Revised scientific names of the genus Hemileia (Pucciniales) based on the new ICN

Based on the new International Code of Nomenclature for algae, fungi, and plants, specifically the effective deletion of Article 59, we provide a list of the revised scientific names of species in the genus Hemileia (Pucciniales). Five new combinations are proposed.


Introduction
At the 18th International Botanical Congress in Melbourne, July 2011, the members agreed to fundamental changes concerning the naming of fungi. Of particular interest is the ending of the dual nomenclature system for fungi as previously governed by Article 59 of the outdated International Code of Botanical Nomenclature (McNeill et al. 2006). In the new International Code of Nomenclature for algae, fungi, and plants (ICN), one species of fungus may have only one scientific name. The use of separate names for the sexual and asexual states is no longer allowed. Thus, the legitimate and validly published names for one species must be considered with priority given to the oldest species epithet. This new rule is effective immediately as dictated by the ICN (McNeill et al. 2012). Although moving to the use of one scientific name is problematic especially for rust fungi, this brings the fungi in line with all other groups of organisms.
The taxonomy of rust fungi has been confused partly because of the difficulty in determining sexual and asexual states and connecting the various states for an individual species. Many genera were defined based on the presence of teliospores that produce basidia. Also serving as the resting or overwintering stage, teliospores are produced on dikaryotic hyphae with meiosis taking place in the basidia that develop from germinating teliospores. Asexual states usually bearing aeciospores or urediniospores were placed in genera such as Aecidium and Uredo that generally lacked phylogenetic meaning i.e. these genera did not unite related species. Some rust species that lacked known teliospores were described using scientific names in the appropriate genus but these were considered invalid names because of the lack of teliospores as dictated by Article 59 of the International Codes of Botanical Nomenclature such as McNeill et al. (2006). Once a specimen with teliospores was discovered, a new name with another type specimen was published in the appropriate genus or, in some cases, the old name was considered to be validated by the species having teliospores. With the changes in the rules of the ICN, the scientific names of many rust fungi must be re-evaluated with priority given to the oldest epithet for a species regardless of the genus in which it was described.
The objective of this paper is to provide a list of scientific names of species in the rust genus Hemileia that brings the scientific names into accordance with the ICN. The genus Hemileia was recently revised by Ritschel (2005). At that time a number of species were placed in Uredo even though they were considered to belong in Hemileia because they lacked teliospores. With the new ICN these names compete for priority and can now be placed in Hemileia. This revision of the scientific names of Hemileia is based entirely on the account of the genus published by Ritschel (2005). Of the 42 names included Ritschel (2005), 26 names need to be changed i.e. the correct scientific name already exists but was not recognized as such in Hemileia. These include five new combinations required because the oldest epithet needs to be placed in the genus Hemileia. The scientific name of the fungus that causes coffee rust, thankfully, remains unchanged, as Hemileia vastatrix.

Material and methods
This account is based on the Ritschel (2005) "Monograph of the genus Hemileia (Uredinales)." Further literature was consulted when the comments in Ritschel (2005) did not provide sufficient information. For the implementation of the new rules concerning dual nomenclature, we consulted Braun (2012), Hawksworth et al. (2011) andNorvell (2011), articles describing the most significant changes and impacts of the new ICN on dual nomenclature of pleomorphic fungi. = Hemileia smalliana Gjaerum, in Gjaerum et al., Lidia 5: 2, 2000. [= Hemileia smallii Fernier, Rev. Mycol. (Paris), Suppl. Colon. 19: 62, 1954hom. illeg. non Wakef. & Hansf. 1949 Comments. Following the new ICN, Hemileia smallii has priority even though the type specimen includes only the asexual stage of this species. Fernier (1954) described a name for the teleomorph using a different type specimen. Because he used the same epithet, that name is an illegitimate, later homonym. For this reason Gjaerum et al. (2000) established another name for this species based on a different type specimen that is here regarded as a taxonomic synonym. Comments. Based on the comments in Ritschel (2005), Uredo solaninum is considered to be the oldest name for this species and is thus transferred to Hemileia. The other names are based on different type specimens that occur on different hosts but are considered by Ritschel (2005) to apply to the same species, thus they are listed here as taxonomic synonyms.

Project features and concepts for the future
For the purpose of giving access to treated taxa, monographic and floristic treatments are often assigned with traditional dicho-or polytomous identification keys. Until today, such keys have proven to be of highest value as they mostly provide identification criteria optimized for a specific set of taxa or an area. Nevertheless, the growing use of the internet as well as the need of open access to information and knowledge on biodiversity at a higher or global scale has led to an increased use of internet-related technologies for identifying taxa (Hagedorn et al. 2010;. Over the past 12 years, the lichen trait database LIAS light (http://liaslight.lias. net) has grown to a considerable pool of descriptive data for lichens with information on morphological, ecological and chemical traits. It is part of the LIAS data platform, which has been initiated as a web project in 1995 (Rambold 1996, Triebel et al. 2007 and actually includes two further active segments: LIAS names (http://liasnames.lias. net/) with more than 40,000 names including synonyms, isonyms and homonyms, as well as LIAS glossary (http://glossary.lias.net/). Data from LIAS names along with data from LIAS light form the Global Species Database LIAS (Rambold 2012 onwards; http://www.catalogueoflife.org/col/details/database/id/79) as taxonomic backbone for lichens in the megascience platform Catalogue of Life (Triebel et al. 2012).
There are two other parts of the site (LIAS main and LIAS checklists) which have been frozen in favour of LIAS light which works based on a reduced number of descriptors (see below) and will incorporate in the near future occurrence data at the country level as well. While the concept of LIAS main was based on lichenologist community driven data input, entry of LIAS light data was primarily taken over by laboratory members of the first author, starting with compilation data from outstanding lichen floras like Wirth (1995) for Baden-Württemberg (Germany), Purvis et al. (1992) for the United Kingdom, Nash et al. ( , 2004Nash et al. ( , 2007 for the Sonoran Desert (U.S.A. and Mexico). By inclusion of a wide range of additional literature with descriptions of lichen taxa from all regions of the world, the project first slowly, then rapidly increased with regard to the number of taxa included and quality of entered data.
The number of taxa is now close to 10,000. Estimate numbers of accepted names of lichens at a global scale range between 13,500 and 20,000. Given the number of 17,000 described, accepted lichen species of the world as proposed by Chapman (2009), LIAS light content now covers about 60% of the accepted taxa. The included traits are based on about 71 different qualitative (i.e. categorical), quantitative (i.e. numerical), and text characters. Several are only used for data management issues, but about 50 only concern diagnostic traits to be used for identification and data analysis purposes. The full list of these characters is provided under http://liaslight.lias.net/ Identification/Navikey/World/en_GB/chars. Outstanding and therefore worthwhile to be mentioned is the categorical trait 'secondary metabolites <compound>', which covers about 1,020 lichen metabolite names as character states. The highly different quality of the individual data records ('species profiles') is due to the most different level of completeness in the literature sources, which include original articles with species descriptions as well as monographic treatments. This means that the coverage and reliability of traits per taxon is relatively unequally distributed. However, the individual data records of each species are being continuously revised by considering additional articles with relevant and reliable species descriptions or trait data. Of the current 10,000 data records, entered by members of the editorial board and co-workers, about 3,000 data records have already been revised at least once. Aside from the purpose of identification, LIAS light data collection may also be applied as background data for reconstructing phylogenetic character evolution by mapping of character states onto phylogenetic trees, as well as for ecological studies with a focus on the distribution of phenotypic traits in a given area by correlating environmental or occurrence data (Zedda et al. 2011).
The taxon names used for the LIAS light dataset are maintained in LIAS names. For most of the records, modern taxonomic concepts are applied when literature references exist. The higher classification backbone is based on MycoNet data (http://fieldmuseum. org/explore/myconet) and updated accordingly. By activating the resulting LIAS light taxon names, access to additional information and links to further resources is provided. Aside form LIAS names (see above), they include a) an interface using the web service of Recent Literature of Lichens (http://www.bio.uni-bayreuth.de/liaslight/service/rll_wrap. php); b) a lichen image accession interface (http://www.bio.uni-bayreuth.de/liaslight/ service/LichenImages.php) to reliable web sites with significant images; and c) several less specific portals like Wikispecies, Google Images, NCBI Search and others.
Associated with LIAS light is LIAS metabolites (http://liaslight.lias.net/Identification/Navikey/Metabolites/index.html), a database compiled by J. A. Elix with additions of K. Kalb (Neumarkt, Germany). This LIAS database component includes the values of TLC, HPLC and MS for about 880 secondary lichen compounds, together with additional information like the substance class and names of related metabolites. It is useful for interpreting chromatographic and spectroscopic analysis results. Soon each data record of LIAS metabolites will be supplemented with the chemical formulas and further representations including SMILE codes.
Multilinguality or internationalization options have become a central challenge of the LIAS light project. At present, versions in 18 languages exist: aside from English, being Arabic, Armenian ( Fig. 1), Chinese, Esperanto, Farsi, French, German, Hebrew, Italian, Korean, Kyrgyz, Polish, Portuguese, Romanian, Russian, Uzbek, Spanish, and Turkish. The translation of the subject-specific vocabulary is intended as a contribution of preventing 'lingual erosion' in the languages aside from English. Regarding the 18 languages actually included in LIAS light, it becomes apparent that several ones (like French, German, Polish, Russian, Spanish etc.) easily cover all general and specific terminologies, while others (like Kyrgyz or Chinese) are only capable of providing the general terms but not or much less the specific ones, (e.g. the scientific names of secondary metabolites). In such cases, supplementation of the respective character state list by terms from English or Russian (in the case of Kyrgyz) or another widely spoken language is required. The use of a native language is desired, especially by students as well as amateur communities, who are involved in monitoring projects in many countries of the world. The translation of the LIAS light character list and the NaviKey web interface contents into the various languages have been achieved by colleagues in biology or lichenology (http://liaslight.lias.net/About/Impressum.html).
LIAS light content is provided via NaviKey, a Java applet and application for accessing descriptive data (http://www.navikey.net/). It provides quite a range of possibilities to modify the queries for obtaining improved results. One of the most important option is 'Retain items unrecorded for the selected characters' being a switch between a 'data query mode' (checkbox "set off", as the default) and an 'identification mode' (checkbox "set on"). While the query mode is strict and only provides the directly matching data according to the selection, the identification mode is more tolerant in the way that it matches positively also in the case, that a respective character state is not specified in the respective data record. During identification, the option to simultaneously select more than one character state, either with OR ('Retain matching at least one selected state' checkbox "set on", as the default) or with AND (checkbox "set off") selection criteria, facilitates one making decisions when specifying states of categorical characters. For instance, the decision to select certain colour nuances may be relatively difficult. However, switching between the two modes is not yet possible during one and the same identification procedure, but will be implemented in the future.
The LIAS light database with its NaviKey web interface still exhibits certain shortcomings and does not fully provide satisfactory results in all cases. Nevertheless, it is foreseeable that by continued data entry and revision and with some improvements of the user interface, LIAS light will become a feasible tool for lichen identification. Especially the intended inclusion of country checklist information from reliable original sources (e.g., the original checklists of Austria, Australia, France, Germany, or North America), will entail improved resolution of data and therefore more reliable identification results, especially for countries with a more or less saturated knowledge of taxon occurrence. The use of checklist information as a prefilter function also beneath country level is being considered. Since October 2013, GBIF offers the option of user-defined filter creation for dynamically generating spatially defined species lists. By implementing access to this service LIAS light will be enabled to use this additional option of taxon preselection. For the next future, functionality of the NaviKey software will be extended by enabling the dynamic generation of taxon description web pages. LIAS metabolites will be optimized and new languages will be added on request.
All contents of the lingual versions are downloadable under a Creative Commons Attribution-Non-Commercial-No Derivative Works 3.0 Unported License as archive data files from the respective LIAS light language version page (e.g., http://liaslight.lias. net/Identification/Navikey/data/download/LIASlightDataFilesWorld.pl_PL.zip for the data set in the Polish language). The data may be evoked by a local installation of Na-viKey and be even used offline, which, for instance, may be of value for work in remote areas with low bandwidth internet. Server logs indicate an increasing acceptance of LIAS light. The relative number of re-visitors or users, respectively, is steadily growing and is a clear argument in favour of internationalizing the project -there is a striking correlation between visitor location at the country level and the language version being selected. A citation of LIAS light by users in their publications is highly appreciated and should be in accordance with the imprint page (http://liaslight.lias.net/About/Impressum.html).
The work was supported in part by the Federal Ministry of Education and Research, Germany (BMBF) under project ID 01 LI 1001 B 'GBIF-D' and by the German Research Foundation (DFG) as a project of the LIS infrastructure program grants (project IDs INST 747/1-1, RA 731/11-2, and TR 290/5-1) and in the context of the funding initiative DFG/ CNPq: German-Brasilian consultative visits under DFG project ID 444 BRA -113 / 58 / 0-1. Support was also granted by the European Union's 7 Framework Programme (project ID FP7/2007-2013 in the 4D4Life project context (grant agreement ID 238988).

Two new species of Ramaria from Arkansas Introduction
The annual national foray of the North American Mycological Association was held in northern Arkansas in 2013, specifically in the lower Boston Mountains of the Ozark Ecoregion. Among collections of fleshy fungi were several species of Ramaria (Gomphaceae, Gomphales, Agaricomycotina), of which two appear to be new to science and are described below.
American literature dealing with the modern genus Ramaria may have begun with the treatment by Burt (1922), followed by that of Coker (1923), both of whom used an expansive concept of Clavaria to shelter taxa of the modern Ramaria. Maxwell Doty (1944) dealt with some taxa of Ramaria in his thesis on Clavaria (s.l.) from Oregon and the Pacific Northwest. Corner (1950Corner ( , 1970 greatly expanded the concept of Ramaria and treated the genus worldwide. The genus in Washington State was taken up by Currie D. Marr in his PhD dissertation, published jointly with his major professor, Daniel E. Stuntz (Marr and Stuntz 1973). Their coverage centered in the Pacific Northwest and not only revealed that region to be unusually speciose, but divided Ramaria species into four subgenera, of which the largest was subg. Laeticolora, which took in the largest, fleshiest, most brightly colored basidiomata theretofore known. The same geographical region was summarized by Exeter (2001) and Exeter et al. (2006). Over the years, Petersen (see bibliography by Exeter et al. 2006) published shorter papers attempting to bring order to the genus and describing some new taxa, chiefly from eastern North America. As a result, although the temperate rain-forests of western North America and, to a lesser extent the temperate forests of eastern North America have been treated scientifically, almost no scientific literature exists for Ramaria in the mid-United States, and this appears especially true of the Ozark Plateau in northern Arkansas. Pine et al. (1999) were the first to explore the clavarioid and cantharelloid fungi using molecular techniques. Among several discoveries, Ramaria was linked to significantly different morphological genera (i.e. Clavariadelphus, Lentaria, Gomphus, Sphaerobolus, Pseudocolus, etc.) in the "gomphoid-phalloid clade," an alliance which has appeared repeatedly in other phylogenies since its discovery. Humpert et al. (2001) dissected the Gomphales in further detail in an attempt to ascertain relationships of and within the genera. At the resolution of nuclear nrLSU, some traditional infrageneric groups (i.e. Laeticolora, Lentoramaria) were found to be polyphyletic, while Echinoramaria seemed to segregate as a discrete clade. Morphologically discordant elements (i.e. Gomphus, Gautieria) were found within the large Ramaria clade. Although Humpert et al. (2001) revealed the considerable heterogeneity within Ramaria, many Ramaria LSU sequences were deposited in GenBank, forming a nucleus for ongoing comparison.
In two seminal publications, Christan (2008) and Christan and Hahn (2005) summarized several previous systematic schemes involving the entire genus. Dealing with the group traditionally referred to as subg. Laeticolora, it was accepted that Laeticolora was best subsumed into subg. Ramaria, which included section Ramaria [with typus generis Clavaria (Ramaria) botrytis] and section Formosae, typified by R. formosa (see Franchi and Marchetti 2001;Christan and Hahn 2005). In the latter section most of the fleshy, coralloid, brightly colored basidiomata could be placed, and according to this outline, the two species described here as new belong in sect. Formosae. A third section, Fennicae (type, R. fennica, see Schild 1995) was reserved for certain species with violaceous coloration and hymenial color change in KOH. Christan and Hahn (2005) detailed further infrageneric taxonomic characters, among them the anatomy of rhizomorphic strands and/or basidiomatal basal mycelium. Several types and complexities of acanthodendroid hyphae were described and illustrated, as well as crystal formation and morphology. Again, with these hyphal characters in mind, the two species described below belong in section Formosae, but are not to be found in the keys by Christan (2008; for European taxa) or Exeter et al. (2006; for Pacific Northwest United States).

Methods
Microscopic observations employed an Olympus BX60 microscope fitted for bright field (BF) and phase contrast (PhC) microscopy. Typical mountant was 3% aqueous KOH, with Melzer's reagent and lactic acid-Cotton blue occasionally used as stains. TFB = Tennessee Fieldbook number, assigned in the field to track notes and photos; TENN = Herbarium, University of Tennessee, permanent accession number; Q = spore length divided by spore width; Q m = mean Q value of number of spores measured; L m = mean length of all spores measured. Colors within quotation marks are from Ridgway (1912) and those within parentheses are from Kornerup and Wanscher (1967).
The term "firm-walled" is intended to describe hyphal walls which are thick enough so that hyphal crumpling or collapse is not observed in microscopic mounts, but there is no measurable distance between inner and outer wall surfaces. "Thick-walled," conversely, describes a hyphal wall thick enough that the inner and outer surface are sufficiently distinct and that the wall thickness can be measured and reported.
Procedures for DNA extraction, PCR and DNA sequencing of the ribosomal ITS and LSU regions was performed as described in Hughes et al. (2013). Initial blast searches of the NCBI database were performed to identify related taxa for which sequences were available. In order to place collections from Arkansas in a larger context, the ribosomal LSU region was selected for analysis. Related sequences were downloaded, combined with similar sequences in our unpublished sequence database, aligned manually and trimmed to 905bp containing D1-D3 variable regions using GCG (2000). Three small areas (4-5bp) of ambiguously aligned sequence were not excluded from the dataset because they were informative in species delineation. The alignment was imported into GENEIOUS (2005). PhyML using the general time reversible model of substitution with parameters estimated and 100 bootstrap replicates was performed within GENEIOUS and the resulting tree exported to FIGTREE (Rambaut 2006). This model requires the fewest assumptions of the data set. The likelihood of the best tree was -2745.59. ITS and LSU sequences were deposited in GenBank (KJ416132-KJ416135). Alignments and the PhyML tree were deposited in TreeBASE (http://purl.org/phylo/treebase/phylows/study/TB2:S15885).  Etymology. admiratia = surprise, astonish, referring to the acanthodendroid hyphae in the outer stipe context. Diagnosis. 1) Member of Ramaria subg. Laeticolora sect. Formosae; 2) clamp connections absent from all tissues; 3) acanthodendroid hyphae common in outer stipe flesh and surface; 4) stipe large, pruinose, white but easily staining to brown where handled or rubbed; 5) branch apices bright orange red; 6) type locality, northern Arkansas; 7) ITS sequence unique in the subgenus; (GenBank ITS accession KJ416133).
Habitat and phenology. Generally second-growth hardwood forest of Quercus with scattered Carpinus, Carya and Acer; sole specimen from late autumn.
As noted by Christan (2008) taxa with cyanophilous dendrohyphidioid stipe tomentum hyphae are more common than originally envisioned (Marr and Stuntz 1973). Most common among them are taxa with yellow branches and apices and rather small stipes. In taxa resembling R. admiratia, centering around R. cystidiophora, varieties have been described based largely on odors (vars. anisata, fabiolens) and rubribrunnescent stipe (var. maculans) (see Marr and Stuntz 1973;Exeter et al. 2006). Conversely, R. admiratia basidiomata exhibit a large, strongly brunnescent stipe and orange to red-orange branches and apices.
The terms "monomitic," "dimitic", etc. were coined by E.J.H. Corner (1932) to describe the anatomy of various polypore basidiomata and have proven useful in subsequent, widely used taxonomic schemes. Dimitic referred only to presence of two hyphal types, specifically generative and skeletal, definition of the latter being philosophically augmented over some decades. Donk (1971) and Petersen (1975), conversely, while accepting the anatomical observations, linguistically considered dimitic to describe any tissue in which two hyphal types were found, whether generative and skeletal, generative and gloeoplerous, generative and acanthanophysoid, etc., as long as the hyphal types were accurately described. In the case of Ramaria admiratia (and other fleshy fungi with various second hyphal types), one is confronted with Cornerian terminology in which "monomitic" actually includes two hyphal types: generative plus cyanophilous, characteristically branched acanthophysoid hyphae. Etymology. Calvus = bald; distalis = referring to the spore wall opposite the hilar appendage.
Habitat and phenology. Possibly associated with deciduous trees from local forests of Quercus, Carya, Carpinus and occasional Pinus, solitary to gregarious, often in troops or rings; fruiting in late autumn.
Hyphae of stipe canescence 2-4 μm diam, relatively brittle and straight, firmwalled, rarely septate, without clamp connections, non-refringent; acanthodendroid hyphae absent; in non-gelatinous areas of stipe flesh hyphae 3-12 μm diam, tortuous, frequently branched, thick-walled (wall -0.7 μm thick), often refringent (PhC), without clamp connections; rare ampulliform swellings (without clamp connection) -16 μm diam, delicately ornamented internally, not unusually thickwalled. Hyphae of upper branch trama appearing subgelatinous under low magnification, but when squashed in KOH shown to be free, 3.5-12 μm diam, without clamp connections, firm-walled (wall -0.5 μm thick); cells filamentous to elongatebarrel-shaped. Basidioles often misshapen, paraphysoid, with various small lobes or sinuate shapes. Basidia (Fig. 7A) 55-72 × 12-13 μm, clavate with somewhat bulbous apex, 4-sterigmate, occasionally with an asymmetric lobe, without clamp connections; contents usually with proximal and distal refringent guttules. Basidiospores (Fig. 7B) (12-)14-15 × 4.5-5.0(-5.5) μm (Q = 2.67-3.33; Q m = 3.03; L m = 14.05 μm), generally boletoid, with scattered small cyanophilous warts and patches through midsection of the spore but absent from the distal end which appears bald; contents with amorphous deposits (PhC); wall slightly thickened through midsection (wall -0.5 μm thick).  Commentary. Care must be taken to ascertain the condition of basidial septa. In clamped basidia, subsequent basidioles arise through the subtending clamp. In clampless basidia (as above), subsequent basidioles arise in precisely the same fashion, arising just below the subtending septum, but without the telltale evidence of a clamp connection. Additional care is required to ascertain that both tramal hyphae and basidia are without clamps. Spores are among the longest in the subgenus and largely as a result, Q m value is high. Spore outline is distinctly boletoid with slight suprahilar depression. First impression was of delicately marbled stipe flesh (i.e. with small, scattered areas of hyphae with gelatinized walls), but closer examination revealed scattered degeneration of inner stipe flesh through maggot infestation. Lower and middle branches as well as apices are essentially unicolorous, straw yellow or dull ochraceous yellow. Juvenile branches exhibit a slight blush of pale pinkish buff, but this soon fades and slowly turns to the adult yellow shades.
In the three days of the NAMA foray, numerous collections of this species were made. Basidiomata seem to occur in troops and "fairy rings" under deciduous trees and are often somewhat bulky. Stipes are not deeply rooted and are easily dislodged, but adult stipes seem consistently maggot-ridden. Because a new taxon was not anticipated, only the type collection was preserved. Although the literature dealing with Ramaria of the Pacific Northwest has been summarized at least twice over the decades (Marr and Stuntz 1973;Exeter et al. 2006), modern literature is unavailable for Ramaria east of the Rocky Mountains except in fragments. Even less adequate is coverage of central United States, including the Ozark Mountains of Arkansas. Marr and Stuntz (1973) described a small group of Ramaria taxa which exhibited cyanophilous "acantho-dendroid" hyphae in the outer stipe flesh, using R. cystidiophora as the focal taxon, but including several infraspecific taxa. In all cases, stipes are white and pruinose and all upper basidiome parts are some shade of yellow. Exeter et al. (2006) illustrated basidiomata of several of the infraspecific taxa in the R. cystidiophora complex, which macroscopically are reminiscent of R. calvodistalis, so a special search was made for acanthodendroidal hyphae. Not only were no such hyphae found, but R. calvodistalis lacks clamp connections, also a violation of the R. cystidiophora complex.
If the key to clampless taxa in Exeter et al. (2006) is employed, no adequate match is found. Ramaria longispora produces spores of appropriate dimensions, but while upper branches and apices are yellow, lower branches are cantaloupe or pinkish salmon. In the working key of one of us (RHP), no match is found because the combination of yellow basidiome coloration, lack of clamp connections and long spores eliminates all candidates.
Ramaria admiratia and R. calvodistalis LSU sequences place them near sequences representing brightly colored Ramaria species (R. aurantiisiccescens and R. araiospora) in subgenus Laeticolora (Fig. 8). ITS divergence within this subgenus is large, however, and Ramaria admiratia and R. calvodistalis ITS sequences are only 86% similar to each other. Ramaria calvodistalis ITS sequences are most closely related (>97%) to two unnamed collections from Mexico (GenBank KC152173 and KC152176). These three collections differ from each other predominantly in the number of bases in repeat areas and probably represent the same lineage. We have previously noted that Mexico may have served as a glacial refugium for taxa now found further north (Hughes et al. 1999;Lickey et al. 2002;Hughes and Petersen 2004). There are no close blast matches to Ramaria admiratia in GenBank or in our sequence data set. A number of taxa in Fig. 8, based on assigned names, appear to be polyphyletic suggesting that morphological species concepts harbor more than one cryptic species or that misidentifications are common.

Introduction
Tremella (both Tremellales) and Syzygospora (Filobasidiales). There are currently 50 lichenicolous species described in the group (Diederich 1986, 2003, Sérusiaux et al. 2003, Diederich 2007, Zamora et al. 2011, Millanes et al. 2012, and an additional five are known but have not yet been formally named (Diederich 1996(Diederich , 2007. The actual diversity and distribution of lichen-inhabiting species is still largely unknown, as comparatively few mycologists and lichenologists collect lichenicolous fungi. Some tremellalean species are relatively easy to spot in the field as they induce the formation of conspicuous galls on the hosts, whereas others grow only inside the host hymenium producing few or no external symptoms. In these cases, study of host herbarium material is extremely useful for diversity surveys, and also improves our knowledge on the geographical distribution of taxa that are otherwise much overlooked in the field. In the Nordic countries, 15 tremellalean lichenicolous species have been reported so far (Nordin et al. 2014). During fieldwork in Sweden and Norway, and herbarium surveys undertaken in BG, S, and UPS, during 2012-2014, we found material allowing one new species to be formally described and several species to be newly reported to the Nordic countries, Iceland, Norway or Sweden.

Methods
Macromorphological traits were observed using a Zeiss Stemi 2000-C dissecting microscope. Microscopic structures were studied using handmade sections stained with Phloxin (1% in water) after pre-treatment with KOH (5%), following the methods of Diederich (1996), and observed with a Zeiss Axioscope-2 microscope. Macro-and microscopic photographs were taken with a Nikon Coolpix 995 camera fitted to both microscopes. The apiculus is not included in basidiospore measurements. Sizes in parentheses represent minimum and maximum observed values.
Distribution, ecology and host. The new species is known from one locality in Denmark, two localities in Norway, and nine in Sweden, all growing inside the hymenium of Rhizocarpon lavatum on siliceous rocks, often along shorelines or near water.
Discussion. The species was tentatively described by Diederich (1996: 170) as Tremella sp. 3. The single specimen known at that time was very small, but the discovery of 12 additional collections from Denmark, Sweden and Norway (Fig. 1A) is now enough for a formal description. There is little to add to the description and illustrations given by Diederich (1996), however. The only novelties are the wider range of basidium width and length (Fig. 1G) and the presence, occasionally, of an anamorph consisting of catenulate conidia. A sequence of the ITS barcode region is not provided, since recently collected specimens were too reduced for DNA extraction, and old specimens could not be amplified by PCR. Diederich (1996) indicated that this species was microscopically very similar to Tremella protoparmeliae, but T. rhizocarpicola could be distinguished by the formation of dark brown swellings on the host apothecia and by small differences in the size of the basidia. The additional specimens studied here show that there is an anamorph formed by catenulate conidia in T. rhizocarpicola, and the basidia of this species can be comparatively shorter and wider than those of T. protoparmeliae.

Tremella pertusariae Diederich
Note. This species was previously recorded in Europe (Belgium, France, Germany, Great Britain, Ireland, Italy, and Spain), where it is very common in the western Pyrenees, and in America (Mexico and U.S.A) (Diederich 1996). Knutsson and Johansson (2011) reported it from Sweden, but with some hesitation since the material could not be identified microscopically. The five additional specimens reported here confirm the presence of the species in the Nordic countries. The fungus forms pale to dark brown or blackish galls, generally on the apothecia of Pertusaria hymenea (Fig. 1B), but also on the thallus of both P. hymenea and P. pertusa, and bears characteristic stalked basidia (Fig. 1H).

Tremella protoparmeliae Diederich & Coppins
Note. This species was previously only recorded in England (Diederich 1996). The fungus grows intrahymenially on Protoparmelia badia, without producing any external symptoms, which suggests that this species could in reality be common, but much overlooked as it is not visible macroscopically (Figs. 1C and 1I).

Syzygospora physciacearum Diederich
Note. New to Iceland and to the Swedish provinces Öland, Södermanland, and Västergötland.
The species was at first tentatively described as Tremella sp. 1 by Diederich (1996) and eventually formally described by Sérusiaux et al. (2003). It has been reported from Austria, Canada, France, Great Britain, Greece, Greenland, and Spain (Diederich 1996, 2007, Sérusiaux et al. 2003, growing inside the hymenia, or exceptionally on the thallus, of different Caloplaca species, viz. C. arenaria, C. arnoldii, C. aurantia, C. carphinea and C. saxicola (Diederich 1996, Sérusiaux et al. 2003, and on the thallus of Xanthoria sorediata (Diederich 2007). It was recently reported as new to the Nordic countries (Thell et al. 2014) from two Swedish collections, growing on new hosts: Caloplaca decipiens (on the thallus; Fig. 1D) and Xanthoria parietina (inside the hymenium; Fig. 1E). Here it is reported on a new host (Xanthoria elegans), in Sweden. All samples below are on X. elegans, growing inside the hymenium or on the thallus. All specimens bear two-celled basidia (Fig. 1J)  Note. New to Sweden.
This fungus was previously known only from Denmark (Diederich 1996). It grows in the hymenium of Rinodina gennarii, forming distinct brown swellings on the host apothecia (Fig. 1F). The basidia have one transverse (Fig. 1K), or exceptionally longitudinal (Fig. 1L)

Biatoropsis usnearum Räsänen
Note. New to Bohuslän. This species has a wide distribution and has been recorded in all Nordic countries except Iceland (i.e., Denmark, Finland, Norway, and Sweden) Christiansen 1994, Nordin et al. 2014).
This material had been placed in S long ago, but as the specimens have apparently not been published or otherwise recorded elsewhere, we report them here. The species had been previously recorded in the Nordic countries, particularly in Sweden and Finland (Diederich 1996).
The species has previously been recorded in the Scandinavian Peninsula, i.e., Norway and Sweden (Holien andTønsberg 1994, Diederich 1996).
In the Nordic countries the species has been so far only reported from Sweden (Millanes et al. 2012).

Pycnopulvinus aurantiacus gen. et sp. nov., a new sporocarp-forming member of Pucciniomycotina Introduction
The majority of Pucciniomycotina (Basidiomycota) species have life cycles that include the production of microscopic fruiting structures (e.g., spermogonia of rust fungi), but only a few species within the subphylum form macroscopic fruiting bodies (Swann et al. 2001;Aime et al. 2014). Although these sporocarps vary in form (e.g., see Aime et al. 2014), many are stilboid or pycnidioid with spore masses produced at the base of the sporocarp and exiting through the tip of the neck. Fungi with such fruiting bodies can be found mostly in Atractiellomycetes (Oberwinkler and Bandoni 1982) and, as is true for many other members of Pucciniomycotina, little is known about their biology or biodiversity.
Heterogastridium pycnidioideum Oberw. & R. Bauer (anamorph Hyalopycnis blepharistoma (Berk.) Seeler) is one such fungus that produces stilboid fruiting bodies. It is a strictly filamentous species with simple septal pores and specialized organelles-colacosomes-associated with mycoparasitism (Oberwinkler et al. 1990). Heterogastridium is placed in Heterogastridiaceae and Heterogastridiales (Oberwinkler et al. 1990), which are now placed in Microbotryomycetes (Weiss et al. 2004).  placed three genera in Heterogastridiaceae in addition to Heterogastridum-Atractocolax R. Kirschner, R. Bauer & Oberw., Colacogloea Oberw. & R. Bauer, and Krieglsteinera Pouzar. Of these, neither sequence data nor reference cultures are available for two of the genera, Atractocolax and Krieglsteinera, both of which are monotypic. Colacogloea, on the other hand, has been demonstrated several times to form a separate clade of Microbotryomycetes, distant from Heterogastridiales (e.g. Aime et al. 2006;Kurtzman et al. 2011). Thus, at present, only Heterogastridium and its sole species, H. pycnidioideum, can be confidently assigned to Heterogastridiales with molecular data.
An unusual stilboid fungus was discovered on the mid-rib of a palm leaf in litter that could not be confidently assigned to any previously described genus. DNA sequence data and phylogenetic analyses indicated that the collection represents a new member of Heterogastridiales. Herein we describe and illustrate Pycnopulvinus aurantiacus gen. et sp. nov. and provide a phylogenetic analysis of Heterogastridiales.

Methods
The specimen (PUL F2679) was collected near Bilsa Biological Station in Ecuador, in the vicinity of N0.350444, W79.732075, on 3 May 2004, where it was growing on the mid-rib of a dead palm leaf in the litter. The fungus was photographed and described in the field after which small pieces of substrate bearing the fruiting bodies were dried on an herbarium drier. Color designations refer to Kornerup and Wanscher (1978). Duplicates are deposited in the Kriebel Herbarium at Purdue University (PUL) and the herbarium of the Pontificia Universidad Catolica del Ecuador (QCA).
Morphological characters of the fruiting bodies were observed first with an Olympus SZ61 dissecting microscope. With the aid of the dissecting microscope, a few fruiting bodies were carefully removed from the substrate and placed in a sterile water droplet on a microscope slide, permitting five minutes of rehydration before further preparation. Thereafter microscopic characters were examined with a Nikon Eclipse 80i microscope with standard differential interference contrast (DIC) settings and with 10×, 20×, 40× and 100× objectives. The length and width of 20 spores was measured from three different fruiting bodies with an ocular micrometer using 100× oilimmersion objective. Images were taken with Nikon Digital Sight DS-Fi1 camera setup and measurements were calibrated with a stage micrometer.
For molecular characterization, five dry fruiting bodies were carefully removed from the leaf surface to avoid the inclusion of leaf material and potential contaminants. These were used for DNA extraction with the E.Z.N.A. High Performance spores, no isolate was obtained. Only an isolate of Ceratocystis paradoxa (deposited in GenBank as AY821864; CBS 116770) was frequently recovered during these attempts.
Of previously sequenced species, PUL F2679 shared the most sequence identity with H. pycnidioideum (= H. blepharistoma) at 92% (414 bp of 452 bp) in the LSU region and 86% (515 bp of 596 bp) in the ITS region. The only other close match was to a previously sequenced but undescribed isolate, CBS 196.95 (GenBank no. LSU-AY323905; ITS-AY323904), which shared 98% identity (446 bp of 453 bp) in the LSU region and 99% identity (609 bp of 616 bp) in the ITS region. Results of phylogenetic analyses are presented in Figure 1.
Type. Pycnopulvinus aurantiacus Toome & Aime Description. Minute, pigmented stilboid sporocarps with a swollen basal region and long tubular neck bearing a mucoid droplet of spores at the tip.
Ecology and distribution. On palm litter in South America (Ecuador); known from sequence data in Central America (Costa Rica). The analysis was performed with combined LSU and ITS sequence data and the topology was rooted with Rhodotorula yarrowii. The numbers above and below branches show the bootstrap and posterior probability values, respectively. LSU and ITS GenBank accession numbers of each used strain are given in the brackets. Superscript letter T indicates sequence data that originate from the type.
Discussion. The genus is closely related to Heterogastridium, but has orange-colored fruiting bodies that are larger and form a distinct basal cushion. Of the other three genera currently recognized in Heterogastridiales (Kirk et al. 2008, Pycnopulvinus can be readily separated from Colacogloea on the basis of DNA sequence data (Aime et al. 2006), and the absence of stilboid fructifications or septate spores in the latter, from Krieglsteinera which does not form fruiting bodies or multicelled spores and parasitizes ascocarps, and from Atractocolax which forms smaller (140 μm diam.), gelatinous, hyaline sporocarps, produces unicellular spores and is associated with bark beetles. Currently, Pycnopulvinus is a monotypic genus. A previously undescribed isolate, CBS 176.95 (isolated from the tropical forest of Costa Rica and accessioned in the GenBank as "Pycnobasidium sp." -a generic name that has not been validly published -appears to represent a member of this genus. In the absence of morphological or other additional data, it cannot be determined whether CBS 176.95 is conspecific with P. aurantiacus or represents a second species of Pycnopulvinus. Despite the rapid accumulation of environmental sequencing data, no studies thus far have published sequences referable to Pycnopulvinus, indicating that members of the genus are probably not widely dispersed in commonly sampled habitats. Figure 2 Diagnosis. Sporocarps minute, orange, with a swollen basal cushion (up to 3 mm wide) and long narrow neck (up to 3 mm long) subtended by a light yellow to orange mucous droplet (up to 1.5 mm diam.) of hyaline, 2-4 celled spores, averaging 3.25 × 11.8 μm. Found on palm leaf litter.
Description. Gregarious, light to dark orange (ca. 5A6-7), superficial, stilboid sporocarps with swollen globose base (0.5-3 mm wide in fresh specimen, drying to 0.2-1 mm), surrounded by hyphae with globular apical cells, 20-30 μm wide and 45-55 μm long. Sporocarp necks erect, long (0.5-2 mm), narrow (up to 110 μm at base, 50-70 μm at middle and widening up to 130 μm at tip), tubular, light yellow to orange, smooth. Hyphae on the outer layer of the neck 5 μm wide, septate; hyphae at the base and inside the neck 2-2.5 μm wide, septate. Ostiolar hyphae extend from the outer layer of the neck cells, hyaline, non-septate, 10-12 μm wide, narrowing at the tip. Clamp connections not observed. Spores accumulate in pale to orange mucous droplets at tips of sporocarp necks, approximately half of the size of the basal cushion in diam. (up to 1.5 mm diam. in fresh material). Spores hyaline, mostly 2-4 celled and cylindrical or somewhat fusiform, measuring 3-4 × 7-18 μm (average 3.25 × 11.8 μm), breaking into smaller compartments. No sporogenous cells were detected at the interior of the sporocarp, but the spores are likely asexually produced. Basidia not observed. Ecology and distribution. On palm leaf litter in tropical forest of Ecuador. Possibly occurring in association with other fungi. Known only from the type locale.
Etymology. aurantiacus = orange, for the color of fresh sporocarps. Specimens examined. PUL F2679. Discussion. The sporocarps most likely represent an asexual stage of P. aurantiacus. Anamorphic stilboid conidiomata have been described for other members of Pucciniomycotina, especially in Atractiellomycetes (e.g. Oberwinkler et al. 2006). However, multicellular spores have not previously been described for any sporocarpforming member of Atractiellomycetes or Microbotryomycetes. The organic matter of palm trees is not a common substrate among Pucciniomycotina and only one other basidiocarp-forming genus, Agaricostilbum Wright, is known to specifically inhabit palm litter (Wright et al. 1981). The recovery of C. paradoxa from the material might hint on a possible mycoparasitic strategy for P. aurantiacus.