Notes on Urnula Campylospora (Pezizales)

Notes on Urnula campylospora (Pezizales)

Matteo CARBONE Summary: The authors present a detailed morphological study of Urnula campylospora (Berk.) Cooke based Carlo AGNELLO on a Tasmanian collection and on four New Zealand specimens housed in the New Zealand Collection of Fungi and Plant Pathogens (PDD). Nomenclatural and taxonomical notes are provided along with photo- graphs and drawings of microscopic and macroscopic features. Keywords: Ascomycota, Sarcosomataceae, Plectania sect. Curvatisporae, Plectania chilensis, Urnula mexicana, Ascomycete.org, 5 (1) : 25-32. Rhizina reticulata, Peziza cinereonigra, Bulgaria sarasinii, Gloeocalyx bakeri, Discina pallidorosea, taxonomy. Janvier 2013 Mise en ligne le 03/01/2013 Riassunto: Gli autori presentano un dettagliato studio morfologico di Urnula campylospora (Berk.) Cooke ba- sato su una raccolta effettuata in Tasmania (Australia) e quattro raccolte conservate presso la New Zealand Collection of Fungi and Plant Pathogens (PDD). Lo studio è corredato da note nomenclaturali e tassono- miche, fotocolor di materiale fresco, di microscopia e disegno al tratto. Parole chiave: Ascomycota, Sarcosomataceae, Plectania sect. Curvatisporae, Plectania chilensis, Urnula mexicana, Rhizina reticulata, Peziza cinereonigra, Bulgaria sarasinii, Gloeocalyx bakeri, Discina pallidorosea, tassonomia.

Résumé : les auteurs présentent une étude morphologique détaillée de Urnula campylospora (Berk.) Cooke basée sur une récolte de Tasmanie (Australie) et sur quatre collections de Nouvelle-Zélande conservées dans la New Zealand Collection of Fungi and Plant Pathogens (PDD). Des notes nomenclaturales et taxinomiques sont fournies avec des photographies, ainsi que des dessins des caractères microscopiques et macrosco- piques. Mots-clés : Ascomycota, Sarcosomataceae, Plectania sect. Curvatisporae, Plectania chilensis, Urnula mexicana, Rhizina reticulata, Peziza cinereonigra, Bulgaria sarasinii, Gloeocalyx bakeri, Discina pallidorosea, taxinomie.

Introduction campylospora (Berk.) Rick, Ann. Mycol., 4 (4): 310 (1906); Plectania campylospora (Berk.) Nannf., in Korf, Mycologia, 49 (1): 110 (1957). = Rhizina reticulata Berk. & Broome, J. Linn. Soc., Bot., 14: 102 Originally described from New Zealand by BERKELEY (in HOOKER, (1875), fide ANNFELDTN (unpub.) and RIFAI (1968); Peziza reticulata 1855) as Peziza campylospora, this species had been transferred to (Berk. & Broome) Petch, Ann. Royal Bot. Gard. Paradeniya, 4: 422 four different genera:Macropodia Fuckel (SACCARDO, 1889); Urnula Fr. (1910), illeg., not Peziza reticulata Grev., Scott. Cryptog. Fl., 3: t. 156 (COOKE, 1892); Discina Fr. (RICK, 1906a); Sarcosoma Casp. (RICK, 1906b), (1825). before being selected by KORF (1957) as the type species of Plectania = Peziza cinereonigra Berk. & Broome, Trans. Linn. Soc. London, II sect. Curvatisporae Korf. Phylogenetic studies conducted by CARBONE ser. Bot., 1 (6): 404 (1879), fide ANNFELDTN (unpub.) and RIFAI (1968); et al. (2013a) on part of the present material placed it in the Urnula Geopyxis cinereonigra (Berk. & Broome) Sacc., Syll. Fung., 8: 66 (1889). clade. The species belonging to this clade show a wide range of mor- = Bulgaria sarasinii Henn. (ut “sarasini”), Monsunia, 1: 38 (1900), phological features so we decided to carry out an in-depth study. fide LE GAL (1953); Sarcosoma sarasinii (Henn.) Boedijn, Bull. Jardin Bot. Buitenzorg, sér. III, 12: 275 (1932). Materials and methods = Gloeocalyx bakeri Massee, Bull. Misc. Inform. Royal bot. Gard. Kew, 175-177: 155 (1901), fide CKBLADE (1968) and RIFAI (1968). Microscopic characters are based on dry specimens rehydrated in water. Two optical microscopes were used: Olympus CX41 trino- Misapplied name: Sarcosoma sarasinii (Henn.) Boedijn sensu BOE- cular and Optika B353 trinocular with plan-achromatic objectives DIJN (1932) = Plectania rhytidia (Berk.) Nannf. & Korf, Mycologia, 49 4×, 10×, 40×, 100× in oil immersion. Microscopic pictures were (1): 110 (1957). For synonymy of the latter see CARBONE et al. (2010). made using a Nikon Coolpix 4500 camera, and a Nikon D3100 camera. The following main reagents were used: Melzer’s reagent, cot- Typification: Type housed in the Kew Gardens Fungarium under ton blue, Congo red. Water mounts were used for the observation of the accession number K(M) 177159 (Dr. Begoña Aguirre-Hudson, the pigmentation and spore size. At least 30 spores were measured pers. comm.). from each apothecium. Original diagnosis: “Peziza campylospora, Berk.; fuliginea, cupula breviter stipitata obliqua extus rugosa subtiliter fibrillosa, carne albida, Taxonomy sporidiis majoribus oblongis curvatis. Hab. On decayed wood, Colenso. Urnula campylospora (Berk.) Cooke, Handbook of Australian Cup above an inch across, oblique, lobed, wrinkled externally, and Fungi: 268 (1892). clothed with inconspicuous myceloid flocci. Stem short, wrinkled. Hymenium fuliginous, like the cup tinged with vinous-red; subs- Basionym: Peziza campylospora Berk., in Hooker, Flora Novae-Ze- tance nearly white, composed of intricate threads. Asci linear. Spo- landiae: 200 (1855). ridia oblong, strongly curved, 1/750 of an inch long; paraphyses Synonyms: slender, very slightly incrassated above. — Allied to the last1, but ≡ Macropodia campylospora (Berk.) Sacc., Syll. Fung., 8: 159 (1889); distinguished at once by the larger curved sporidia, its different ha- Discina campylospora (Berk.) Rick, Broteria, 5: 31 (1906); Sarcosoma bitat, etc.”

1 Peziza rhytidia. 25 Description Macroscopic characters Apothecia up to 5 cm in diameter, funnel-shaped cup, subsessile to definitely stipitate.Hymenium dark brown to blackish with age. External surface velvety to definitely warty, dark brown to brownish black, horizontally corrugated especially near the margin. Stem up to 2 cm high, usually vertically corrugated.

Microscopic characters Asci 410–490 × 14–18 μm, cylindrical, operculate, inamyloid, eight-spored, with very thick walls (up to 2–2.2 μm), and a tapered, flexuous base which sometimes shows a generative hypha connected more or less distantly from the basal septum. Paraphyses not, or only slightly, exceeding the length of the asci, 2–3 μm wide, cylindrical, closely septate, sometimes anastomosing; tips straight to flexuous and sometimes lobed/diverticulated; an extracellular, brownish, amorphous pigment is present in the upper part (in dried specimens observed in water mounts), ag- glutinating them to bundles. Hymenial hairs cylindrical, as long as the paraphyses, 3.5–4 (–5) μm wide, non-septate but with a single septum at the very base; tips simple to slightly subcapitate, curved to al- most hooked; pale brownish, especially in the upper part due to the same kind of pigment as in the paraphyses. Spores smooth, curved with rounded poles being allantoid to bean-shaped, 26–32 (–35) × 9.5– 11.5 (–13) μm, av. 29.5 × 10.5 μm, Q = (2.5–) 2.7–3.1 (– 3.6), av. Q = 2.9, subhyaline, with 0 to 4 guttules, walls up to 1 (–1.5) μm thick; very young spores are glo- bose, smooth and thick walled. Subhymenium of a dense textura intricata of cylindrical, closely septate hyphae, up to 4 μm in diameter with thickened, more or less dark brown walls; at low magnification, it appears uniformly brownish to brown. Medullary excipulum of loose textura intricata immersed in a Urnula campylospora. Picture: G. Gates gelatinous matrix, with cylindrical, hyaline, septate hyphae, 2.5–3.5 (–6) μm in diameter, with walls very slightly thickened up to 0.5 μm. In some observations of dried spe- Discussion cimens few encrusted hyphae have been observed. Ectal excipulum of textura angularis made up of elements up to 27 μm wide and/or high, very dark brown due to the colored walls and the pre- The examined material agrees perfectly both with the original sence of an abundant encrusting, amorphous to crystalline brown description (BERKELEY, in HOOKER, 1855) and Rifai’s type revision (RIFAI, pigment. External hairs mainly of two types, although interme- 1968). Urnula campylospora is easy to recognize morphologically, diates can be found: 1) short, hyphoid, cylindrical, av. 5–6 μm wide especially because of the allantoid spores. As reported by CARBONE et (in some mounts up to 8 μm), septate, subhyaline to very pale yel- al. (2013b), Urnula mexicana (Ellis & Holw.) M. Carbone, Agnello & lowish-brown colored, thin to medium-walled, heavily encrusted by A.D. Parker surely belongs to this species complex but differs in pos- an amber brown to dark brown pigment; 2) long, true hairs, 5–6 μm sessing only hyphoid external hairs, straight hymenial hair tips and wide, thick walled up to 1 μm, brown to dark brown, paler in the in having spores with a lower Q ratio. These morphological diffe- upper part, mainly smooth but also slightly encrusted by the same rences, together with the disjuncted geographical distribution, led pigment of the previous type, especially in the basal part. Subicu- CARBONE et al. (2013b) to consider it as a distinct species, although lum made up of cylindrical, 5–7 μm wide septate hyphae, brown this has yet to be confirmed genetically. due to an epimembranaceous pigmentation, with smooth walls thickened up to 0.8–1 μm. Based on the description of GAMUNDÍ (1971), Plectania chilensis (Mont.) Gamundí, originally described from Chile (MONTAGNE, 1850), Studied collections may be another member of this group. According to SPEGAZZINI AUSTRALIA. Tasmania, Hobart, Myrtle Gully, Mount Wellington, on (1918), LE GAL (1953) and GAMUNDÍ (1971), and also to macro-micro- dead wood of Eucalyptus obliqua, 15.IV.2012, leg. et det. G. Gates (HO morphological pictures of a collection made by Prof. Goetz Palfner 567025). NEW ZEALAND. Bay of Plenty, Te Urewera, Mangapae, UA1 (University of Concepción, Chile), this species, despite having a si- W2, on decaying wood, 24.V.2005, leg. B. Paulus & M. Fletcher, det. milar coloring as U. campylospora, has a more regularly funnel-sha- B. Paulus (PDD-83522 and PDD-83528, as Plectania campylospora). ped cup and narrower spores which are much less curved and Bay of Plenty, Te Urewera, Te Waiiti, on fallen log, 17.05.2006, leg. mostly asymmetrical. It is so far known from wood of Nothofagus P.R. Johnston & B. Paulus, det. B. Paulus (PDD-88805, as P. campylos- and Aextoxicum only (GAMUNDÍ, 1971). At present we retain it in the pora). Waikato, Rangitoto Station, on fallen wood, 16.V.2006, leg. et det. P.R. Johnston (PDD-89182, as P. campylospora). genus Plectania until molecular studies will establish its correct position. 26 Plate 1 – Historical plates. A: Peziza (Geopyxis) cinereo-nigra in BERKELEY & BROOME (1879); B: Peziza rhytidia in MASSEE (1896); C: Urnula campylospora in COOKE (1892); D: Bulgaria sarasinii in HENNINGS (1900). With permission by “Biblioteca dell’Orto Botanico dell’Università degli studi di Padova” 27 Fig. 1 – Urnula campylospora. Microscopic characters of the collection PDD 88805. A: Medullary excipulum in Congo red; B: Ectal excipulum in Congo red; C-D: Hyaline external hairs in water; E-F: True external hairs in water. Scale bars = 10 μm. Pictures: C. Agnello & M. Carbone. 28 Fig. 2 – Urnula campylospora. Microscopic characters of the collection PDD 88805. A-B: Hymenium with asci, paraphyses and hymenial hair in Congo red; C: Asci apex in Congo red; D: Ascus and spores in Congo red; E: Spores in water mount. Scale bars = 10 μm. Pictures: C. Agnello.

Comparing the inequilateral spores of Plectania yunnanensis W.Y. from their more regular spores shape. For a quick overview on these Zhuang with those of P. campylospora and P. chilensis, it is pretty taxa see CARBONE & AGNELLO (2012, 2013) and CARBONE et al. (2009). clear that the spores of the first are definitely not curved (ZHUANG & WANG, 1998) and that again more studies are required to figure out Notes on synonyms and misapplications the real position of this species. Rhizina reticulata Berk. & Broome was described from Sri Lanka With regards to other Urnula species such as Urnula hiemalis with the following diagnosis (BERKELEY & BROOME, 1875): “E cupuliformi Nannf. and Urnula mediterranea (M. Carbone, Agnello & Baglivo) expansa, substipitata, subtus velutina, reticulata, brunnea; hymenio vi- M. Carbone, Agnello & P. Alvarado, they are easily distinct mainly noso; sporidiis oblongis curvulis obtusissimis (no. 321). On wood. Hab- 29 Fig. 3 – Urnula campylospora. Microscopic characters of the collection PDD 88805. A: Section; B: Spores; C: Ascus; D: Paraphyses; E: Hymenial hairs; F: Hairs of the ectal excipulum. Drawing: C. Agnello. 30 galla, Dec. 1867, 1868. Neilgherries, E.S. Berkeley. At first obconic, inaequilateralibus, 2-guttulatis, hyalinis 19-21 × 10-12 μ”. We believe then cup-shaped, at lenght expanded, 1-2 inches across; asci linear; that there are no sufficient data to support this synonymy. As the paraphyses slightly clavate, sporidia .001 long, .0005 wide”. original diagnosis appears to describe a different species and mo- This species was illegitimately transferred to Peziza Fr. by PETCH reover because of the geographic difference, we doubt the syno- (1910), who was not aware of the earlier homonym Peziza reticulata nymy of these two taxa. According to HEIN (1988) the type of this Grev. (GREVILLE, 1825). Nannfeldt, according to his annotations on the species is also lost. herbarium envelope of the type specimen, and then RIFAI (1968) placed R. reticulata in synonymy with P. campylospora. It is noteworthy Geographical distribution that MASSEE (1896) had earlier used this synonymy, but in this publi- According to PADEN (1983), U. campylospora occurs in “Australasia, cation he also regarded Plectania rhytidia as another synonym. Brazil, Buthan, Ceylon and Jamaica”, and further reports come from Similarly, Peziza cinereonigra Berk. & Broome, a species described Mexico (POMPA-GONZÁLES & CIFUENTES, 1991), India (PANT & PRASAD, from Australia (BERKELEY & BROOME, 1879), was regarded as a synonym 2008) and China (TENG, 1963; ZHUANG & WANG, 1998). For the reasons of U. campylospora by NANNFELDT (unpubl.) and RIFAI (1968). The ori- mentioned above, we are not currently able to determine its exact ginal diagnosis of P. cinereonigra seems to support this point of view: range of distribution because further combined morphological- “Cupula infundibuliformis, (sicca) cinereo-nigra, extus laevis, profunde phylogenetic studies are required in order to figure out if similar but et irregulariter rugosa, margine incurva; stipes laevis, concolor, sursum distinct species could exist. For the time being, we therefore prefer dilatatus; asci lineares, 8 sporidia curvata continentes; paraphyses li- to keep a restricted area to Australasia for U. campylospora. neares, apice furcati. A fine species, allied to P. corium, from which it differs in its more tender substance, and consequently deeply wrin- Acknowledgements kled cups and curved, somewhat sausage-shaped, sporidia, which measure 0.001 inch in lenght by about half that in width. It grows on wood, while P. corium occurs on the ground; but its affinities are with We are grateful to Peter Johnston (New Zealand Landcare Re- the section Geopyxis. The cups are about 1 inch 2 lines across, and search Herbarium, PDD) for the critical and linguistic review of the with the stem are about 1 ¼ inch high”. manuscript and for his improving suggestions; Wanda Daley (New Bulgaria sarasinii Henn. was described from Indonesia, Sulawesi Zealand Landcare Research Herbarium, PDD) for arranging the loan of the New Zealand collections; Genevieve Gates (Hobart Univer- Island, with the following diagnosis (HENNINGS, 1900): “Ascomatibus gelatinosis, substipitatis, cupulatis, atris, 2 cm diametro, extus rugosis, sity, Tasmania) for sending us her collection and picture; Goetz Palf- pruinosis, intus laevibus; ascis cylindraceis, obtusis 8-sporis ca. 250- ner (University of Concepción, Chile) for sharing his photograph of 300 μ longis p. sporif 150-180 × 15-17 μ; paraphysibus filiformibus, Plectania chilensis; Dr. Begoña Aguirre-Hudson (Kew Gardens Her- multiguttulatis 3-3½ μ crassis, fuscidulis; sporis monostichis, ellipsoi- barium, London) for the information about the type specimen of deis interdum curvatis, utrinque obtusis, primo multiguttulatis intus P. campylospora. fuscidulis, dein 1-2 guttulatis 20-24 × 14-16 μ”. Although the type material has been lost (HEIN, 1988), we agree with LE GAL (1953) that References HENNINGS’ (1900) description and plate show features perfectly coin- cident with P. campylospora. BOEDIJN (1932) validly transferred this BERKELEY M.J. 1855. — Nat. Ord. CII. Fungi. In: HOOKER J.D. The botany species in the genus Sarcosoma, but it must be noted that he was of the Antarctic Voyage of H.M. discovery ships Erebus and Terror in using this name in the sense of our concept of P. rhytidia. the years 1839-1843 under the command of captain Sir James Clark Gloeocalyx bakeri Massee was described from Australia with the Ross. II. Flora Novæ-Zelandiæ. Part II. Flowerless plants: 172-210. following diagnosis (MASSEE, 1901): “Ascomata sessilia, cupulata, sub- London, L. Reeve. gelatinosa, glabra, sicco cornea, corrugata, extus reticulato-venosa, BERKELEY M.J. & BROOME C.E. 1875. — Enumeration of the Fungi of Cey- ubique atra vel disco pallida, 1-2 cm. lata. 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Rivista di Micologia, 53 For the time being, we have no reason to doubt in all these syno- (2): 119-135. nymies. Conversely, we disagree with the synonymy of Discina pal- CARBONE M., AGNELLO C. & ALVARADO P. 2013a. — Phylogenetic studies lidorosea Henn. proposed by RICK (1906b). D. pallidorosea was in Sarcosomatacae (Ascomycota, Pezizales). Ascomycete.org, 5 (1): described from Blumenau in the south of Brazil (HENNINGS, 1902, as 1-12. D. pallide-rosea) with the following diagnosis: “Ascomatibus aquoso- CARBONE M., AGNELLO C. & PARKER A.D. 2013b. — Urnula padeniana (Pe- carnosis vel subgelatinosis (Möller teste), subcupulato-explanatis, dein zizales), and the type study of Bulgaria mexicana. Ascomycete.org, discoideis, brevissime stipitatis, extus pallidis rugoso-plicatis venosis (in 5 (1): 13-24. alcohole), margine recurvatis, disco concavo vel plano, levi, glabro pal- COOKE M.C. 1892. — Handbook of Australian Fungi. 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Matteo Carbone Via Don Luigi Sturzo 173 16148 Genova Italy [email protected] Carlo Agnello Via Antonio Gramsci 11 72023 Mesagne Italy [email protected]