Morphology and phylogenetic interrelationships of the Asteraceae,Calyceraceae, Campanulaceae,Goodeniaceae, and related families (Asterales)

In search for the sister group of the Asteraceae, morphological evidence was assembled for investigating the relationship between the Asteraceae and those families most frequently considered to be their closest relatives, in particular the Calyceraceae, Campanulaceae (along with the frequently included Lobeliaceae, Cyphiaceae, Cyphocarpaceae, and Nemacladaceae), and Goodeniaceae (and the sometimes included Brunoniaceae). Several other families that have been associated with this group of families, the “Asterales-Campanulales-complex,” were also considered: Pentaphragmataceae, Sphenocleaceae, Stylidiaceae, Donatiaceae, Menyanthaceae, and Argophyllaceae. In order to delineate the complex more precisely, another eight putatively related families were also included in the analysis. Cladistic parsimony analysis of 46 morphological and chemical characters for the 23 families was undertaken. Stability of the branches was estimated by the number of extra steps necessary to lose the group, as well as by the number of reweighted extra steps (using rescaled consistency indices) necessary to lose the group (a new approach). The results indicate that there is a monophyletic group of 14 families comprising those of the Asterales-Campanulales-complex as well as Pentaphragmataceae, Sphenocleaceae, Stylidiaceae, Donatiaceae, and Menyanthaceae; this group is recognized as the order Asterales. Within the order, the Asteraceae, Calyceraceae, Brunoniaceae, and Goodeniaceae form one comparatively well-supported clade and the five families of the Campanulaceae sensu lato form another well-supported clade.     

Review of the sporoderm ultrastructure of members of the Asterales

Palynomorphological characteristics of the order Asterales are discussed. Particular attention is paid to the pollen morphology of basal families of this group and to that of problematic taxa that are considered as sister groups to the group under study. Ultrastructurally similar sporoderms of several families, including (1) Asteraceae, Calyceraceae, and Goodeniaceae; (2) Campanulaceae, Phellinaceae, and Menyanthaceae; (3) Rousseaceae, Abrophyllaceae, and Columelliaceae, are described. Pollen grains of Alseuosmiaceae and Stylidiaceae show unique ultrastructural features of the exine.     

Evolutionary origin of the Asteraceae capitulum: Insights from Calyceraceae

? Premise of the study: Phylogenies based on molecular data are revealing that generalizations about complex morphological structures often obscure variation and developmental patterns important for understanding the evolution of forms, as is the case for inflorescence morphology within the well-supported MGCA clade (Menyanthaceae + Goodeniaceae + Calyceraceae + Asteraceae). While the basal families share a basic thyrsic/thyrsoid structure of their inflorescences, Asteraceae possesses a capitulum that is widely interpreted as a racemose, condensed inflorescence. Elucidating the poorly known inflorescence structure of Calyceraceae, sister to Asteraceae, should help clarify how the Asteraceae capitulum evolved from thyrsic/thyrsoid inflorescences. ? Methods: The early development and structure of the inflorescence of eight species (five genera) of Calyceraceae were studied by SEM, and patterns of evolutionary change were interpreted via phylogenetic character mapping. ? Key results: The basic inflorescence structure of Calyceraceae is a cephalioid (a very condensed botryoid/thyrsoid). Optimization of inflorescence characters on a DNA sequence-derived tree suggests that the Asteraceae capitulum derives from a simple cephalioid through two morphological changes: loss of the terminal flower and suppression of the cymose branching pattern in the peripheral branches. ? Conclusions: Widely understood as a condensed raceme, the Asteraceae capitulum is the evolutionary result of a very reduced, condensed thyrsoid. Starting from that point, evolution worked separately only on the racemose developmental control/pattern within Asteraceae and mainly on the cymose developmental control/pattern within Calyceraceae, producing head-like inflorescences in both groups but with very different diversification potential. We also discuss possible remnants of the ancestral cephalioid structure in some Asteraceae.     

Studies in the Goodeniaceae and the Brunoniaceae with a discussion of their relationship to Asteraceae and Calyceraceae

The relationships between Asteraceae, Calyceraceae, Goodeniaceae, and Brunoniaceae are discussed, and some SEM-observations of microcharacters within the two latter families are presented. Based on morphological evidence neither Goodeniaceae s.str. nor Goodeniaceae s.lat. (i.e., incl. Brunonia) appear to be the putative sister taxon of Asteraceae. A cladistic analysis demonstrates that Brunonia australis may constitute a separate, monotypic family or alternatively be deeply nested within Goodeniaceae s.lat Brunonia shares a number of potential synapomorphies with Asteraceae. Of special interest are the peculiar hairs and feathery pappus bristles which very much resemble those found within Asteraceae subfam. Barnadesioideae. However, studies with SEM disclose that these structures rather constitute autapomorphies for the taxa in question. In conclusion, Calyceraceae remain to be the best sister candidate of the sunflower family.     

从叶绿体DNA trnL-F序列论双参属的归属问题

双参属Triplostegia Wall．ex DC．由分布于东南亚地区的2个种组成,为多年生草本植物。它的归属一直存在争议,有时置于川续断科Dipsacaceae或败酱科Valerianaceae,有时单立一科,即双参科Triplostegiaceae。本研究对广义川续断目Dipsacales s．l．的21种植物(分别来自于败酱科、川续断科、双参属、刺参属Morina、广义忍冬科Caprifoliaceae s. l．、五福花科 Adoxaceae)和外类群人参Panax schin-seng Nees．的叶绿体 DNA trnL-F区进行了测序,并建立系统发育树状图。结果显示,败酱科、川续断科、双参属、刺参属和广义忍冬科的4个属(双盾木属Dipelta、虫胃实属Kolkwitzia、六道木属Abelia和北极花属Linnaea)形成 了一个单系群并得到了很强的支持(100％ bootstrap);双参属与川续断科有更近的关系,建议作为一个亚科置于川续断科;广义忍冬科为一多系类群;而刺参属与其他广义川续断目类群之间的关系尚不能确定。     

Phylogenetic relationships of the disputed genus Triplostegia based on trnL-F sequences

The phylogenetie relationships of Triplostegia Wall. ex DC., comprising two species of perennial herbs from southeastern Asia, have long been in dispute. This genus was placed in either Dipsacaceae or Valerianaceae or in a family of its own, Triplostegiaceae. In this paper, the chloroplast DNA (cpDNA) trn L-F regions of 21 species in the Dipsacales s. l. (including Valerianaceae, Dipsacaceae, Triplostegia, Morina, Caprifoliaceae s. l. and Adoxaceae) and an outgroup Panax schin-seng Nees. were amplified and sequenced. The phylogenetic relationships among these 22 species were constructed based on trn L-F sequences. The results demonstrated that Valerianaceae, Dipsacaceae, Triplostegia, Morina and four genera from the Caprifoliaceae s. l. form a monophyletic group with a strong support (100% bootstrap). Triplostegia, a sister group to Dipsacaceae, is close enough to be placed in the Dipsacaceae as a subfamily. The traditional Caprifoliaceae s.l. are polyphyletic, and relationships of Morina among the groups within Dipsacales s. l. are uncertain. Key words Triplostegia; Caprifoliaceae s. l.; Morina; Dipsacales s. l.; trnL-F sequences; Sys-tematic position     

Fructose oligosaccharides: Possible markers of phylogenetic relationships among dicotyledonous plant families

Stems from more than 555 species representing 440 genera of approximately 210 dicotyledonous families were surveyed for the presence of fructose oligosaccharides. The tissues from 372 species representing 312 genera and 187 families did not contain such oligosaccharides. The sugars occurred sporadically or in small amounts in members of the following families: Crossosomataceae, Lauraceae, Malpighiaceae, Cornaceae, Myrisinaceae, Melianthaceae, Greyiaceae, and Simaroubaceae. The fructosyl sucrose trisaccharide, isokestose, aswell as its fructose tetrasaccharide homolog (but not higher inulin oligosaccharides) were present in members of the families Hippocastanaceae and Limnathaceae. Stems from members of the families Clethraceae and Monotropaceae, and some members of the Polemoniaceae, contained two series of fructose oligosaccharides: one based on isokestose (the inulin series) and the other one based on kestose (the levan series). The inulin or isokestose-based series of fructose oligosaccharides, alone, was present in members of the following families: Menyanthaceae, Boraginaceae, Campanulaceae, Goodeniaceae, Stylidiaceae, Brunoniaceae, Calyceraceae and Compositae. The primary metabolites occur ubiquitously and consistently in tissues of plants of these families. We hypothesize that this conservative character represents several phylogenetic alliances among them.     

Phylogeny of the Dipsacales s.l. based on chloroplast trnL-F and ndhF sequences

Sequences of the chloroplast trnL-F region and 3(') end ndhF gene were used to elucidate phylogenetic relationships and the delimitation of families within Dipsacales s.l. Parsimony analyses of individual and combined data were conducted using maximum parsimony method. The most parsimonious tree based on combined trnL-F and 3(') end ndhF data set recognizes seven major clades of Dipsacales s.l. with the following relationships: Apiales (Adoxaceae ((Diervillaceae, Caprifoliaceae s.str.) (Linnaeaceae (Morinaceae (Dipsacaceae, Valerianaceae))))). Both Sambucus and Viburnum have close relationships with Adoxaceae, supporting their inclusion in this family. Caprifoliaceae s.l. (excluding Sambucus and Viburnum) is polyphyletic, and comprises three clades or families, i.e., Linnaeaceae (Abelia, Dipelta, Kolkwitzia, and Linnaea), Diervillaceae (Weigela and Diervilla) and Caprifoliaceae s.str. (Heptacodium, Leycesteria, Lonicera, Symphoricarpos, and Triosteum). This study focuses on the systematic position of Heptacodium, Triplostegia, and Morinaceae; and suggests that Heptacodium is closely related to the other Caprifoliaceae s.str.; Triplostegia is a sister to Dipsacaceae; Morinaceae, which has an affinity with Dipsacaceae, is possibly a sister group with Dipsacaceae-Valerianaceae clade. Our results are highly congruent with those of and.     

Phylogeny of theAsterales sensu lato based onrbcL sequences with particular reference to theGoodeniaceae

TherbcL gene of 25 taxa was sequenced and analyzed cladistically in order to define more precisely the orderAsterales s.l. and to reconstruct the phylogeny ofGoodeniaceae. The cladistic analyses show that theAsterales comprise the familiesAbrophyllaceae, Alseuosmiaceae, Argophyllaceae, Asteraceae, Calyceraceae, Campanulaceae s.l.,Donatiaceae, Goodeniaceae (includingBrunoniaceae),Menyanthaceae, Pentaphragmataceae, andStylidiaceae. Abrophyllaceae, Alseuosmiaceae, Brunoniaceae, andDonatiaceae have previously not been studied in this respect. Within theGoodeniaceae, four groups supported by therbcL data can be distinguished: the genusLechenaultia, theAnthotium-Dampiera-group, the genusBrunonia, and a group formed by the remaining genera, theScaevola-Goodenia-group.     

Leaf Venation and Its Systematic Significance in Sapindaceae of China

Leaf venation of 27 species representing 25 genera of Sapindaceae (sstr.) of China was investigated for the first time. The pinnate venation pattern in most species is either camptodromous, or craspedodromous. Three types of leaf blade margin were observed, ie., entire, toothed and partite. The secondary veins are branched or unbranched. Most species have intersecondary veins. The tertiary veins of most species are reticulate and percurrent. The areoles are regular or irregular. Veinlets are simple, branched or absent. The delimitations of Xanthoceroideae and Lepisanthes sensu lato are supported by leaf venation characters. The close relationships among Dimocarpus, Litchi and Nephelium are supported by the evidence from leaf venation. A key to the species of Sapinaceae based on leaf venation characters is presented.     

中国无患子科植物的叶脉形态及其系统学意义

对国产狭义无患子科25属27种植物的叶脉形态特征进行了研究报道。结果表明：叶脉均属于羽状脉类型,其中多数为曲行羽状脉,部分为直行羽状脉;叶缘有全缘、具齿和深裂3种类型;二级叶脉具有分支和不分支两种类型;大部分种类具二级间脉,少数不具间脉或间脉不明显;多数种类的三级脉为结网型和贯串型并存;网眼的发育有完善和不完善2种类型;盲脉有简单、具分支和无盲脉3种类型。叶脉形态研究结果支持文冠果亚科以及广义鳞花木属概念,观察发现龙眼属、荔枝属与韶子属从脉序特征方面表现出较近的亲缘关系。编写了国产无患子科叶片脉序特征检索表。     

Preliminary phylogeny of Valerianaceae (Dipsacales) inferred from nuclear and chloroplast DNA sequence data

Valerianaceae is a relatively small (ca. 350 species), but morphologically diverse angiosperm clade. Sequence data from the entire ndhF gene, the trnL-F intergenic spacer region, the trnL intron, the matK region, the rbcL-atpB intergenic spacer region and internal transcribed spacer (ITS) region of nuclear ribosomal DNA were collected for 21 taxa within Dipsacaceae and Valerianaceae (1 and 20, respectively). These data were included in several phylogenetic analyses with previously published sequences from Dipsacales. Results from these analyses (maximum parsimony, maximum likelihood, and Bayesian analysis) are in strong agreement with many of the conclusions from previous studies, most importantly: (1) Valerianaceae is sister to Dipsacaceae; (2) Triplostegia is more closely related to species of Dipsacaceae than to Valerianaceae; and (3) Valeriana appears not to be monophyletic, with Valeriana celtica falling outside the remainder of the species of Valeriana sampled here (with very strong support). With the exception of V. celtica, these data support two major clades within Valeriana; one that is exclusively New World and another that is distributed in both the Old and New World. Although the species of Valerianaceae and its sister group Dipsacaceae plus Triplostegia, are widely distributed in the Northern Hemisphere, and the data imply that Valerianaceae diversified initially in Asia (the Himalayan Patrinia and Nardostachys falling at the base of the clade), the center of modern species diversity for the group is in the Andes of South America with as many as 175 species restricted to that region. Although the exclusively South American taxa form a clade in the chloroplast and combined ITS and chloroplast analyses, support values tend to be low. Future studies will need to include additional data, in the form of both characters and taxa, before any strong conclusions about the character evolution, diversification, and biogeography of the South American valerians can be made.     

Revisiting the phylogeny of Dipsacales: New insights from phylogenomic analyses of complete plastomic sequences

Phylogenetic relationships in Dipsacales have long been a major challenge. Although considerable progress has been made during the past two decades, questions remain; the uncertain systematic positions of Heptacodium, Triplostegia, and Zabelia, in particular, impede our understanding of Dipsacales evolution. Here we use 75 complete plastomic sequences to reconstruct the phylogeny of Dipsacales, of which 28 were newly generated. Two primary clades were recovered that form the phylogenetic backbone of Dipsacales. Seven of the primary clades correspond to the recognized families Adoxaceae, Caprifoliaceae s. str., Diervillaceae, Dipsacaceae, Linnaeaceae, Morinaceae, and Valerianaceae, and one corresponds to Zabelia, which was found to be the closest relative of Morinaceae in all analyses. Additionally, our results, with greatly increased confidence in most branches, show that Heptacodium and Triplostegia are members of Caprifoliaceae s. str. and Dipsacaceae, respectively. The results of our study indicate that the complete plastomic sequences provide a fully‐resolved and well‐supported representation of the phylogenetic relationships within Dipsacales.     

Comparative anatomy and systematics of woody Saxifragaceae. Aphanopetalum Endl.

The floral and vegetative anatomy of the small Australian genus Aphanopetalum were studied. Wood is described for the first time and is characterized by predominantly solitary pores, scalariform vessel element perforation plates with low bar numbers, imperforate tracheary elements with distinctly bordered pits, sparse axial parenchyma, and a combination of homocellular and heterocellular rayS. Starch occurs in both axial and ray parenchyma of the wood. Stems possess unilacunar, one-trace nodes and the uncommon feature of an endodermis with well-defined Casparian stripS. Leaves have anomocytic stomata, a bifacial mesophyll and semicraspedodromous venation or a combination of semicraspedodromous and brochidodromous venation. The tetramerous flowers are apetalous or have minute petals. The compound, half-inferior gynoecium consists of essentially totally united carpels. The pattern of floral vascularization resembles different Saxifragaceae sensu lalo in that the compound sepal-plane and petal-plane traces give rise to staman bundles as well as sepal, petal, and carpel wall venation in their respective planes. The ventral ovarian bundles are fused into a single ventral complex that subdivides at the top of the ovary to form ventral bundles and to supply the one ovule in each locule. Vegetative and floral features provide compelling evidence to suggest that Aphanopetalum has its nearest relatives among the Saxifragaceae sensu lato rather than Cunoniaceae. The genus is probably best treated as forming its own subfamily (or family) among the saxifragaean alliance.     

The complete external transcribed spacer of 18S-26S rDNA: amplification and phylogenetic utility at low taxonomic levels in asteraceae and closely allied families

For molecular phylogenetic reconstruction of some intrageneric groups of plants, a DNA region is needed that evolves more rapidly than the internal transcribed spacer (ITS) of the 18S-26S nuclear ribosomal DNA (nrDNA) repeat. If the region identified is nuclear, it would also be desirable for it to undergo rapid concerted evolution to eliminate problems with coalescence. The external transcribed spacer (ETS) of the nrDNA repeat has shown promise for intrageneric phylogenetic reconstruction, but only the 3' end of the region has been utilized for phylogenetic reconstruction and "universal" primers for PCR amplification have been elusive. We present a method for reliably amplifying and sequencing the entire ETS throughout Asteraceae and some closely allied families. We also show that the ETS is more variable and phylogenetically informative than the ITS in three disparate genera of Asteraceae-Argyranthemum (tribe Anthemideae), Asteriscus (tribe Inuleae), and Helianthus (tribe Heliantheae). The full ETS was amplified using a primer (ETS1f) within the intergenic spacer in combination with a primer (18S-2L) in the 5' end of the highly conserved 18S gene. ETS1f was designed to correspond to a highly conserved region found in Helianthus and Crepis, which are in separate subfamilies of Asteraceae. ETS1f/18S-2L primed in all of the tribes of Asteraceae as well as exemplar taxa from Campanulaceae, Goodeniaceae, and Calyceraceae. For both Argyranthemum and Asteriscus, we were able to directly sequence the ETS PCR products when a single band was produced. When multiple bands were produced, we gel-purified and occasionally cloned the band of interest before sequencing. Although PCR produced single bands for Helianthus species, it was necessary to clone Helianthus amplifications prior to sequencing due to multiple intragenomic ETS repeat types. Alignment of ETS sequences for Argyranthemum and Asteriscus was straightforward and unambiguous despite some subrepeat structure in the 5' end. For Helianthus, different numbers of large tandem subrepeats in different species required analysis of the orthology of the subrepeats prior to alignment. In all three genera, the ETS provided more informative variation for phylogenetic reconstruction and allowed better resolution of relationships than the ITS. Although cloned sequences from Helianthus differed, intragenomic clones consistently formed clades. This result indicated that concerted evolution was proceeding rapidly enough in ETS that species-specific phylogenetic signal was retained. It should be now be possible to use the entire ETS for phylogenetic reconstruction of recently diverged lineages in Asteraceae and at least three other families (approximately 26,000 species or about 8% of all angiosperms).     

7-Polyacylated delphinidin 3,7-diglucosides from the blue flowers of Leschenaultia cv. Violet Lena

The triacyl anthocyanins, Leschenaultia blue anthocyanins 1 and 2 (LBAs 1 and 2) were isolated from the blue flowers of Leschenaultia R. Br. cv. Violet Lena (Goodeniaceae), in which LBA 1 was present as a dominant pigment. The structure of LBA 1 was elucidated to be delphinidin 3-O-[6-O-(malonyl)-beta-D-glucopyranoside]-7-O-[6-O-(4-O-(6-O-(4-O-(beta-D-glucopyranosyl)-trans-caffeoyl)-beta-D-glucopyranosyl)-trans-caffeoyl)-beta-D-glucopyranoside] by application of chemical and spectroscopic methods. Since LAB 2 was isolated in small amount, its structure was tentatively assigned as either delphinidin 3-(malonylglucoside)-7-[(glucosyl-p-coumaroyl)-(glucosylcaffeoyl)-glucoside] or delphinidin 3-(malonyl-glucoside)-7-[(glucosyl-caffeoyl)(glucosyl-p-coumaroyl)-glucoside]. This is the first report of the occurrence of 7-polyacylated anthocyanins in the family of Goodeniaceae, although others have been found in the families of the Ranunculaceae, Campanulaceae, and Compositae. Moreover, delphinidin 3-glycoside-7-di-(glucosylcaffeoyl)-glucoside has been reported only in the flowers of Platycodon grandiflorum (Campanulaceae). From a chemotaxonomical viewpoint, the Goodeniaceae may be closely related to the Campanulaceae.     

Pollen Morphology of Caprifoliaceae from China and Its Taxonomic Signi-ficance

The pollen morphology of 54 samples representing 12 genera and 31 species was investigated with the aid of scanning electron microscope. Observed were pollen grains of Sambucus, Viburnum, Lonicera, Leycesteria, Heptacodium, Linnaea, Abelia, Dipelta, Kolkwitzia, Symphoricarpos, Triosteum, Weigela. Based on the shape, size, position and number of aperture, exine sculpture, three types are recognized: 1. Pollen grains subprolate, less frequently prolate, rather small, 3-colporate, exine reticulate, as in Sambucus, Viburnum. 2. Mostly spheroidal, subolate, bigger than the former, also 3-colporate, exine spinulose as in Lonicera, Leycesteria, Heptacodium, Triosteum, Linnaea, Abelia, Dipelta, Kolkwitzia, Symphoriocarpos, Weigela. 3. Spheriodal, more or less flattend, exine scabrous as in Abelia section Zabelia and Lonicera section Isoxylosteum. 1. The systematic position of Caprifoliaceae: It has been generally treated as a member of the order Rubiales together with Rubiaceae, Valeriaceae and Dipsacaceae on floral characters. In respect to serological character, it has a close relationship with Cornaceae, and was placed in Araliales. The above stated 2nd and 3rd types of pollen grains are similar to those of Patrinia (Valerianaceae), Scabiosa (Adoxaceae), Cornus (Cornaceae), and the pollen grains of the 1st type are similar to those of Styraceae, Genetianaceae and Araliaceae. Taking the information so far available into consideration, the authors agree to the Cronquists treatment retaining Caprifoliaceae in the order Dipsacales together with Adoxaceae, Valerianaceae and Dipsacaceae. 2. The division of tribes: Formerly Sambuceae included the genera Sambucus and Viburnum. Fritsch (1891) segregated Viburnum from Sambuceae and suggested a new tribe Viburneae including Triosteum. There is distinct difference in palynological features between these two genera. The exine sculpture of Viburnum is reticulate, but that of Triosteum is spinulose. It is reasonable to separate another new tribe, Triosteae, from Viburneae. 3. The pollen morphorlogy of several Chinese endemic genera, such as Heptacodium, Dipelta, Kolkwitzia resembles that of Lonicera, Leycesteria, Linnaea, Symphoricarpos, Abelia, Triosteum. This evidence supports the foregoing treatment including them in Caprifoliaceae. 4. Two different exine sculptures are shown in sections of the genera Abelia and Lonicera. In Abelia the exine of the section Euabelia is spinulose, but that of the section Zabelia is scabrous. Likewise, in Lonicera, the exine of the section Isoxylosteum is scabrous, while that of other sections such as Nintooa, Isika, Lonicera, subgenus Caprifolium, is spinulose. It shows that pollen morphology is one of diagnostic characters for section division.     

Phylogenetic affiliations of members of the heterogeneous lichen-forming fungi of the genus Lecidea sensu Zahlbruckner (Lecanoromycetes, Ascomycota)

The genus Lecidea Ach. sensu lato (sensu Zahlbruckner) includes almost 1200 species, out of which only 100 species represent Lecidea sensu stricto (sensu Hertel). The systematic position of the remaining species is mostly unsettled but anticipated to represent several unrelated lineages within Lecanoromycetes. This study attempts to elucidate the phylogenetic placement of members of this heterogeneous group of lichen-forming fungi and to improve the classification and phylogeny of Lecanoromycetes. Twenty-five taxa of Lecidea sensu lato and 22 putatively allied species were studied in a broad selection of 268 taxa, representing 48 families of Lecanoromycetes. Six loci, including four ribosomal and two protein-coding genes for 315- and 209-OTU datasets were subjected to maximum likelihood and Bayesian analyses. The resulting well supported phylogenetic relationships within Lecanoromycetes are in agreement with published phylogenies, but the addition of new taxa revealed putative rearrangements of several families (e.g. Catillariaceae, Lecanoraceae, Lecideaceae, Megalariaceae, Pilocarpaceae and Ramalinaceae). As expected, species of Lecidea sensu lato and putatively related taxa are scattered within Lecanoromycetidae and beyond, with several species nested in Lecanoraceae and Pilocarpaceae and others placed outside currently recognized families in Lecanorales and orders in Lecanoromycetidae. The phylogenetic affiliations of Schaereria and Strangospora are outside Lecanoromycetidae, probably with Ostropomycetidae. All species referred to as Lecidea sensu stricto based on morphology (including the type species, Lecidea fuscoatra [L.] Ach.) form, with Porpidia species, a monophyletic group with high posterior probability outside Lecanorales, Peltigerales and Teloschistales, in Lecanoromycetidae, supporting the recognition of order Lecideales Vain. in this subclass. The genus name Lecidea must be redefined to apply only to Lecidea sensu stricto and to include at least some members of the genus Porpidia. Based on morphological and chemical similarities, as well as the phylogenetic relationship of Lecidea pullata sister to Frutidella caesioatra, the new combination Frutidella pullata is proposed here.