To explore strange new worlds - The diversification in Tremella caloplacae was linked to the adaptive radiation of the Teloschistaceae.

Lichenicolous fungi are a heterogeneous group of organisms that grow exclusively on lichens, forming obligate associations with them. It has often been assumed that cospeciation has occurred between lichens and lichenicolous fungi, but this has been seldom analysed from a macroevolutionary perspective. Many lichenicolous species are rare or are rarely observed, which results in frequent and large gaps in the knowledge of the diversity of many groups. This, in turn, hampers evolutionary studies that necessarily are based on a reasonable knowledge of this diversity. Tremella caloplacae is a heterobasidiomycete growing on various hosts from the lichen-forming family Teloschistaceae, and evidence suggests that it may represent a species complex. We combine an exhaustive sampling with molecular and ecological data to study species delimitation, cophylogenetic events and temporal concordance of this association. Tremella caloplacae is here shown to include at least six distinct host-specific lineages (=putative species). Host switch is the dominant and most plausible event influencing diversification and explaining the coupled evolutionary history in this system, although cospeciation cannot be discarded. Speciation in T. caloplacae would therefore have occurred coinciding with the rapid diversification - by an adaptive radiation starting in the late Cretaceous - of their hosts. New species in T. caloplacae would have developed as a result of specialization on diversifying lichen hosts that suddenly offered abundant new ecological niches to explore or adapt to.

Digging Up the Roots: Taxonomic and Phylogenetic Disentanglements in Corticiaceae s.s. (Corticiales, Basidiomycota) and Evolution of Nutritional Modes.

Corticiaceae is one of the traditional families of the Agaricomycetes and served for a long time as a convenient placement for basidiomycetes with a resupinate, corticioid form of fruiting body. Molecular studies have helped to assign many corticioid fungi to diverse families and orders; however, Corticiaceae still lacks a phylogenetic characterization and modern circumscription. Here, we provide the first comprehensive phylogenetic and taxonomic revision of the family Corticiaceae based on extensive type studies and sequences of nLSU, ITS, IGS, nSSU, and mtSSU regions. Our analyses support the recognition of ten monophyletic genera in the Corticiaceae, and show that nutritional mode is not a robust basis for generic delimitations in the family. The genus Mycobernardia and the species Corticium thailandicum, Erythricium vernum, and Marchandiomyces allantosporus are described as new to science, and five new combinations are proposed. Moreover, ancestral character state reconstruction revealed that saprotrophy is the plesiomorphic nutritional mode in the Corticiaceae, while several transitions have occurred to diverse nutritional modes in this family. Identification keys are provided to the genera in Corticiaceae s.s. as well as to the species in Corticium, Erythricium, Laetisaria, and Marchandiomyces.

Lichenicolous species of Hainesia belong to Phacidiales (Leotiomycetes) and are included in an extended concept of Epithamnolia.

The lichenicolous taxa currently included in the genus Hainesia were studied based on the nuclear rDNA (18S, 28S, and internal transcribed spacer [ITS]) genes. The authors found that lichenicolous taxa form a distinct lineage sister to Epiglia gloeocapsae (Phacidiales, Leotiomycetes), only distantly related to the type species of Hainesia (Chaetomellaceae, Helotiales). Owing to morphological similarities, the authors include the lichenicolous species into the previously monotypic genus Epithamnolia. A new species, Epithamnolia rangiferinae, is described, several names are reduced into synonymy, and a key to the species of Epithamnolia is provided. The incorporation of public environmental ITS sequences showed that the closest relatives of these lichenicolous taxa are various endophytic, endolichenic, and soil-inhabiting fungi.

Contribution to the phylogeny and taxonomy of the genus Taeniolella, with a focus on lichenicolous taxa.

Taeniolella is a genus of asexual ascomycetes with saprophytic, endophytic, and lichenicolous life styles. A phylogeny of representative species is presented, with a focus on lichenicolous taxa. We obtained mtSSU and nuLSU sequence data from culture isolates of Taeniolella and from freshly collected specimens of other taxa. Taeniolella is recovered as strongly polyphyletic. The type species, Taeniolella exilis, is placed in Kirschsteiniotheliaceae within Dothideomycetes. Other saprophytic/endophytic Taeniolella species previously assigned to Sordariomycetes based on sequences were found to represent either contaminants or species that cannot be assigned to Taeniolella for morphological reasons. Lichenicolous species are restricted to Asterotexiales (Dothideomycetes) where the sequenced species of Taeniolella do not form a monophyletic group, but are related to species of Buelliella s. lat., Karschia, Labrocarpon, Melaspilea s. lat., and Stictographa. Molecular data are, however, not sufficient to reallocate the lichenicolous Taeniolella species to other genera so far. Anamorph-teleomorph relationships between these taxa and lichenicolous Taeniolella are discussed but could not be demonstrated. The type of Buelliella is placed in Asterotexiales, and the genus recovered as polyphyletic. Three new lichenicolous Taeniolella species are described, Taeniolella hawksworthiana, Taeniolella pyrenulae and Taeniolella toruloides. Taeniolella rudis is transferred to Sterigmatobotrys.

Cyphobasidium gen. nov., a new lichen-inhabiting lineage in the Cystobasidiomycetes (Pucciniomycotina, Basidiomycota, Fungi).

Pucciniomycotina is a highly diverse group of fungi, showing a remarkably wide range of lifestyles and ecologies. However, lichen-inhabiting fungi are only represented by a few species included in the genera Chionosphaera and Cystobasidium, and their phylogenetic position has never been investigated. Phylogenetic analyses using the nuclear SSU, ITS, and LSU ribosomal DNA markers reveal that the lichenicolous members of Cystobasidium (C. hypogymniicola, C. usneicola) form a monophyletic group distinct from Cystobasidium and outside the Cystobasidiales. The new genus Cyphobasidium is consequently described to accommodate these lichen-inhabiting species. Cyphobasidium is characterized by producing conspicuous galls on the host lichen thalli, by having distinctive basidia that arise from a thick-walled, cup-like structure, the probasidium, that persists after the senescence of the actual basidium (meiosporangium), and by its lichenicolous occurrence on species of Hypogymnia and Usnea. Cyphobasidium is one of the few representatives of the Cystobasidiomycetes in which the sexual stage predominates in nature, whereas most species in the group are known only from an asexual yeast phase. This is the first time the position of lichen-inhabiting taxa within the Pucciniomycotina is investigated using molecular data.

The phylogenetic analysis of fungi associated with lichenized ascomycete genus Bryoria reveals new lineages in the Tremellales including a new species Tremella huuskonenii hyperparasitic on Phacopsis huuskonenii.

The basidiomycete order Tremellales includes many species parasitic on or cohabiting with lichen-forming fungi. In this study, we examined the phylogenetic position of Tremellales obtained from Bryoria thalli using nSSU, 5.8S, and partial nLSU sequence data. Both Bayesian and maximum likelihood analyses revealed the presence of basidiomycetous fungi in three separate clades within Tremellales. Tremellales sp. A and Tremella sp. B exist asymptomatically in Bryoria thalli and should thus be regarded as endolichenic rather than lichenicolous fungi. The third lineage represents a new species and is described here as Tremella huuskonenii. It is hyperparasitic over galls induced by Phacopsis huuskonenii, a lichenicolous fungus growing in Bryoria thalli. We also examined the genetic diversity of Tremella sp. B and Tremella huuskonenii with an extended taxon sampling using ITS and partial nLSU sequence data. For comparison, ITS, GAPDH, and Mcm7 regions were used for phylogenetic analyses of the host lichen specimens. According to our results, phylogenetic structure within the two Tremella species does not appear to correlate with the geographic distribution nor with the phylogeny or the secondary chemistry of the host lichen. However, ITS haplotype analysis of T. huuskonenii revealed some genetic differences between European and North American populations as some haplotypes were more common in Europe than in North America and vice versa.

Host switching promotes diversity in host-specialized mycoparasitic fungi: uncoupled evolution in the Biatoropsis-usnea system.

Fungal mycoparasitism-fungi parasitizing other fungi-is a common lifestyle in some basal lineages of the basidiomycetes, particularly within the Tremellales. Relatively nonaggressive mycoparasitic fungi of this group are in general highly host specific, suggesting cospeciation as a plausible speciation mode in these associations. Species delimitation in the Tremellales is often challenging because morphological characters are scant. Host specificity is therefore a great aid to discriminate between species but appropriate species delimitation methods that account for actual diversity are needed to identify both specialist and generalist taxa and avoid inflating or underestimating diversity. We use the Biatoropsis-Usnea system to study factors inducing parasite diversification. We employ morphological, ecological, and molecular data-methods including genealogical concordance phylogenetic species recognition (GCPSR) and the general mixed Yule-coalescent (GMYC) model-to assess the diversity of fungi currently assigned to Biatoropsis usnearum. The degree of cospeciation in this association is assessed with two cophylogeny analysis tools (ParaFit and Jane 4.0). Biatoropsis constitutes a species complex formed by at least seven different independent lineages and host switching is a prominent force driving speciation, particularly in host specialists. Combining ITS and nLSU is recommended as barcode system in tremellalean fungi.

Naming and outline of Dothideomycetes-2014 including proposals for the protection or suppression of generic names.

Article 59.1, of the International Code of Nomenclature for Algae, Fungi, and Plants (ICN; Melbourne Code), which addresses the nomenclature of pleomorphic fungi, became effective from 30 July 2011. Since that date, each fungal species can have one nomenclaturally correct name in a particular classification. All other previously used names for this species will be considered as synonyms. The older generic epithet takes priority over the younger name. Any widely used younger names proposed for use, must comply with Art. 57.2 and their usage should be approved by the Nomenclature Committee for Fungi (NCF). In this paper, we list all genera currently accepted by us in Dothideomycetes (belonging to 23 orders and 110 families), including pleomorphic and nonpleomorphic genera. In the case of pleomorphic genera, we follow the rulings of the current ICN and propose single generic names for future usage. The taxonomic placements of 1261 genera are listed as an outline. Protected names and suppressed names for 34 pleomorphic genera are listed separately. Notes and justifications are provided for possible proposed names after the list of genera. Notes are also provided on recent advances in our understanding of asexual and sexual morph linkages in Dothideomycetes. A phylogenetic tree based on four gene analyses supported 23 orders and 75 families, while 35 families still lack molecular data.

Phylogeny and character evolution in the jelly fungi (Tremellomycetes, Basidiomycota, Fungi).

The Tremellomycetes (Agaricomycotina, Basidiomycota, Fungi) are a nutritionally heterogeneous group comprising saprotrophs, animal parasites, and fungicolous species (fungal-inhabiting, including lichen-inhabiting). The relationships of many species, particularly those with a lichenicolous habit, have never been investigated by molecular methods. We present a phylogeny of the Tremellomycetes based on three nuclear DNA ribosomal markers (nSSU, 5.8S and nLSU), representing all main taxonomic groups and life forms, including lichenicolous taxa. The Cystofilobasidiales, Filobasidiales, Holtermanniales, and Tremellales (including the Trichosporonales) are recovered as monophyletic, but this is not the case for the Tremellomycetes. We suggest, however, that the Cystofilobasidiales tentatively continue to be included in the Tremellomycetes. As currently circumscribed, the Filobasidiaceae, Sirobasidiaceae, Syzygosporaceae and Tremellaceae are non-monophyletic. Cuniculitremaceae, Sirobasidiaceae and Tetragoniomycetaceae are nested within Tremellaceae. The lichenicolous species currently included within the Tremellomycetes belong in this group, distributed across the Filobasidiales and Tremellales. Lichen-inhabiting taxa do not form a monophyletic group; they are distributed in several clades and sometimes intermixed with taxa of other nutritional habits. Character state reconstruction indicates that two morphological traits claimed to characterize groups in the Tremellomycetes (the basidium habit and basidium septation) are highly homoplastic. Comparative phylogenetic methods suggest that the transitions between single and catenulate basidia in the Tremellales are consistent with a punctuational model of evolution whereas basidium septation is likely to have evolved under a graduational model in the clade comprising the Holtermanniales, Filobasidiales, and Tremellales.

The obligately lichenicolous genus Lichenoconium represents a novel lineage in the Dothideomycetes.

Lichenicolous fungi are obligately lichen-associated organisms that have evolved many times throughout the Ascomycota and Basidiomycota. Approximately 20% of lichenicolous ascomycetes are recognized only from asexual (anamorphic) characteristics, so the phylogenetic position of many groups has never been resolved. Here we present the first molecular phylogeny of Lichenoconium, a genus of strictly asexual, obligately lichenicolous species with broad geographic distributions and diverse host ecologies. We obtained nuclear and mitochondrial rDNA sequences from fungal cultures isolated from four species in the genus, including a new species, Lichenoconium aeruginosum sp. nov., collected in France, Luxembourg and Netherlands. Our multilocus phylogeny supports the monophyly of fungi in the genus Lichenoconium, and places the genus in the Dothideomycetes, an ascomycete class made up mainly of saprobes and plant-associated endophytes and pathogens. There are only a few recognized groups of lichen-formers in the Dothideomycetes, but Lichenoconium is not supported as being closely related to any of these, nor to any other recognized order within the Dothideomycetes. Given that Lichenoconium is but one of over 100 genera of anamorphic lichenicolous fungi, most of which have never been studied phylogenetically, we suggest that asexual lichenicolous fungi may represent novel and evolutionarily significant phylogenetic groups in the Kingdom Fungi.

A new lichenicolous teleomorph is related to plant pathogens in Laetisaria and Limonomyces (Basidiomycota, Corticiales).

Molecular and morphological data were used to assess the taxonomic placement of an undescribed lichenicolous basidiomycete teleomorph collected in Luxembourg, Belgium and Germany. The new species is ecologically and morphologically similar to Marchandiobasidium aurantiacum, teleomorph of the common bulbilliferous lichen pathogen Marchandiomyces aurantiacus. However phylogenetic analysis of nuclear and mitochondrial rDNA sequences indicated a close relationship of the new species-not to M. aurantiacum but to turf grass pathogens in genera Laetisaria and Limonomyces, including the generic type of Laetisaria. We are including the new species in Laetisaria based on strong statistical support for the clade containing these teleomorphs and several Marchandiomyces anamorphs. The morphological and ecological diversity of this clade indicates a potentially significant evolutionary role played by lichen-associated species in the Corticiales.

Towards a new classification of the Arthoniales (Ascomycota) based on a three-gene phylogeny focussing on the genus Opegrapha.

A multi-locus phylogenetic study of the order Arthoniales is presented here using the nuclear ribosomal large subunit (nuLSU), the second largest subunit of RNA polymerase II (RPB2) and the mitochondrial ribosomal small subunit (mtSSU). These genes were sequenced from 43 specimens or culture isolates representing 33 species from this order, 16 of which were from the second largest genus, Opegrapha. With the inclusion of sequences from GenBank, ten genera and 35 species are included in this study, representing about 18% of the genera and ca 3% of the species of this order. Our study revealed the homoplastic nature of morphological characters traditionally used to circumscribe genera within the Arthoniales, such as exciple carbonization and ascomatal structure. The genus Opegrapha appears polyphyletic, species of that genus being nested in all the major clades identified within Arthoniales. The transfer of O. atra and O. calcarea to the genus Arthonia will allow this genus and family Arthoniaceae to be recognized as monophyletic. The genus Enterographa was also found to be polyphyletic. Therefore, the following new combinations are needed: Arthonia calcarea (basionym: O. calcarea), and O. anguinella (basionym: Stigmatidium anguinellum); and the use of the names A. atra and Enterographa zonata are proposed here. The simultaneous use of a mitochondrial gene and two nuclear genes led to the detection of what seems to be a case of introgression of a mitochondrion from one species to another (mitochondrion capture; cytoplasmic gene flow) resulting from hybridization.

The Ascomycota tree of life: a phylum-wide phylogeny clarifies the origin and evolution of fundamental reproductive and ecological traits.

We present a 6-gene, 420-species maximum-likelihood phylogeny of Ascomycota, the largest phylum of Fungi. This analysis is the most taxonomically complete to date with species sampled from all 15 currently circumscribed classes. A number of superclass-level nodes that have previously evaded resolution and were unnamed in classifications of the Fungi are resolved for the first time. Based on the 6-gene phylogeny we conducted a phylogenetic informativeness analysis of all 6 genes and a series of ancestral character state reconstructions that focused on morphology of sporocarps, ascus dehiscence, and evolution of nutritional modes and ecologies. A gene-by-gene assessment of phylogenetic informativeness yielded higher levels of informativeness for protein genes (RPB1, RPB2, and TEF1) as compared with the ribosomal genes, which have been the standard bearer in fungal systematics. Our reconstruction of sporocarp characters is consistent with 2 origins for multicellular sexual reproductive structures in Ascomycota, once in the common ancestor of Pezizomycotina and once in the common ancestor of Neolectomycetes. This first report of dual origins of ascomycete sporocarps highlights the complicated nature of assessing homology of morphological traits across Fungi. Furthermore, ancestral reconstruction supports an open sporocarp with an exposed hymenium (apothecium) as the primitive morphology for Pezizomycotina with multiple derivations of the partially (perithecia) or completely enclosed (cleistothecia) sporocarps. Ascus dehiscence is most informative at the class level within Pezizomycotina with most superclass nodes reconstructed equivocally. Character-state reconstructions support a terrestrial, saprobic ecology as ancestral. In contrast to previous studies, these analyses support multiple origins of lichenization events with the loss of lichenization as less frequent and limited to terminal, closely related species.

Remarkable nutritional diversity of basidiomycetes in the Corticiales, including a new foliicolous species of Marchandiomyces (anamorphic Basidiomycota, Corticiaceae) from Australia.

Fungi in the basidiomycete order Corticiales are remarkably diverse nutritionally, including a variety of saprotrophs, plant and fungal pathogens, and lichen-forming fungi. Tracing the origin of this diversity depends on a clearer understanding of the phylogenetic relationships of fungi in the order. One of its core members is the genus Marchandiomyces, originally established for lichen pathogens that form orange or coral bulbils. We describe here a new species in the genus, M. marsonii sp. nov., which is unusual in its appearance, habit, and geographic provenance. It is foliicolous on leaves of Pandanus (screw pines, Pandanaceae) and produces flattened, coral bulbils resembling apothecia of the ascomycete genus Orbilia. It is also the first member of the genus to be collected from Australia. An isolate of the new fungus and several additional cultures of related plant pathogenic fungi were obtained and investigated phylogenetically using parsimony, likelihood, and Bayesian analyses of nuclear small and large subunit ribosomal sequences. Our phylogeny makes clear that Marchandiomyces species and their close relatives contribute significantly to the ecological diversity of the Corticiales and that this diversity is derived mainly from lignicolous ancestors.

Phylogenetic diversity of lichen-associated homobasidiomycetes.

The vast majority of lichenicolous fungi are relatively host-specific, nonvirulent ascomycetes and heterobasidiomycetes. A few known lichenicolous homobasidiomycetes (mushroom-forming fungi) generally exhibit broad host ecologies and in some cases, high virulence. Many produce conspicuous sclerotia or bulbils, thought to be adaptive in dispersal and survival. To more clearly understand the evolution of these atypical behaviors in lichenicolous basidiomycetes, we isolated or acquired specimens or cultures of 23 lichenicolous homobasidiomycetes and their relatives, from which we obtained mainly nuclear and some mitochondrial rDNA sequences. Phylogenetic analyses in this study indicate that a lichenicolous habit arose in four major clades. In two of these clades the habit represents a major evolutionary theme linked to the origin of well-known basidiolichens. The phylogenetic diversity of these fungi indicates that the lichenicolous habit arose recently and independently in the mushroom-forming fungi.

New insights into classification and evolution of the Lecanoromycetes (Pezizomycotina, Ascomycota) from phylogenetic analyses of three ribosomal RNA- and two protein-coding genes.

The Lecanoromycetes includes most of the lichen-forming fungal species (> 13500) and is therefore one of the most diverse class of all Fungi in terms of phenotypic complexity. We report phylogenetic relationships within the Lecanoromycetes resulting from Bayesian and maximum likelihood analyses with complementary posterior probabilities and bootstrap support values based on three combined multilocus datasets using a supermatrix approach. Nine of 10 orders and 43 of 64 families currently recognized in Eriksson's classification of the Lecanoromycetes (Outline of Ascomycota--2006 Myconet 12:1-82) were represented in this sampling. Our analyses strongly support the Acarosporomycetidae and Ostropomycetidae as monophyletic, whereas the delimitation of the largest subclass, the Lecanoromycetidae, remains uncertain. Independent of future delimitation of the Lecanoromycetidae, the Rhizocarpaceae and Umbilicariaceae should be elevated to the ordinal level. This study shows that recent classifications include several nonmonophyletic taxa at different ranks that need to be recircumscribed. Our phylogenies confirm that ascus morphology cannot be applied consistently to shape the classification of lichen-forming fungi. The increasing amount of missing data associated with the progressive addition of taxa resulted in some cases in the expected loss of support, but we also observed an improvement in statistical support for many internodes. We conclude that a phylogenetic synthesis for a chosen taxonomic group should include a comprehensive assessment of phylogenetic confidence based on multiple estimates using different methods and on a progressive taxon sampling with an increasing number of taxa, even if it involves an increasing amount of missing data.

Marchandiomyces lignicola sp. nov. shows recent and repeated transition between a lignicolous and a lichenicolous habit.

The anamorphic basidiomycete genus Marchandiomyces presently includes two common lichenicolous (lichen-inhabiting) species, M. corallinus and M. aurantiacus (teleomorph Marchandiobasidium aurantiacum). We describe here a new species, M. lignicola sp. nov., that is similar to M. corallinus in the colour of its sclerotia, but differs in having a wood-inhabiting (lignicolous) habit. The phylogenetic position of this lignicolous fungus was compared with the lichenicolous species of Marchandiomyces and related species currently placed in the basidiomycetous families Corticiaceae and Ceratobasidiaceae using parsimony, likelihood, and Bayesian analyses of complete sequences of the nuclear small subunit and internal transcribed spacers ribosomal DNA, and a portion of the nuclear large subunit ribosomal DNA. These DNA sequences were obtained from isolated cultures of freshly collected specimens. Significant Bayesian posterior probabilities, as well as maximum likelihood and parsimony analyses, indicate that the new lignicolous species is closely related to M. corallinus, the type species of Marchandiomyces. In most analyses these two species are monophyletic with the lichenicolous M. aurantiacus, although this relationship is not strongly supported. Since M. lignicola is more closely related to M. corallinus than to M. aurantiacus, either a transition to the lignicolous habit occurred recently within an ancestral lichenicolous group or, more likely, transition to the lichenicolous habit arose recently and in parallel from an ancestral lignicolous habit.

Assembling the fungal tree of life: progress, classification, and evolution of subcellular traits.

Based on an overview of progress in molecular systematics of the true fungi (Fungi/Eumycota) since 1990, little overlap was found among single-locus data matrices, which explains why no large-scale multilocus phylogenetic analysis had been undertaken to reveal deep relationships among fungi. As part of the project "Assembling the Fungal Tree of Life" (AFTOL), results of four Bayesian analyses are reported with complementary bootstrap assessment of phylogenetic confidence based on (1) a combined two-locus data set (nucSSU and nucLSU rDNA) with 558 species representing all traditionally recognized fungal phyla (Ascomycota, Basidiomycota, Chytridiomycota, Zygomycota) and the Glomeromycota, (2) a combined three-locus data set (nucSSU, nucLSU, and mitSSU rDNA) with 236 species, (3) a combined three-locus data set (nucSSU, nucLSU rDNA, and RPB2) with 157 species, and (4) a combined four-locus data set (nucSSU, nucLSU, mitSSU rDNA, and RPB2) with 103 species. Because of the lack of complementarity among single-locus data sets, the last three analyses included only members of the Ascomycota and Basidiomycota. The four-locus analysis resolved multiple deep relationships within the Ascomycota and Basidiomycota that were not revealed previously or that received only weak support in previous studies. The impact of this newly discovered phylogenetic structure on supraordinal classifications is discussed. Based on these results and reanalysis of subcellular data, current knowledge of the evolution of septal features of fungal hyphae is synthesized, and a preliminary reassessment of ascomal evolution is presented. Based on previously unpublished data and sequences from GenBank, this study provides a phylogenetic synthesis for the Fungi and a framework for future phylogenetic studies on fungi.

Marchandiobasidium aurantiacum gen. sp. nov., the teleomorph of Marchandiomyces aurantiacus (Basidiomycota, ceratobasidiales).

The name Marchandiobasidium aurantiacum gen. sp. nov. is introduced for the teleomorph of Marchandiomyces aurantiacus. Dolipore septa and septal pore caps of the closely related Marchandiomyces corallinus are typical of the Ceratobasidiales, and the basidiomatal characters of Marchandiobasidium aurantiacum are reminiscent of those of the monotypic genus Waitea. Morphological, ultrastructural and molecular data suggest that Marchandiobasidium should not be included in Waitea, but should be treated as a distinct genus.