Laboulbeniomycetes (Fungi, Ascomycota) of Denmark Author Santamaria, Sergi Unitat de Botànica, Departament de Biologia Animal, de Biologia Vegetal i d’Ecologia, Facultat de Biociències, Universitat Autònoma de Barcelona, 08193 Cerdanyola del Vallès (Barcelona), Spain. sergi.santamaria@uab.cat Author Pedersen, Jan Natural History Museum of Denmark (Zoological Museum), University of Copenhagen, Universitetsparken 15, 2100 København Ø, Denmark. japedersen@snm.ku.dk text European Journal of Taxonomy 2021 2021-11-26 781 1 425 http://dx.doi.org/10.5852/ejt.2021.781.1583 journal article 10.5852/ejt.2021.781.1583 2118-9773 5828924 Key to the Danish genera of thallus-forming Laboulbeniomycetes 1. On cockroaches ( Blattodea ). Dioecious. In female thalli, cell II gives rise to a secondary axis which forms perithecia and connects directly with host integument by means of a series of narrow cells perforating the integument directly by small haustoria; some of these cells may differentiate into shield-like structures................................. Herpomycetales Haelew. & Pfister. Herpomyces Thaxt. – On other arthropods (species on Blattodea in this section are unknown in Europe). Dioecious or monoecious. Thalli not as above. Typically attached to the host integument by means of a single cell or foot (usually black) developed by transformation of the ascospore basal cell. If not a blackened foot, a conspicuous haustorium is present (except for Laboulbenia hyalopoda De Kesel ) ................................................................................................ 2 ( Laboulbeniales Lindau ) 2. Outer wall of perithecium consisting of more than eight equal or subequal superposed cells for each vertical row. (See also the genus Compsomyces . ) ............................................................................ 3 – Outer wall of perithecium consisting of 4–5(–8) unequal superposed cells for each vertical row........................................................................................................... 9 ( Laboulbeniaceae Peyr. ) 3. Axis of receptacle consisting of a series of superposed cells. Most species on aquatic Coleoptera .......................................................................................... 4 ( Ceratomycetaceae S.Colla ) – Receptacle massive, multiseriate. On terrestrial Coleoptera Staphylinidae of the genera Lathrobium Gravenhorst, 1802 and Lobrathium Mulsant & Rey, 1878 (less frequently on other genera of Coleoptera as Patrobus , a carabid)..................................... Euceratomycetaceae I.I.Tav. Euzodiomyces Thaxt. 4. Receptacle consisting of a series of superposed cells, those in the lower part distinctly broadened. On Coleoptera Hydraenidae .................................... Drepanomyceteae I.I.Tav. Thripomyces Speg. – Cells of the lower receptacle similar to each other................................. 5 ( Ceratomyceteae Thaxt. ) 5. Perithecial apex bearing four slender and rather long ligules surrounding the ostiole. On Coleoptera Dryopidae .......................................................... Helodiomycetinae I.I.Tav. Helodiomyces F.Picard – Perithecial apex simple or with subapical horns. On Coleoptera Hydrophilidae ................................ .............................................................................................................. 6 ( Ceratomycetinae Thaxt.) 6. Perithecium with a very long slender neck and an inflated venter ....... Rhynchophoromyces Thaxt. – Perithecium with indistinct neck and venter..................................................................................... 7 7. Outer wall cells similar in height in all vertical rows....................................... Autoicomyces Thaxt. – Two vertical rows of flattened cells alternating with narrow elongated cells in the other two rows... .......................................................................................................................................................... 8 8. Normal forms with a multicellular (more than five cells) lower receptacle. Bifurcate horns on perithecium. Thalli from claws without horns.............................................. Eusynaptomyces Thaxt. – Lower receptacle consisting of 2–4(–5) superposed cells. Simple horns on perithecium................... ............................................................................................................................ Ceratomyces Thaxt. 9. Receptacle massive, multiseriate. Perithecia ± hidden, immersed in a cavity of the thallus upper region ......................................................... Zodiomycetoideae (Thaxt.) I.I.Tav. Zodiomyces Thaxt. – Receptacle different. Perithecia obvious, well visible .................................................................... 10 10. Antheridia unicellular, simple, consisting of a phialide; or not evident or unknown (supposedly some taxa may be apomictic)........................................................................ 19 ( Laboulbenioideae Peyr.) – Antheridia multicellular, compound, consisting of several fertile cells or phialides located inside a common cavity. Spermatia are ejected through a pore of that cavity.............................................. 11 11. Secondary receptacles giving rise to antheridia and perithecia.Antheridia consisting of 3–4 tiers of paired cells, where the median tier or 2 nd + 3 rd tiers give rise to 2–8 phialides. Spermatia are ejected through a duct between the upper tier cells... Monoicomycetoideae (Thaxt.) I.I.Tav. Monoicomyces Thaxt. – Otherwise.......................................................................... 12 ( Peyritschielloideae (Thaxt.) I.I.Tav.) 12. Dioecious. Cells VII and basal cells of perithecium not visible at maturity ....................................... ........................................................................................................ 13 ( Dimorphomyceteae Thaxt. ) – Monoecious. Cells VII and basal cells of perithecium well-distinguished at maturity .................. 15 13. Neither secondary appendages nor blackened foot. Haustorium well-developed. Mostly on Mallophaga (bird lice) ......................................................................................... Trenomyces Chatton & F.Picard – Secondary appendages and normal blackened foot present ........................................................... 14 14. Female thalli with a laterally elongated cell I, underlying a series of cells bearing both appendages and perithecia................................................................................................ Dimorphomyces Thaxt. – Female thalli without a laterally elongate cell I supporting cells which give rise to lateral perithecia and appendages.................................................................................................. Dimeromyces Thaxt. 15. Receptacle flattened and multiseriate. Antheridia flask-shaped ............ 16 ( Peyritschielleae Thaxt. ) – Receptacle otherwise, with superposed and contiguous cells I, II and III........................................... ............................................................................................................... 17 ( Haplomyceteae Thaxt. ) 16. Receptacle consisting of a basal cell with three superposed tiers of horizontally arranged cells.................................................................................................................... Peyritschiella Thaxt. – Receptacle consisting of a basal cell and 2–3 vertical series of superposed cells....... Rickia Cavara 17. Compound antheridium in the basal cell of primary appendage which continues above by simple or ramified sterile branches ................................................................................ Cantharomyces Thaxt. – Compound antheridium terminal on primary appendage ............................................................... 18 18. Compound antheridium terminated with an efferent neck................................. Camptomyces Thaxt. – Compound antheridium spinose, pore for release of spermatia in a sublateral position ..................... ............................................................................................................................. Haplomyces Thaxt. 19. Perithecium terminal on a uniseriate receptacle consisting of cells I, II, III and a few flattened cells. Primary appendage disappearing when perithecium develops. On Heteroptera Corixidae ................ ....................................................................................... Coreomyceteae Thaxt. Coreomyces Thaxt. – Perithecium lateral, arising from cell II or its derivatives. Primary appendage present. On other hosts ................................................................................................................................................ 20 20. Primary appendage is a continuation of the same axis of the primary receptacle.Antheridia unknown. Perithecial outer wall consisting of four vertical rows each with four cells of unequal height. On Col. Staphylinidae Pselaphinae of the genus Pselaphus Herbst, 1792 ........................................................ ....................................................................................................... Tanmaurkiella Santam. gen. nov. – Primary appendage seen as a lateral branch of the thallus ............................................................. 21 21. Perithecial vertical wall cell rows consisting of more than five cells, ± equal in height..................... ..................................................................................................... 22 ( Compsomyceteae Thaxt. p.p.) – Perithecial vertical wall cell rows consisting of three, four or five cells, unequal in height .......... 23 22. Receptacle multiseriate, with the cell II subdivided by horizontal and vertical septa. Perithecia supported on uniseriate multicellular stalks, with an apical rostrum and a subterminal outgrowth.... .................................................................................... Kainomycetinae I.I.Tav. Kainomyces Thaxt. – Receptacle uniseriate, two-celled. Perithecium pedicel consisting of the stalk cell (cell VI) and an additional underlying cell which may be regarded as part of the secondary receptacle. Neither rostrum nor outgrowths on perithecium. On Astenus Dejean, 1833 (Col. Staphylinidae )................................ ................... Compsomycetinae Thaxt. Compsomyces Thaxt. p.p. ( C. verticillatus (Thaxt.) Thaxt. ) 23. Perithecium borne on secondary axis. On Lesteva Latreille, 1797 (Col. Staphylinidae ) .................... ............................................................ Compsomycetinae . Compsomyces p.p. ( C. lestevae Thaxt. ) – Perithecia borne on primary axis .................................................................................................... 24 24. One elongate accessory cell on the outer side of the perithecial venter wall (study of immature thalli may be necessary). Only on aquatic beetles ........................................................................................ .................................................................... Hydrophilomyceteae I.I.Tav. Hydrophilomyces Thaxt. – Not such elongate accessory cell on the perithecium ..................................................................... 25 25. Cell wall of cell VII and at least one of the perithecial basal cells (m, n, n’) well distinguished at maturity........................................................................................................................................... 28 – Cell wall of cell VII and perithecial basal cells (m, n, n’) vanishing, not visible at maturity ............. ........................................................................................................ 26 ( Euphoriomyceteae I.I.Tav.) 26. Primary receptacle, below cell VI, two-celled (i.e., cells I and II).................... Phaulomyces Thaxt. – Primary receptacle, below cell VI, consisting of more than two cells, by subdivision of cell II ... 27 27. Neither branches nor antheridia below the perithecium. Receptacle uniseriate.................................. ..................................................................................................... Siemaszkoa I.I.Tav. & T.Majewski – Secondary appendages and antheridia below perithecia. Receptacle looking multiseriate................. ....................................................................................................................... Euphoriomyces Thaxt. 28. Receptacle multicelled, ± multiseriate or corticated, with cells II and III horizontally and vertically divided, developing a cellular complex bearing lateral or terminal abundant crowded branchlets............................................................................................... 29 ( Teratomyceteae Thaxt. ) – Receptacle multicelled, uniseriate, or only with few vertical septa in the distal end .......................... ................................................................................................................ 34 ( Laboulbenieae Thaxt. ) 29. Receptacle broadened distally into a cellular complex bearing numerous lateral and/or terminal crowded appendages which in turn bear intercalary seriated antheridia............................................. ............................................................................................................. 30 ( Teratomycetinae Thaxt.) – Receptacle seemingly uniaxial or corticated, the latter with a conspicuous haustorium. Antheridia terminal ........................................................................................................................................... 32 30. Receptacle almost symmetrical, four-layered, with cell II subdivided by horizontal septa into two superposed cells (IIa and IIb), and an uppermost fourth tier where cell III is divided repeatedly by vertical and oblique septa ............................................................................................................... 31 – Receptacle clearly asymmetric, unilateral, with cell II subdivided into three cells (IIa, IIa’ and IIb), IIb and IIa’ separated by a vertical septum. On Deleaster Erichson, 1939 (Col. Staphylinidae )......................................................................................................... Idiomyces Thaxt. 31. Appendages with pointed branchlets ................................................................... Teratomyces Thaxt. – Appendages without pointed branchlets...................................................... Symplectromyces Thaxt. 32. Blackened foot absent. Haustorium conspicuous. On Col. Silvanidae ............. Microsomyces Thaxt. – Blackened foot present. Lacking a noticeable haustorium ............................................................. 33 33. Thallus consisting of a primary bicellular receptacle (cells I and II) and a secondary receptacle formed by division of cell II, which represents the main axis, where lateral cells give rise to appendages. All perithecial basal cells (m, n, and n’) well-distinguished ..................................................................... ................................................................ Rhachomycetinae (Thaxt.) I.I.Tav. Rhachomyces Thaxt. – Thallus only with one primary axis. Of the perithecial basal cells, only cell m is visible. On Col. Leiodidae .............................................................. Asaphomycetinae I.I. Tav. Asaphomyces Thaxt. 34. Thalli dioecious. Male thalli consisting of 3–4 superposed cells, the uppermost cell functioning as a phialide. Female thalli with a uni- or bicellular primary appendage, sometimes embedded among thallial base cells and externally invisible ....................... 35 ( Amorphomycetinae (Thaxt.) I.I.Tav.) – Thalli monoecious, or if dioecious, female and male thalli different from above.......................... 36 35. Female thalli consisting of a three-celled receptacle, cell III supporting a unicellular primary appendage. Male thalli four-celled, the uppermost cell functioning as a phialide .............................. ............................................................................................................................ Dioicomyces Thaxt. – Female thalli consisting of a two-celled receptacle, with cells II–III not separated; primary appendage usually indistinguishable. Male thalli three-celled, the uppermost cell functioning as a phialide ........................................................................................................... Amorphomyces Thaxt. 36. Antheridia below the primary septum or the primary appendage consisting of a campanulate basal cell and a filiform distal cell. Only on aquatic beetles.................... 37 ( Chitonomycetinae I.I.Tav.). – Antheridia on a well-developed primary appendage, borne above the primary septum ................ 38 37. Perithecial outer wall consisting of four vertical rows, each with 3–4 cells of unequal height .......... ............................................................................................................................ Chitonomyces Peyr. – Perithecial outer wall consisting of four vertical rows of cells organized as two adjacent rows with 4–5 cells (formed from n) and the other two rows with 7–8 cells................... Hydraeomyces Thaxt. 38. Lower receptacle two-celled (below the perithecial stalk cell), consisting of superposed cells I and II. An inner sterile cell is formed from the appendage axis cell just above the perithecium (usually the third cell if only one perithecium is developed), as a corner cell. On Chaetarthria Stephens, 1833 (Col. Hydrophilidae )............................ Chaetarthriomycetinae I.I.Tav. Chaetarthriomyces Thaxt. – Lower receptacle (below the perithecial stalk cell) consisting of three or more cells, in a uniseriate row (by subdivision of cell II). If two-celled, characteristics are different from above. ................ 39 39. Lower receptacle (below the perithecial stalk cell) three- to multicelled....................................... 40 – Lower receptacle (below the perithecial stalk cell) two-celled (i.e., cells I and II)........................ 43 40. Antheridial cells embedded in an irregular mass of cells above the primary septum, bearing short efferent necks .................................................................................................................................. 41 – Antheridial cells are free phialides (i.e., flask shaped cells), usually on appendage branches....... 42 41. Cell III subdivided irregularly into several small cells. Perithecium with four outer wall cells of unequal height in each vertical row.............. Misgomycetinae (Thaxt.) I.I.Tav. Misgomyces Thaxt. – Cell III subdivided into three cells, by a horizontal septum and the upper cell divided again by a vertical septum. Perithecium with three outer wall cells in two adjacent vertical rows and four in other two rows. On Col. Heteroceridae ............................................................................................... ............................................. Laboulbeniinae Thaxt. p.p. Botryandromyces I.I.Tav. & T.Majewski 42. Antheridia are elongate phialides arranged in clusters supported by short branchlets arising from corner cells in lower receptacle. On Col. Heteroceridae ..... Laboulbeniinae p.p. Botryandromyces – Antheridia sessile or on short stalk cells in lower part of the primary appendage. On Col. Ptiliidae ........................................................................... Laboulbeniinae p.p. Ecteinomyces Thaxt. 43. Receptacle usually five-celled, with a two-celled lower receptacle (below perithecial stalk cell) and a three-celled upper receptacle formed by subdivision of the terminal cell (cell III) into three cells (namely III, IV and V). Perithecium with four outer wall cells in each vertical row, the cells distinctly unequal in height. Lowermost cell of primary appendage is the insertion cell, typically flat and usually blackened ..................................................... Laboulbeniinae p.p. Laboulbenia Mont. & C.P.Robin – Receptacle three-celled ..................................................... 44 ( Stigmatomycetinae (Thaxt.) I.I.Tav.) 44. Perithecium with five outer wall cells in each vertical row, the cells equal or slightly unequal in height, but the two cells of the venter somewhat longer. Primary appendage usually unbranched, bearing lateral sessile or subsessile antheridia, or the cells functioning wholly as phialides.Mostly on Col. Staphylinidae Pselaphinae , also in Col. Cryptophagidae (so far only in Denmark !) ........ Cryptandromyces Thaxt. – Perithecium with outer wall cells distinctly unequal in height (cells of the venter twice or more longer than any of the apical cells, especially the uppermost ones in the series) ...................................... 45 45. Cell III supporting an unbranched, free appendage, consisting of two superposed cells and a terminal, spinose, flask-shaped phialide. On Col. Staphylinidae Pselaphinae ............................. Bordea Maire – Appendage with different characters as above ............................................................................... 46 46. Primary appendage consisting of three superposed cells, with the median and upper cells giving rise to antheridia, the upper cell bearing a dorsal spinous process. On Cryptophagus Herbst, 1863 (Col. Cryptophagidae ) (see also Cryptandromyces cryptophagi Santam. sp. nov. ) ..................................... ...................................................................................................................... Rossiomyces R.K.Benj. – Otherwise........................................................................................................................................ 47 47. Cell III narrow and adnate to the side of the perithecium, reaching one half or beyond its height. On Diplopoda Julida (millipedes)............................................................................ Troglomyces S.Colla – With other characters. On insects ................................................................................................... 48 48. Primary appendage bent in sharp angle towards the perithecium, consisting of two superposed cells and a terminal, spinose phialide. Perithecium bearing two apical outgrowths. On Dermaptera Forficulidae (earwigs)........................................................................................ Distolomyces Thaxt. – On other insects and with other characters ..................................................................................... 49 49. Primary appendage consisting of a short branch which on the inner side of its basal cell produces an appendage giving rise many short lateral branches. On Heteroptera Hebridae (true bugs) ............................................................................................................ Tavaresiella T.Majewski – On other insects and with other characters ..................................................................................... 50 50. On Diptera ...................................................................................................................................... 51 – On Coleoptera ................................................................................................................................. 52 51. Cell I inflated, without a blackened foot but with a conspicuous haustorium. Cell II flattened. Cell III free. On Nycteribiidae (parasitic flies living on bats)................................... Arthrorhynchus Kolen. – Cells I and II elongate, cell III laterally adnate to cell VI ........................... Stigmatomyces H.Karst. 52. Primary appendage limited in growth, compact, terminating with antheridia ............................... 53 – Primary appendage not limited in growth, ± ramified and terminating with a sterile branch ........ 56 53. Primary appendage caducous by breaking off the collapsing third cell which supports two cells and four antheridia. On Brachypterus Kugelann, 1794 (Col. Kateretidae ).. Aphanandromyces W.Rossi – Primary appendage perennial ......................................................................................................... 54 54. Cell III in contact with cell I........................................................................................................... 55 – Cell III not in contact with cell I. On Col. Staphylinidae Paederinae ............ Sphaleromyces Thaxt. 55. Perithecium with five outer wall cells in each vertical row. Trichogyne V-shaped, with arms covered by globoid papillae ......................................................................................... Acompsomyces Thaxt. – Perithecium with four outer wall cells in each vertical row, bearing apical lobes. Trichogyne different. Mostly on Col. Coccinellidae .......................................................................... Hesperomyces Thaxt. 56. Branchlets bearing antheridia arising from corner cells of appendage axial cells.......................... 57 – Branchlets bearing antheridia not arising from corner cells........................................................... 58 57. Cell III always bearing antheridial branchlets, rarely on appendage cells above. On Lathrobium (Col. Staphylinidae )................................................................................................... Rhadinomyces Thaxt. – Cell III sterile, rarely bearing antheridial branchlets, which are abundant on appendage cells above. On Sepedophilus Gistel, 1856 (Col. Staphylinidae ) ............................................ Stichomyces Thaxt. 58. Cell II adnate or parallel to cell VI. Mostly on Col. Leiodidae (less commonly on Col. Staphylinidae )..................................................................................................... Diphymyces I.I.Tav. – Cell II supporting the free cell VI. Mostly on Col. Staphylinidae .................. Corethromyces Thaxt.