic position of these fungi within the Erysiphaceae. To ascertain valid taxonomic conclusions, and to determine whether potential introductions of exotic Rhododendron powdery mildews in North America have occurred, it was necessary to put the new North American phylogenetic data into a worldwide context. Therefore, available phylogenetic data from all Erysiphe spp. on Rhododendron have been included in our analyses. Based on analyses of numerous new internal transcribed spacer (ITS) and 28S rDNA sequences and already available sequences deposited in GenBank retrieved from evergreen and deciduous Rhododendron spp., the following Erysiphe spp. could be phylogenetically confirmed (all belonging to Erysiphe sect. Microsphaera): Erysiphe azaleae nom. cons. (Oidium ericinum could be verified as a synonym), E. digitata (holotype sequenced), E. izuensis, and E. vaccinii. Erysiphe azaleae and E. vaccinii are epitypified with sequenced specimens, and an ex-neotype sequence has been obtained for Oidium ericinum. Erysiphe rhododendri (Erysiphe sect. Erysiphe), only known from two collections in India (Himalayan region), was not available for phylogentic analyses. ARTICLE HISTORY