Holwayaceae Quijada, Matočec & I. Kušan, in Quijada, Matočec, Kušan, Tanney, Johnston, Mešić & Pfister, Biology 11(4, no. 583): 13 (2022)

Index Fungorum number: IF 842621, MycoBank number: MB 842621, Facesoffungi number: FoF 15026

Type genus – Holwaya Sacc., Syll. fung. (Abellini) 8: 646 (1889)

Etymology – named after the type genus Holwaya.

Sexual morph: Ascoma apothecioid, paragymnohymenial or eugymnohymenial, superficial or erumpent, scattered to gregarious, solitary or sharing a common stroma-like base; pulvinate-turbinate to discoid or cup-shaped, pale yellowish to black, hymenium margin and receptacle concolorous, margin indistinct, thin and smooth, disc ± circular when fresh, sometimes slightly irregular by mutual pressure; sessile to short- to rather long-stipitate, stipe tapering downward; surfaces smooth or slightly rugulose due to the protrusion of paraphyses tips in the hymenium or protruding cells in the ectal excipulum. Asci cylindric-clavate, 4–8-spored, (1–)2–4-seriate; apex dome-shaped with a subapical OLR (best visible by the DIC of *ripe unopened and in ↑asci) with its thinner, lateral wall staining in CR but subapical ring and walls at apex unstained, slightly subconical and with moderately to strongly swollen wall in dead state, apex wall also partly laterally thickened; wall IKI and MLZ negative; base narrowed to a short to medium-long stalk and arising from croziers. Ascospores ellipsoid-ovoid-fusoid to cylindrical-fusiform-acicular, hyaline, straight or slightly to moderately curved, smooth to ornamented with coarse ridges or crests that can form a reticulum (KOH stable), eguttulate or multiguttulate (tiny oil drops); aseptate to (10–)16–20 transversal septa when septate and overmature with a small lateral wart-like to rod-shaped protuberance at each cell, on which one or two small conidia are formed. Paraphyses filiform, apically moderately to strongly clavate-capitate, exceeding living mature asci, sparsely branched at lower cells, ±equidistantly and sparingly septate; apical cells hyaline or partly olive-brown, pigmented with or without refractive globules, wall surface smooth to finely warted; cells apically free or agglutinated and embedded in a brownish gelatinous matrix and exudate. Subhymenium composed of compact ascogenous cells. Medullary excipulum very reduced and sometimes undifferentiated from ectal excipulum; if differentiated, then composed of dense textura intricata, slightly gelatinized, of thin-walled cells with a ±vertical orientation to the subhymenium. Ectal excipulum of ±textura globosa–angularis to t. prismatica, cells hyaline to olive-brown; cortical layer with groups of angular to clavate cells slightly protruding, thick-walled, covered by a dark, brownish exudate. Anchoring hyphae at base present or not. Asexual morph: Only known for Holwaya.

Specimens examined – Holwaya mucida—CANADA: State and province not recorded, on the bark of Tilia sp., 3 October 1896, J. Macoun (CUP-D-02006).—CROATIA: Primorje – Gorski kotar County, Nadvučnik village, near town Vrbovsko, the vegetation of young Carpinus betulus with Pinus nigra and Picea abies, on cut, semi-decorticated lying trunks of Tilia sp. covered with moss, 6 November 2010, R. Kranjčev, N. Matočec and I. Kušan (CNF 2/8749).—CZECH REPUBLIC: South Bohemian Region, NE of Hluboká nad Vltavou, Libochovka Nature Reserve, a forest of Fagus sylvatica with admixed Picea abies and Tilia sp., on a fallen trunk of Tilia ap. covered with mosses, 16 October 2016, J. Holec (PRM 944637).—DENMARK: Sjælland Region, Ringsted Municipality, Allindemagle Skov, forest with Tilia sp., on Tilia sp., 20 September 2011, M. Vestergaard (det. Thomas Læssøe) (C-F-91657)—SLOVENIA: Gorizia Region, Idrija, 370 m S-SE from the Kres peak (521 m), urban alley of Tilia sp. trees, on a rotten lying trunk of Tilia sp., 23 November 2001, A. Piltaver and N. Matočec (CNF 2/5423).—SWEDEN: Västmanland County, Västerås-Barkarö parish, Flaten island near Ridön island, on a fallen trunk of Tilia cordata, 20 October 1975, S. Ryman (PRM 869874).—USA: New Hampshire State, Carroll County, Chocorua, on Acer sp., September 1910, W.G. Farlow (PRM 685875, as H. gigantea), idem (PRM 685876, as H. gigantea); New York State, Slaterville Springs, Lloyd-Cornell Preserve, on a rotten log of Tilia sp., 25 September 1954, R.P.Korf (PRM 919564); idem, Coy Glen, Ithaca, on unidentified wood, 2 October 1982, T. Capiello and N. Shishkoff (CUP-60122); idem, Tompkins, Enfield, on unidentified wood, 14 October 1905, G. Atkinson (CUP-A-019509); Massachusetts, Concord, Estabrook Woods, on a fallen trunk of Populus sp., 3 Feburary 2019, Luis Quijada (L.Q.H.-102 in FH). Ramgea ozimecii—CROATIA: Lika-Senj County, near Perušić, about 2.6 km E-SE of Gornji Kosinj, pit Čardačina jama, in the dark zone of the karstic pit, on bat dropping, 30 September 2016, R. Ozimec (CNF 2/9997, holotype). Patinella hyalophaea—BOSNIA AND HERZEGOVINA: Sutjeska National Park, riverine canyon bottom, wet dicot woody remnant, 28 June 2015, N. Omerović (FAMU-1390, N.O.280615-03); Sarajevo County, Bijambare Protected Landscape, Brodić stream, periodical karstic watercourse in alti-montane conifer forest, Picea abies wet remnant, 4 August 2014, N. Omerović (no voucher).

Note – Phylogenetically isolated within Thelebolales. Ascomata open; outer layer of the excipulum with globose to angular cells; capitate paraphyses with intra- or extracellular pigments and asci with forcible spore discharge augmented by an opening limitation ring-type apical apparatus; ascospores hyaline, smooth, or ornamented; either aseptate and ellipsoid-ovoid-fusoid, not producing conidia, or with more than ten septa, acicular-cylindrical-fusiform-acicular, and producing conidia. Ramgea differs from apothecioid members of Thelebolaceae by ornamented ascospores and paraphyses with capitate tips bearing granular, firmly cemented pigment exudates. Patinella and Holwaya differ from Tympanis and Durandiella (Tympanidaceae) and Aotearoamyces and Bulgaria (Phacidiaceae) in excipular composition of mostly globose-angular cells in combination with non-amyloid asci and ascospores.

Figure 1. Phenotypic features of Holwayaceae: Holwaya (A1–6,B1–3,C1–6,D1–3,E1–2), Patinella (F1–6,G,H,I,J1–3) [84], and Ramgea (K,L1–2,M1–4,N1–2,O1–3) [64]. Comparative morphological features among genera in the family: A, F, and K. Macromorphology of the apothecia: B, J, and N. Excipulum in transverse section: C, G, and M. Asci and details of ascus apex: D, H, and L. Paraphyses: E, I, and O. Ascospores. A1, A3 from CNF 2/8749; A2, A4–6 from L.Q.H.-102; B1–3, C3–4, D2–E2 from CUP-60122; C1–2, C6, D1 from CUP-A-019509; C5 from CUP-D-02006. A1, A3 phot. N. Matočec, A2, A4–6 phot. J. Warfel, B1–E2 phot. Luis Quijada. Scale bars: A1–6 = 5 mm; F1–6, K = 0.5 mm; B1, N1 = 100 µm; B3, C1–2, D1, E2, J1 = 50 µm; J2 = 20 µm; B2, C3–6, D2–3, E1, G, H, J3, L1–2, M1–4, N2, O1–3 = 10 µm; I = 5 µm.