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In this paper we present an updated checklist of thallus-forming Laboulbeniomycetes (Ascomycota, Pezizomycotina), that is, the orders Herpomycetales and Laboulbeniales, from Belgium and the Netherlands. Two species are newly described based on morphology, molecular data (ITS, LSU ribosomal DNA) and ecology (host association). These areHesperomyces halyziaeonHalyzia sedecimguttata(Coleoptera, Coccinellidae) from both countries andLaboulbenia quarantenaeonBembidion biguttatum(Coleoptera, Carabidae) from Belgium. In addition, nine new country records are presented. For Belgium:Laboulbenia aubryionAmara aranea(Coleoptera, Carabidae) andRhachomyces spinosusonSyntomus foveatus(Coleoptera, Carabidae). For the Netherlands:Chitonomyces melanurusonLaccophilus minutus(Coleoptera, Dytiscidae),Euphoriomyces agathidiionAgathidium laevigatum(Coleoptera, Leiodidae),Laboulbenia fasciculataonOmophron limbatum(Coleoptera, Carabidae),Laboulbenia metabletionSyntomus foveatusandS. truncatellus(Coleoptera, Carabidae),Laboulbenia pseudomaseionPterostichus melanarius(Coleoptera, Carabidae),Rhachomyces canariensisonTrechus obtusus(Coleoptera, Carabidae), andStigmatomyces hydrelliaeonHydrellia albilabris(Diptera, Ephydridae). Finally, an identification key to 140 species of thallus-forming Laboulbeniomycetes in Belgium and the Netherlands is provided. Based on the combined data, we are able to identify mutual gaps that need to be filled as well as weigh the impact of chosen strategies (fieldwork, museum collections) and techniques in these neighboring countries. The aim of this work is to serve as a reference for studying Laboulbeniomycetes fungi in Europe.

Laboulbeniales is one of the most peculiar orders of Ascomycota. These fungi are characterized by an ectoparasitic life-style on arthropods, determinate growth, lack of an asexual stage, high species richness, and intractability to culture. The order Laboulbeniales, sister to Pyxidiophorales, has only recently been assigned a separate class, the Laboulbeniomycetes, based on very few ribosomal DNA sequences. So far, DNA isolations and PCR amplifications have proven difficult. Here, we provide details of isolation techniques and the application of commercially available kits that enable efficient and reliable genetic analyses of these fungi. We provide 43 newly generated Laboulbeniales ribosomal DNA sequences, among which are the first published sequences for species in the genera Gloeandromyces, Herpomyces, Laboulbenia, Monoicomyces, and Polyandromyces. DNA extractions were possible using from 1 to 30 thalli from hosts preserved in ethanol (70–100 %). In two cases, we successfully isolated DNA from thalli on dried insect collections. Laboulbeniales molecular systematics could be substantially enhanced through these improved methods by allowing more complete sampling of both taxa and gene regions.

The ant-specific fungiAegeritella, Laboulbenia, Rickia, Hormiscium,andMyrmicinosporidiumin the Holarctic region—nine species—are reviewed. Present knowledge is highly biased geographically, as shows the single record for Holarctic Asia, and this is to solve. The phylogenetic position ofAegeritella, Hormiscium, andMyrmicinosporidiumis unknown. Hosts seem to be also skewed phylogenetically although this may be a true pattern.

Five species of the Laboulbeniales, including two unrecorded species are reported from South Korea. They are as follows; Dioicomyces anthici Thaxter on Anthicus confucii Marseul, Laboulbenia melanaria Thaxter on Anisodacthius tricuspidatus Morawitz, L. philonthi Thaxter on Philonthus wuesthoffi Bernhauer, Peyritschiella japonicus Terada on Philonthus japonicus Sharp, and Scaphidiomyces baeocerae Thaxter on Scaphisoma unicolor Achard. Among these species, L. melanaria Thaxter, S. baeocerae Thaxter and the male thallus of D. anthici Thaxter are newly described from South Korea. L. Philonthi Thaxter and P. japonicus Terada are newly collected in some places where were unlike with the examined region ago.

A new parasitic fungus, Laboulbenia curtonoti sp. n., associated with the endemic Madagascan fly Curtonotum balachowskyi Tsacas (Diptera: Curtonotidae), is described and figured. The new species is one of the very few Laboulbeniales which penetrate the insect's cuticle by means of more-or-less developed rhizoids. A brief review of the 23 species of the genus Laboulbenia associated with Diptera is presented in a tabulated form.

The myrmecophilous fungi Rickia wasmannii Cavara, 1899 and Laboulbenia camponoti S. W. T. Batra, 1963 (Ascomycota: Laboulbeniales) are reported for the first time from Bulgaria. Rickia wasmannii was found on Myrmica scabrinodis Nylander, 1846 ant workers (Hymenoptera: Formicidae) in South-eastern Bulgaria near to the Black Sea coast. This is the easternmost record of Rickia wasmannii in Europe. Laboulbenia camponoti was found in six different localities in Bulgaria on the carpenter ants Camponotus aethiops (Latreille, 1798), C. universitatis Forel, 1890 and C. pilicornis (Roger, 1859). Camponotus aethiops and C. universitatis are new hosts for the fungus. For both fungi species the known distribution and host ranges summarized. This is the first record of the ant species Camponotus pilicornis for the Bulgarian fauna.

Twenty-nine Laboulbeniales are reported for the first time from Greece. These are Aphanandromyces audisioi W. Rossi, Botryandromyces heteroceri (Maire) I. I. Tavares et Majewski, Cantharomyces numidicus Maire, Cantharomyces orientalis Spegazzini, Chitonomyces bakeri Thaxter, Cryptandromyces elegans (Maire) W. Rossi et Castaldo comb. nov., Cucujomyces rotundatus Majewski, Dioicomyces anthici Thaxter, D. italicus Spegazzini, Dimorphomyces myrmedoniae Thaxter, Hesperomyces virescens Thaxter, Hydrophilomyces coneglianensis Spegazzini, H. gracilis Majewski, Idiomyces peyritschii Thaxter, Laboulbenia cristata Thaxter, L. flagellata Peyritsch, L. gyrinicola Spegazzini, L. inflata Thaxter, L. philonthi Thaxter, L. pseudomasei Thaxter, L. stilicicola Spegazzini, Monoicomyces britannicus Thaxter, M. homalotae Thaxter, Rhadinomyces pallidus Thaxter, Rickia peyerimhoffii Maire, Smeringomyces anomalus (Thaxter) Thaxter, Stichomyces conosomatis Thaxter, Teratomyces mirificus Thaxter, Zodiomyces vorticellarius Thaxter. ...

Specimens of the type collection of Laboulbenia polymorpha Sugiyama (1978) were reexamined. The holotype, M. Ishikawa 673, is composed of three slides and includes three morphologically different forms of thalli, of which two forms were illustrated (Sugiyama 1978, fig. 1-D, 1-E). On the other hand, one of the paratypes, M. Ishikawa 674, has now been lost but photographs were made earlier from this slide, in which one mature individual illustrated as fig. 1-C (Sugiyama 1978) is included. This individual was not correctly shown in Sugiyama’s illustration, but actually has a strong resemblance to a form in slide 673-b that was not illustrated by Sugiyama (1978). Thus, three different forms have been recognized as variations of L. polymorpha. In the present article, each variation was termed C-form, D-form, and E-form because Sugiyama (1978) used the same notation in his figures. A mature specimen of C-form in slide 673-b has been selected as a lectotype. Slide 673-d includes only young thalli, one of which was illustrated as fig. 1-G (Sugiyama 1978). This young thallus undoubtedly belongs to another species; mature thalli of the same species were also found in slide 673-b. Another paratype, K. Sugiyama 2101, includes C- and D-forms of L. polymorpha. Infection sites of the C- and D-forms have been determined: the C-form grows mainly on the lateral margins of the elytra of the host, and the D-form occurs mainly on the basal part of the elytra and the mesothorax.

Fungi of the order Laboulbeniales are obligate ectoparasites that are easy to detect on the integument of their arthropod hosts. This apparency, and other characteristics that suit them for the role of ‘model’ group in the exploration of parasite biodiversity patterns, are discussed. Salient features of the general biology of these organisms are summarized and past patterns of species description reviewed. In recent years, much as in the fungi as a whole, roughly equal proportions of new species have been described from tropical and temperate regions. Comparison of the known Laboulbeniales mycobiotas of Europe, Asia and ‘tropical Asia’ reveals that most of the more significant parasite genera are more or less equally well represented in each region. Patterns of host utilization are also largely similar, with more than 50% of host species recorded from each region belonging to the two beetle families Carabidae and Staphylinidae. However, there are indications that parasite species that are not confined to a single host species have significantly broader geographical ranges than any of their hosts. The pattern of host utilization in the relatively fully-inventoried assemblage of beetles of a tropical forest in North Sulawesi is described, in terms of the taxonomic group membership, habitat affiliation and feeding biology of recorded hosts. On the basis of the Sulawesi dataset and other available data on host exploitation patterns at various spatial scales, we suggest that between 2 and 5%, i.e. between ca 40000 and ca 100000 of an estimated 2 million extant beetle species, are hosts of one or more species of Laboulbeniales. Known and predicted levels of host specificity imply that a global figure for Laboulbeniales species exploiting Coleoptera is likely to be between 10000 and 50000 species. A smaller number of Laboulbeniales species, probably no more than half the number associated with beetles, are to be found on other arthropod hosts. Ways in which these estimates might be refined are discussed.

Laboulbenia ophioneae is described as a new species with illustrations. It is closely related to Laboulbenia celestialis and Laboulbenia asiatica because of some similarities in the morphology of appendages and perithecia. The present species from the latter two species can be distinguished by the shorter, inflated perithecia, the shorter receptacles, and the appendages consisting of more or less darkly colored, broader branches. This new species was found on elytra of Ophionea indica from Taiwan, which has been also known as a host of Laboulbenia polymorpha. A mature thallus noticed by Terada (2004) on the slide 673b (M. Ishikawa collection) and a young thallus illustrated by Sugiyama (1978, fig. 1-G as L. polymorpha) on 673d (M. Ishikawa collection) are both identified as L. ophioneae.

Four new species of Laboulbeniales occurring on beetles (Coleoptera) from Sulawesi (Celebes), Indonesia, are described and illustrated: Cucujomyces celebensis sp.nov. parasitic on Hyliota sp. (Cucujoidea, Cucujidae), Dimeromyces storkii sp.nov. parasitic on Eustra matanga Andrewes and Eustra plagiata Schm.-Goebel (Carabidae, Paussinae), Laboulbenia hammondii sp.nov. parasitic on Omadius spp. (Cleridae), and Rickia anthribidicola sp.nov. parasitic on Hypseus vestitus Jordan (Anthribidae, Anthribinae). All of the new species are compared with existing taxa. Of particular note is R. anthribidicola, which represents the first confirmed instance of parasitization by a laboulbenialean fungus on members of the hyperdiverse beetle family Anthribidae. Keywords: Laboulbeniales, Indonesia, Anthribidae.

The type-species of four genera of Pseudococcidae are studied. Fonscolombia Lichtenstein is redefined, the type-species, Fonscolombia graminis Lichtenstein is assigned to the Pseudococcidae, redescribed and a lectotype designated. Laboulbenia Lichtenstein is accepted as an objective synonym of Antonina Signoret. Tetrura Lichtenstein (non: Lesson) is accepted as an objective synonym of Coccura Sulc. The adult female of the material of this species which Lichtenstein misidentified as Tetrura rubi (Schrank) is redescribed and illustrated. Brevennia Goux is redefined, and the type-species Ripersia (Brevennia) tetrapora Goux redescribed. The characteristics of Asphodelococcus Morrison are discussed, the type-species Ripersia asphodeli Bodenheimer redescribed and a neotype designated; A.meoconcitae Tranfaglia & Marotta is a new synonym of A. asphodeli.

Laboulbeniales (Ascomycetes) new for Italy. In the present paper are quoted 19 species of Laboulbeniales not jet reported for Italian flora: Asaphomyces cholevae Th., Corethromyces henroti Balazuc, C. pallidus (Th.), C. stilici Th., Euzodiomyces lathrobii Th., Helodiomyces elegans Picard, Laboulbenia acupalpi Speg., L. inflata Th., L. philonthi Th., L. picardi Maire, Misgomyces dyschirii Th., M. lavagnei Picard, Monoicomyces californicus (Th.), Peyritschiella protea Th., Rhachomyces lasiophorus (Th.), R. furcatus (Th.), R. philonthinus Th., R. pilosellus (Robin). Also the genera Asaphomyces Th., Euzodiomyces Th., Helodiomyces Picard and Peyritschiella Th. are new for Italy. All the quoted species are parasites of Insects of the order of the Coleoptera.

Three new Laboulbeniales occurring on Carabidae (ground beetles) are described: Laboulbenia sciakyi, parasitic on Planetes elegans Nietner (Galeritinae, Planetini) from Sri Lanka, Rhachomyces basilewskyi, parasitic on Eunostus burgeoni Alluaud (Galeritinae, Galeritini) from Zaire, and Rhachomyces leleupidiae, parasitic on Leleupidia kahuziana Basilewsky (Zuphiinae, Leleupidiini) from Zaïre.

Two new Laboulbeniales occurring on African Zuphiini are described: Eucantharomyces basilewskyi, parasitic on Zuphium spp., and Laboulbenia cajanii, parasitic on Paraleleupidia uvirana Basilw.

The Laboulbeniales are predominantly parasites of true insects (Hexapoda), infesting mostly Coleoptera. Two members of the Laboulbeniales are reported from South Africa for the first time; Polyandromyces coptosomalis Thaxter and Laboulbenia monoleptae Balazuc. Both fungi are illustrated and their distribution, morphology, host specificity and nutrition are discussed.