Johnston, P.R. 1986: Rhytismataceae in New Zealand. 1. Some foliicolous species of Coccomyces de Notaris and Propolis (Fries) Corda. New Zealand Journal of Botany 24(1): 89-124.
Details
Associations
Descriptions
Ascocarps intraepidermal. In vertical section upper stromatal layer 15-30 µm wide, comprising brown to dark brown, angular cells, 5-8 .µm diam. A few cylindric, hyaline, periphysis-like cells, 10-15 x 2-3 µm, near opening. Lower stromatal layer 20-30 µm wide, of 3-5 layers of dark brown, thick walled cells 5-8 µm diam. A 15-20 µm wide layer of gelatinised tissue present between the lower stromatal wall and the sub-hymenium, extending a short distance up the sides of the hymenium, giving rise to few, branching, excipular-like elements.
Paraphyses 1.5-2 µm diam., swollen to 4-6 µm at the clavate apices, extending 10- 15 µm beyond asci. Asci 101-118 x 7.5-9 µm, subclavate, tapering in to a broadly truncate apex, wall uniform in thickness, non-amyloid, 8-spored. Ascospores 35-43.5 x 2-3 µm, clavate, tapering to the base, 0(-1) septate, hyaline, with a gelatinous cap.
ETYMOLOGY: clavatus = clavate; refers to ascospore shape.
C. clavatus is similar to C. phyllocladi, also found on Phyllocladus. The two species are indistinguishable macroscopically, and are similar in appearance in cross-section, both having a characteristic, gelatinous layer between the lower stromatal wall and the sub-hymenium. They differ in the size and shape of the asci and ascospores.
No rhytismataceous species have previously been reported on Phyllocladaceae. Two Hypoderma-like species have also been found on this host in New Zealand.
South America: COLOMBIA: Dpto. Antioquia, Medellin Pto.-Valdivia Rd., on unidentified fern, K. P. Dumont, S. E. Carpenter, M. A. Sherwood, 12.VIII.1976 (holotype - NY).
Ascocarps subepidermal. In vertical section upper stromatal layer poorly developed, 20-25 µm wide, not becoming wider around opening, composed of brown to very pale brown, thick walled, angular to globose cells, 3-6 µm diam. Inside of stromatal layer lined with periphyses when immature. Excipulum becoming well-developed as ascocarps mature, arising from inner layers of upper stroma, comprising hyaline, closely septate hyphae, which become swollen, and embedded in dark brown material at apices. Lower stromatal layer 5-15 µm wide, the outer 1-2 layers of cells dark brown and thick walled, with hyaline, globose cells to the inside.
Paraphyses 1-1.5 µm diam., branching several times near apices not or only slightly swollen. intermixed with crystals, extending 15-20 µm beyond asci. Asci cylindric to subclavate, (132-)150-195(-210) x 7.5-11 µm, tapering to an acute apex, wall not thickened at apex, non-amyloid, 8-spored. Ascospores filiform, (95-)110-125 x 1.8-2.5 µm, tapering slightly to both ends, 0-1 septate, often coiling when released, surrounded by poorly developed gelatinous sheath.
Pycnidia subepidermal. In vertical section lenticular in shape, wall 5-10 µm wide, of brown, angular cells, lined with hyaline, thin walled cells on which the conidiogenous layer is held. The lower wall is wider and darker than the upper. Conidiogenous cells lining both upper and lower walls, comprising flask-shaped, sympodial, solitary conidiogenous cells, often with two conidia held at the ends. Conidia short-cylindric, 3.5-5.5 x 1 µm, hyaline, non-septate.
The ascocarps of the type specimen of C. crystalligerus are larger, have a rounder, more regular-shaped opening, and better developed layer of crystals on the hymenium, compared with New Zealand collections. Pycnidia are not present on the type specimen, but were found on some, but not all, of the New Zealand collections. Otherwise the New Zealand collections of this fungus fit the description of the type specimen well, especially with regard to the host, the appearance and size of the hymenial elements, and the structure of the ascocarp margin.
The only other rhytismataceous taxon so far consistently found on ferns in New Zealand is a Hypoderma-like species on Gleichenia cunninghamii Heward ex Hook. C. radiatus was found once on Dicksonia squarrosa
Ascocarps intraepidermal. In vertical section upper stromatal layer 15-30 µm wide, comprising pseudoparenchymatous, thin walled cells, 5-8 µm diam., cells dark brown to carbonaceous around opening of ascocarp, paler brown near outer edge. An extra layer of hyaline, gelatinous cells is present between the upper stromatal layer and the covering host tissue in immature ascocarps. This layer is often not visible in mature ascocarps. Lower stromatal layer separate from upper, 8-15 µm wide, of 2-4 layers of dark brown, thick walled, globose cells, 4-10 µm diam. Subhymenium 10-15 µm wide, of hyaline, thin walled cells. Layer of gelatinised tissue up to 40 µm wide developing between subhymenium and lower stromatal layer, and sometimes also present on the inside of the upper stromatal layer. Excipulum-like structure becoming well developed, arising from the layer of gelatinised tissue, up to 40-80 µm wide, elements 1.5-2.5 µm diam., closely septate and often capped by a dark brown substance.
Paraphyses 1.5-2 µm diam., gradually becoming wider toward apex, up to 2.5-4.5 µm diam., not branching, extending 15-25 µm beyond asci. Asci 131-163 x 7- 10 µm, cylindric, basal stalk short and broad, tapering to a rounded to slightly truncate apex. Ascus wall slightly thickened at apex, 1.5-2 µm thick, with a small apical pore, non-amyloid, 8-spored. Ascospores 90-119 x 1.5-2.2 µm, tapering to basal end, 0-1 septate, straight when released, well-developed gelatinous sheath.
Pycnidia intraepidermal, flat, lenticular in shape, lower wall of dark brown, angular cells, upper wall absent or comprising a few, scattered, pale brown cells. Conidiogenous layer lining the lower wall, comprising a palisade of solitary, cylindric to flask-shaped, hyaline, sympodial, conidiogenous cells, 12-18 x 2-3 µm. Sterile elements, filiform and slightly swollen at the apices, connected by hyphal bridges near bases, 45-80 x 2-2.5 µm, intermixed with the conidiogenous cells. Conidia short-cylindric, rounded ends, hyaline, 0-septate, 4.5-6.5 x 1-1.2 µm.
ETYMOLOGY: cupressinum; refers to host plant.
The ascocarps of C. cupressinum look superficially similar to those of C. araucariae Butin & Speer and C. radiatus Sherwood. C. araucariae (Brasil, Vossoroca, Porana, on Araucaria angustifolia, H. Butin, E. Speer, 11.IX. 1976 - Holotype, ZT) differs in ascus and ascospore size, and in excipular structure. C. radiatus lacks the layer of gelatinised tissue found between the stroma and hymenium in C. cupressinum, has a different excipular structure, and narrower asci.
Two rhytismataceous species have been described from Podocarpaceae - Hypoderma podocarpi Butin from Podocarpus species in Chile (Butin 1970), and Lophodermellina dacrydii Sydow from Dacrydium sp. in Borneo (Sydow 1921) but this is the first record of Coccomyces on Podocarpaceae.
Also found on Dacrydium in New Zealand is a Hypoderma-like species. which is very similar to Hypoderma podocarpi in ascocarp shape and appearance, with both having more or less round ascocarps and white lip cells along the opening slits. The New Zealand species differs from H. podocarpi in having larger asci and ascospores, and 4-spored asci.
Ascocarps intraepidermal. In vertical section upper stromatal layer 30-50 µm wide, comprising pale brown to brown, thin to thick walled, more or less globose cells, 4-8 µm diam. Cells becoming paler and thinner walled near outer edge of ascocarp. Periphyses absent. Lower stromatal layer joining the upper, 15-25 µm wide, of 4-5 layers of dark brown, thick walled, more or less globose cells, 5-9 µm diam., the outer layer of cells with darker and thicker walls. Subhymenium 10-20 µm wide, of hyaline, thin walled, rounded cells, 3-8 µm diam. Layer of disorganised hyphal tissue, 20-50 µm wide, between subhymenium and lower wall. Excipulum poorly developed, comprising a few marginal paraphyses becoming closely septate and sometimes capped by dark brown material.
Paraphyses 2-2.2 µm diam., swollen to 4-7 µm diam. at the clavate or fusoid apices, extending 20-30 µm beyond asci. Asci 104-140 x (5-)5.5-7(7.5) µm, cylindric to subclavate, tapering slightly to rounded apex, non-amyloid, 8-spored. Ascus apical thickening and broad pore sometimes seen when immature. Ascospores filiform, 55-78 x 1.2-1.8 µm tapering slightly to basal end, more or less straight when released, 0-1 septate, gelatinous sheath present (see notes below).
Pycnidia depressed-globose, intraepidermal to subepidermal, ostiole not well-defined, often opening through a stomatal cell. Pycnidial wall continuous, not divided into upper and lower parts, 5-12 µm wide, of 1-3 layers of brown, somewhat flattened, pseudoparenchymatous cells, 2-3.5 µm diam. Conidiogenous layer lining whole pycnidium, comprising hyaline, flask-shaped, sympodial conidiogenous cells, 9-18 x 1.5-3.8 µm, with a long, flexuous neck, often with two conidia held at the ends. Conidiogenous cells solitary or borne on poorly developed, irregularly branching conidiophores, intermixed with filiform, sterile elements, especially in area beneath ostiole. The sterile elements may form a column of tissue, almost reaching to the top of the pycnidium. Conidia short-cylindric, rounded ends, 0-septate, hyaline, (3.8-)4.5-6 x 0.8-1(-1.2) µm.
CHARACTERISTICS IN CULTURE: Ascospores of PDD 43034 and PDD 43964 germinated within 5-6 days. Cultures on oatmeal agar 3.5-4 cm diam. after 4 weeks; aerial mycelium sparse, felted, white; agar surface grey toward centre of colony, yellowish toward edge; reverse, yellow. Pycnidia forming near centre of colony, globose, black walled; conidial ooze yellow. Conidiogenous cells and conidia of same appearance and size as those found in nature.
ETYMOLOGY: globosus = globose, spherical; refers to shape of anamorph pycnidium.
NOTES: Some of the collections on Weinmannia racemosa (PDD 44649, 44650, 44653, 44928, 44962) can be separated from typical C. globosus in having asci 7-9 µm wide, and broadly truncate at the apices, and ascospores (71-)80-98 µm long. In all other respects, including the anamorph, they are identical to C. globosus. They have been included here in C. globosus.
The anamorph pycnidium of C. globosus is unusual when compared to those of most other Rhytismataceae, in not being flattened, and in having the conidiogenous layer lining the whole pycnidial cavity. It is similar to that described for Lophodermium camelliicola by Minter & Sharma (1982).
C. globosus is similar in hymenial dimensions to C. phyllocladi. C. phyllocladi differs in that it lacks pycnidia, and has an excipulum which forms from a layer of gelatinised tissue between the hymenium and the outer stromatal layers. Cultural characteristics also differ.
C globosus is also similar to C. antillarum Sherwood and C. spegazzinii Saccardo. From Sherwood's 1980 descriptions, C. antillarum and C. spegazzinii are more or less the same in ascus and ascospore size and shape, and ascocarp margin structure. They appear to differ only in C antillarum lacking zone lines around the lesions, having an excipulum which remains hyaline, and having periphyses lining the upper stromatal wall. The specimens of C antillarum studied (R. Thaxter, 1912-13, on unidentified leaves, Grand Etang, Grenada - holotype FH; Pfister 1006, on unidentified leaves, Grand Etang, Pfister, Carpenter, Sherwood 6.1.1974 - FH) agreed well with the description of Sherwood (1980), but the only specimens of C. spegazzinii examined (Brasil, Apiahy, Puiggari, 1889 - LPS 34990; Venezuela, Edo. Sucre, Dumont et al., 15.VII.1972, NY-VE5083) were small and in poor condition, and did not match the description of the hymenium of C. spegazzinii given by Sherwood. Compared with C. globosus these species have the same macroscopic appearance and hymenial element dimensions, and the excipulum and the way it develops are also similar. C globosus can be separated from C. antillarum in having an anamorph, and in lacking periphyses. C spegazzinii is apparently associated with zone lines, and lacks an anamorph. However, considering the small number of collections of C. antillarum and C. spegazzinii in herbaria and the relatively minor differences between all three taxa, they may in fact be found to be conspecific when more South American collections are available for comparison.
Ascocarps intraepidermal. In vertical section upper stromatal layer 20-80 µm wide, becoming wider around opening. Wall composed of dense, black tissue around opening ascocarp, otherwise of brown to pale brown, pseudoparenchymatous cells, 4-7 µm diam. Lip cells cylindric, hyaline, becoming dark brown, embedded in gel. Short, hyaline periphyses often present, lining the upper wall. Lip cells and periphyses often elongating and becoming septate, forming an extensive excipulum-like layer in older ascocarps. Lower stromatal layer joining the upper, 8-15 µm wide, of 2-3 layers of thick walled, dark brown cells, 2.5-4 µm diam. Subhymenium 19-30 µm wide, of hyaline, thin walled cells, 3-6 µm diam.
Paraphyses 2-2.2 µm diam., becoming slightly swollen at apices, 3-4.5 µm diam., extending 10-20 µm beyond asci. Asci subclavate, gradually tapering toward the base from near the apex, tapering at top to a more or less acute apex, uniformly thin walled, 104-142 x 7.5-11 µm, non-amyloid, 8-spored. Ascospores acicular, 41-60 x 2.2-3 µm, tapering slightly to the basal end, 0-septate, poorly developed gelatinous sheath.
ETYMOLOGY: lauraceus; refers to host plant family.
NOTES: C. lauraceus is more or less intermediate between C delta (Kunze) Saccardo, and C. dentatus (Kunze & Schmidt:Fries) Saccardo, two species discussed by Sherwood (1980). All three species have subclavate asci and paraphyses with slightly swollen apices.
C lauraceus agrees well with C. delta (Azores, Sao Miguel, Hickling Park, on Lauraceae, R. P. & N. Korf, L. M. Kohn, A. Y. Rossman, 5.IV. 1978 CUP-MM1845; Canary Is., La Palma, Los Tilos, on Ocotea sp., R. P. Korf, W. C. Dennison, L. M. Kohn, M. A. Sherwood, 14.1.1976 - CUP-MM654; ibid., on Laurus sp. - CUP-MM647; Madiera, Chao dos Louros, on Lauraceae, R. P. & N. Korf, L. M. Kohn, A. Y. Rossman, 26.IV.1978 - CUPMM2455) in possessing lip cells, in lacking pycnidia, in ascus size, and in being found exclusively on Lauraceae. It differs from C. delta in having periphyses which develop into an excipulum-like layer, and in having significantly shorter and broader ascospores (41-60 x 2.2-3 µm versus 80-100 x 2 µm).
The ascospore length of C. lauraceus matches that of C. dentatus (Sydow, Mycotheca Germanica 1249- PDD; USA, Oregon, on Castanopsis sp., Sherwood, 8.Vlll. 1978 - FH; Ellis and Everhart, North American Fungi 3336, on Berberis sp. - PDD). However, C. dentatus lacks lip cells, is associated with pycnidia, has shorter asci (80-105 µm), and, according to Sherwood (1980) is found on a wide range of host plants.
This is the only Coccomyces species found on Lauraceae in New Zealand.
Ascocarp intraepidermal. In vertical section upper stromatal layer 15-20 µm wide, comprising rounded, pale brown, thin walled cells. An extra laver of hyaline, gelatinous cells developing between the upper stromatal layer and the covering host tissue. Lower stromatal layer 15 µm wide, comprising 3-4 layers of dark brown, thick walled cells. Excipulum absent.
Paraphyses 2 µm diam., becoming irregularly tangled and swollen up to 4.5µm diam. at apices, extending 15-20 µm beyond asci. Asci cylindric, 195-235 x 9-10.5 µm, tapering slightly to a broad, truncate apex, uniformly thin walled, non-amyloid, 8-spored. Ascospores filiform, 142-206 x 1.8-2 µm, not tapering, 0-3 septate, surrounded by a poorly developed gelatinous sheath.
Pycnidia intraepidermal, flat and lenticular in shape, upper wall absent. Lower wall 7-12 µm wide, of 1-3 layers of dark brown, thick walled cells, 3-5 µm diam., lined with 1-2 layers of hyaline cells, on which the conidiogenous layer forms. Conidiogenous cells 8-12 x 1.5-2 µm, solitary, cylindric to flask-shaped, sympodial. Conidia cylindric with rounded ends, 4-6 x 1 µm, hyaline, non-septate.
CHARACTERISTICS IN CULTURE: Ascospores germinating on agar plates after 48 h. Cultures on oatmeal agar 3.5-4 cm diam. after 6 weeks; aerial mycelium tufted, pale grey, in two concentric bands; agar surface undulate, dark brown in colour, fruiting bodies appear to be forming near edges of colonies, but remaining sterile.
ETYMOLOGY: libocedri; refers to host plant.
NOTES: The ascocarp is similar in external appearance to C. cupressinum, but the two species are easily distinguished microscopically. Lophodermium juniperi (Greville) Darker is recorded from Libocedrus decurrens Torr. (Hunt & Ziller 1978).
This species has lip cells lining the single opening slip, asci 110-130 x 15-17 µm, and ascospores 70-90 x 2-3 µm. Two species, described as Hypoderma by Butin (1970) from Pilgerodendron species, have saccate asci and ascospores less than 50 µm long.
New Zealand: NORTHLAND: Puketi S.F., Picnic Area, Bush Walk, on Knightia excelsa, P. R. Johnston, G. J. Samuels, 16.IV. 1982 (PDD 44664); Mangamuka Summit, on undetermined leaf, P. R. Johnston, G. J. Samuels, 15.IV.1982 (PDD 44665). AUCKLAND: Waitakere Ra., Centennial Track, on Astelia solandri, P. R. Johnston, 12.X. 1982 (PDD 43317); Waitakere Ra., Cowan Track, on Knightia excelsa, P. R. Johnston, G. J. Samuels, R. H. Petersen, 27.IV.1983 (PDD 44670); Clevedon, Morehau Reserve, on Knightia excelsa, P. R. Johnston, G. J. Samuels, R. H. Petersen, 5.V.1983 (PDD 44672); ibid., on Metrosideros fulgens, P. R. Johnston, E. M. Gibellini, 22.V. 1984 (PDD 45196); Waitakere Ra., Sharp's Bush, on Knightia excelsa, P. R. Johnston, 28.XI.1983 (PDD 44673). COROMANDEL: Papa Aroha, Rautawhake Stream, on Metrosideros fulgens, P. R. Johnston, E. M. Gibellini, 12.VIII.1984 (PDD 45362). TARANAKI: Mt. Egmont, Puniho Track, on Knightia excelsa, P. R. Johnston, G. J. Samuels, R. H. Petersen, 25.IV.1983 (PDD 44669); Mt. Egmont, Waiweranui Track, on Weinmannia racemosa, P. R. Johnston, G. J. Samuels, R. H. Petersen, 25.IV.1983 (PDD 44671). NELSON: Karamea, Umere Rd., on Pseudopanax sp., P. R. Johnston, G. J. Samuels, R. E. Beever, R. H. Petersen, 14.IV. 1983 (PDD 44668). WESTLAND: Fox Glacier, Lake Gault, on Pseudopanax sp., P. R. Johnston, 10.IV.1983 (PDD 44667); Fox Glacier, Mt. Fox Track, on Pseudopanax colensoi (Hook. f.) Philipson, P. R. Johnston, G. J. Samuels, R. E. Beever, R. H. Petersen, 8.IV.1983 (PDD 44666).
South America: COLOMBIA: Dpto. Valle del Cauca, Buga-Buenaventura Rd, on unidentified leaves, Dumont, Molina, 30.VIII.1976 (NYC07552). VENEZUELA: Edo. Aragua, Parq. Nac. Henry Pettier, Maracay, on unidentified leaves, Dumont, Haines, Samuels, 13.VI.1971 (NY-VE2112). Central America: CUBA: Cuba Fungi 442, C. Wright (Phacidium limitatum - K - holotype).
Ascocarp intraepidermal. In vertical section upper stromatal layer 20-40 µm wide, not extending as far as the lower wall. Small cap of dense, black tissue around opening, otherwise of brown to dark brown, pseudoparenchymatous cells, 4-7 µm diam. Lower stromatal layer 8-15 µm wide, of 1-2 layers of thick walled, brown to dark brown, more or less globose cells, 5-10 µm diam. Subhymenium 10-20 µm wide, of thin walled, hyaline, globose cells, 7- 10 µm diam., a layer, up to 50 µm wide, of loose hyphae and crystals present between subhymenium and lower wall. Excipulum becoming well developed, up to 50 µm wide, arising from inner layers of upper wall, comprising several rows of hyaline, closely septate hyphae, becoming slightly swollen and covered by a dark brown substance at apices. Excipulum formed after the ascocarps have opened, and may not be apparent in young, recently opened ascocarps. In young ascocarps has the appearance of a well developed layer of short, hyaline, cylindric periphyses. These become longer and develop septa to form the excipulum.
Paraphyses 2-3 µm diam., gradually increasing in width to 3.5-4.5 µm diam. at apices, extending 15-20 µm beyond asci. Asci cylindric, tapering slightly to a rounded apex, 95-125 x 4-5(-5.2) µm, either uniformly thin walled, or sometimes with an apical thickening, with a short, broad pore, non-amyloid, 8-spored. Ascospores filiform, 44-75 x 0.8-1.0 µm, straight or irregularly curving when released, not tapering, 0-1 septate, gelatinous sheath poorly developed.
Pycnidia intraepidermal, lenticular. Lower wall 8- 10 µm wide, of 2-3 layers of thick walled, dark brown, flattened cells, 2-4 µm diam. Upper wall almost obsolete, consisting of a few hyaline, thin walled cells beneath the cuticle, and a small group of dark walled cells around the ostiole. Conidiogenous layer not seen (see notes below).
HABITAT: Found on fallen leaves of Knightia excelsa, Pseudopanax species, Astelia solandri A. Cunn., Clematis paniculata Gmel., Elaeocarpus dentatus J. R. et G. Forst.) Vahl., Metrosideros fulgens, and Weinmannia racemosa. Although found on a wide range of host plants, most collections were found on either Knightia excelsa (12) or Pseudopanax species (4).
Has previously been reported on various hosts from Central and South America, and Java (Sherwood 1980).
The pycnidia assumed to be those of the C. limitatus anamorph were found in association with the teleomorph in all collections. The pycnidia are typical in structure of Leptostroma, but all those examined were sterile. Many Rhytismataceous taxa form an anamorph and teleomorph within the same lesion, but at different times. It is not possible to identify C. limitatus unless the teleomorph state is present, as its most common host species also provide a substrate for a number of other Rhytismataceous taxa with similar anamorphs.
C. limitatus is the only species of Coccomyces found in New Zealand with ascospores less than 1.0 µm wide. It is similar to a number of the species included in the "C. leptosporus complex" described by Sherwood (1980). Species in this group which have 3-4 sided, angular ascocarps are C. tesselatus Sherwood (Thaxter 2612, on unidentified leaves, Grand Etang, Grenada, Thaxter 1912-13 - FH; Pfister 844, on unidentified leaves, Guadeloupe, Pfister, Carpenter, Sherwood, 4.1.1974 paratype - FH), C. antillarum (see specimens cited under C globosus), C philippinus (Schmidt & Kunze) Karsten (Baker, Fungi Malayana, on Neolitsea, Mt. Maquiling, Prov. Laguna, Philippines, June 1914 - type - S), and C. limitatus. C. limitatus can be separated from these other species by its very narrow asci (4-5.5 µm wide) and by its paraphyses which are broad along their entire length, rather than being swollen near the apex.
The type specimen of C limitatus has ascocarps lacking an excipulum, and asci slightly wider (5-6 µm) than those found in New Zealand collections. However the type specimen is quite small, and in all other respects, both macroscopic and microscopic, is identical to the New Zealand fungus. Other collections from South America cited by Sherwood as C. limitatus (e.g., NY-VE2112) did have narrower asci, and an excipulum developing in the same way as that described above for the New Zealand collections.
Ascocarps initially subcuticular, with some epidermal cells becoming filled with fungal tissue as the ascocarp ages. In vertical section upper stromatal layer 15-20 µm wide, comprising brown to pale brown, thin walled, rounded cells, 4-6 µm diam. This layer does not extend as far as the outer edge of the ascocarp. The area between the upper and lower stromatal layers is filled with hyaline, gelatinous tissue. Cylindric, vertically orientated, hyaline, gelatinous-walled cells develop between the upper stromatal layer and the covering host tissue. These cells form a well-developed layer in immature ascocarps, but are often difficult to distinguish after the ascocarp has opened. A few periphyses, 12-17 x 2-3 µm, line the inside of the upper wall. The lower stromatal layer is 60-70 µm wide; the outer 30-40 µm is composed of dark brown, thick walled, globose cells, 5-8 µm diam., the inner part composed of hyaline, globose cells, 6-10 µm diam. Subhymenium 10- 15 µm wide, of hyaline, gelatinous-walled cells.
Paraphyses 2-2.5 µm diam., gradually increasing in width to 3-4 µm diam. at the apex, not branching, not gelled together, extending 20-30 µm beyond the asci. Asci 137-155 x 8.5-10 µm, subclavate, tapering in slightly to the broadly truncate to slightly rounded apex, the wall sometimes with a short, broad apical pore, non-amyloid, 8-spored. Ascospores 58-72 x 2.5-3 µm, tapering slightly to basal end with a small swelling at the very base, 0-septate, straight when released, gelatinous cap at both ends.
Pycnidia intraepidermal, flat, lenticular in shape, lower wall of brown, angular cells, upper wall absent. Conidiogenous layer lining the lower wall, comprising a palisade of solitary, cylindric, hyaline, sympodial conidiogenous cells, 8-16 x 2-3 µm. Intermixed amongst the conidiogenous cells, especially near the centre of the pycnidia are filiform, hyaline, sterile elements, 45-60 x 1-2 µm. Conidia, 5-6 x 1-2 µm, short-cylindric, rounded at both ends, hyaline, 0-septate.
ETYMOLOGY: longwoodicus; refers to type locality.
NOTES: The ascocarps of C. longwoodicus are similar in macroscopic appearance to two other species found on southern conifers, C. cupressinum and C. libocedri, with all having a pale, poorly developed upper ascocarp wall. The three species can be distinguished using ascus and ascospore size.
Ascocarps intraepidermal. In vertical section upper stromatal layer up to 18-30 µm wide, cap of dense, black tissue around opening, otherwise comprising hyaline to brown, thin walled cells, 4-9 µm diam., paler near the outer edge of the ascocarp. Lower stromatal layer 20-30 µm wide, of 2-4 layers of dark brown, thick walled cells, 4-8 µm diam. Subhymenium seated on a 45-60 µm wide layer of gelatinised hyphae which also extends up the sides of the hymenium. Excipulum present, arising from the gelatinous layer at the sides of the hymenium, elements 2-2.5 µm diam., irregularly branching, septa about 10- 15 µm apart.
Paraphyses 1.5-2 µm diam., swollen to 4-6.5 µm at the fusoid to clavate apex, extending 15-25 µm beyond asci. Asci 119-137 x 6-7.5 µm, cylindric to subclavate, apex rounded, wall slightly thickened at apex, with small central pore, non-amyloid, 8-spored. Ascospores filiform, 62-77 x 1.2-1.5(-2) µm, tapering to basal end, 0-1 septate, more or less straight when released, surrounded by gelatinous sheath.
CHARACTERISTICS IN CULTURE: Ascospores of PDD 44674 and PDD 44676 germinated on agar plates after 3 days. Growth of colony slow, 0.5-1 cm diam. after 4 weeks, 3 cm diam. after 14 weeks; aerial mycelium cottony, white, sparse; agar brownish in reverse. Remaining sterile.
ETYMOLOGY: phyllocladi; refers to host plant.
NOTES: Many of the infected phylloclades have the appearance of being dissected by numerous zone lines. These zone lines however are not narrow and black as found in other species, but are formed by a darkening of the host tissue around the separate fungal lesions, and are brown, broad, and diffuse at the edges. They appear to be formed by the host as a reaction to infection, rather than by the fungus.
C. phyllocladi is similar to C. globosus in hymenial dimensions. The two species can be distinguished by host, absence of pycnidia in C. phyllocladi, and in C. globosus lacking the gelatinised layer lining the stromatal wall of C. phyllocladi. Cultural characteristics also differ.
See also notes under C. clavatus.
New Zealand: NORTHLAND: Puketi S.F., Picnic Area, Bush Walk, on Knightia excelsa, P. R. Johnston, G. J. Samuels, 16.IV.1982 (PDD) 44688). AUCKLAND: Waitakere Ra., Piha Valley, Cowan Track, on Rubus cissoides A. Cunn., P. R. Johnston, G. J. Samuels, R. H. Petersen, 27.IV.1983 (PDD 44687); Waitakere Ra., Swanson, University Hut, on Knightia excelsa, P. R. Johnston, 31.111.1983 (PDD 44686); Waitakere Ra., Kauri Knoll Track, on Dracophyllum latifolium A. Cunn., P. R. Johnston, E. M. Gibellini, IO.VII.1983 (PDD 44678); Waitakere Ra., Walkers Bush Track, on Cordyline sp., P. R. Johnston, G. J. Samuels, E. H. C McKenzie, 4.VIII.1982 (PDD 44689); Beachhaven, on Dracophyllum sp., P. R. Johnston, 30.X.1983 (PDD 44681); Waitakere Ra., Sharps Bush, on Leptospermum scoparium, P. R. Johnston, 14.XI.1983 (PDD 44680); Waitakere Ra., Kauri Knoll Track, on Dracophyllum latifolium, P. R. Johnston, 28.XI.1983 (PDD 44679). COROMANDEL: Papa Aroha, Rautawhake Stream, on Metrosideros fulgens, P. R. Johnston, E. M. Gibellini, 12.VIII.1984 (PDD 45363); Little Barrier Island, vic. E. Cape, 100 in., Track 16, on Knightia excelsa, P. R. Johnston, R. E. Beever, E. P. Laracy, 16.VI.1984 (PDD 47066). TARANAKI: Mt. Egmont, Puniho Track, on Weinmannia racemosa, P. R. Johnston, G. J. Samuels, R. H. Petersen, 25.IV.1983 (Pt)D 44685); ibid., on Knightia excelsa (PDD 44684). NELSON: Mt. Dunn Walkway, on Weinmannia racemosa, P. R. Johnston, G. J. Samuels, R. H. Petersen, R. E. Beever, 20.IV.1983 (PDD 44683). WESTLAND: Fox Glacier, Minehaha Track, on Dicksonia squarrosa, P. R. Johnston, G. J. Samuels, R. E. Beever, 8.IV.1983 (PDD 44682); Haast Pass, Robinsons Creek, on Nothofagus menziesii, P. R. Johnston, G. J. Samuels, R. H. Petersen, R. E. Beever, 12.IV.1983 (PDD 43961).
North America: USA: Virginia, Dismal Creek, on Rhododendron sp., C. L. Shear, 16. X. 1935 (Holotype, BPI).
Ascocarps subepidermal. In vertical section upper stromatal layer 10-12 µm wide, of three to four layers of pale brown to hyaline cells, thin walled. rounded, 2.5-5 µm diam. Lower stromatal layer 20-30 µm wide, of three to four layers dark brown, more or less globose cells, thick walled, up to 8 µm diam. Subhymenium 20-30 µm wide, of hyaline, thin walled cells. Excipulum developing from the marginal paraphyses. Excipular elements becoming swollen, 4-5 µm diam., and closely septate near apices, septa 4-10 µm apart.
Paraphyses 2 µm diam., swollen to 4-6 µm diam. at the clavate apices, embedded in yellow gel, extending 15-20 µm beyond asci. Asci cylindric. 102-156 x 4.5-6.5(-7) µm, apex rounded wall thickened to 1.5-2.5 µm at apex with a small central pore, non-amyloid, 8-spored. Ascospores filiform, (77-)88-108 x 1-1.8 µm, tapering to one end, 0-1 septate, more or less straight when released, distinct gelatinous sheath.
CHARACTERISTICS IN CULTURE: A low percentage of the ascospores of PDD 43961 and PDD 44685 germinated on agar plates. Colonies on oatmeal agar 5-8 cm diam. after 10 weeks; aerial mycelium dense, erect, cottony, white, with small, scattered patches of black, stringy hyphae; agar not discoloured. Remaining sterile.
Ascocarps intraepidermal to subepidermal. In vertical section upper wall up to 50 µm wide, comprising a 20-30 µm wide layer of dark brown, thick walled, hyphal cells, 5-12 µm diam. Inner layer of wall lined by a 10-20 µm wide layer of crystals. A layer of periphyses, 10-15 x 2-3 µm, forms on the inside of the lower part of the wall. Lower wall 15-30 µm wide, similar in structure to the upper wall, with some hyphae extending below the wall layer amongst the host hypodermal cells. Subhymenium 5-8 µm wide, of hyaline, thin walled, more or less globose cells.
Paraphyses 1-2 µm diam., irregularly swollen and branching near the apices, capped by scattered clumps or crystals, not extending beyond asci. Asci cylindric or wider toward the base, narrowing abruptly to a short, basal foot, tapering gradually to the rounded apex, 71-76 X 5.5-6.5 pm (up to 8- 1 0 pm wide near the base), uniformly thin walled, non-amyloid, 8-spored. Ascospores filiform, 51-67 x 1-1.5 µm, 1-septate, with a small swelling (up to 2 µm diam.) above the septum, gelatinous cap at both ends.
ETYMOLOGY: dendrobii; refers to host plant.
NOTES: Distinguished from P. emarginata, the other dark walled Propolis species found in New Zealand, by ascospore shape.
Ascocarps deeply immersed in hypodermal tissue. In vertical section more or less orbicular in shape. Upper wall poorly developed, comprising 3-4 layers of hyaline, gelatinised hyphae, 2.5-4 µm diam. Inside of wall lined with a layer of unbranching periphyses, 8-10 x 2-3 µm. Crystals form amongst the periphyses and wall hyphae near the top of the margin. Lower wall not present, subhymenium 10-20 µm wide, forming directly on disintegrating host tissue.
Paraphyses 0.8-1.2 µm diam., propoloid, tangled together, intermixed with crystals near the apices, often anastomosing near bases, extending 15-20 µm beyond asci. Asci 114-147 x 7.5-9 µm, cylindric, tapering to the small truncate, or broadly rounded, apex, non-amyloid, 8-spored. Ascus wall thickened near apex, with a broad, central pore. Ascospores filiform, not tapering, often coiling on release, 80-112 x 1.2-1.5 µm, 0-1 septate, with a gelatinous cap at both ends.
CHARACTERISTICS IN CULTURE: Ascospores germinated on agar plates after 2-3 days. Colonies on oatmeal agar 2-3 cm diam. after 8 weeks; aerial mycelium white, cottony, erect; agar pink, with yellow patches near edges of colony. Remaining sterile.
ETYMOLOGY: dracophylli; refers to host plant.
NOTES: The structure of the ascocarp margin and the appearance of the ascus apex of this species are similar to those of some Ostropalean genera such as Stictis and Propolidium. However 0-1 septate ascospores with a gelatinous sheath, and anastomosing paraphyses are distinctive Rhytismataceous characters. The external appearance of the ascocarp conforms to that of Propolis.
SPECIMENS EXAMINED: New Zealand: AUCKLAND: Waitakere Ra., Walkers Bush Track, on Metrosideros robusta, P. R. Johnston, G. J. Samuels, E. H. C. McKenzie, 4.VIII. 1982 (PDD 43951); Waitakere Ra., Huia Dam Foot-bridge, on Metrosideros excelsa, P. R. Johnston, 27.IV.1982 (PDD 43952); Waitakere Ra., Kaitarakihi Beach, on Metrosideros excelsa, P. R. Johnston, 8.III.1983 (PDD 43953); ibid., 20.VI. 1983 (PDD 43958); Waitakere Ra., Cowan Track, on Metrosideros robusta, P. R. Johnston, G. J. Samuels, R. H. Petersen, 27.III.1983 (PDD 43954); Mt. Albert Research Centre, on Metrosideros sp., P. R. Johnston, V. 1983 (PDD 43955). COROMANDEL: Little Barrier Island, Titoki Point on Metrosideros excelsa, P. R. Johnston, 11.IV. 1984 (PDD 45364). NELSON: Karamea, I km. S. of Granite Rd., on Metrosideros robusta, P. R. Johnston, G. J. Samuels, R. E. Beever, 14.111.1983 (PDD 43956). TARANAKI: Mt. Egmont, Waiweranui Track, on Metrosideros robusta, P. R. Johnston, G. J. Samuels, R. H. Petersen, 25.III.1983 (PDD 43957).
Australia: VICTORIA: on Eucalyptus sp., Martin 439 (K - Holotype of Stictis emarginata Cooke & Massee).
Ascocarps subepidermal. In vertical section the immature ascocarp has an upper wall of thin walled, dark brown hyphae, lined inwardly with a layer of downward projecting, hyaline periphyses, 5-10 x 2-3 µm, with a few crystals forming in the wall near the centre of the ascocarp. The lower wall comprises 3-4 layers of dark brown, thin walled hyphae. When ascocarps open the margin is 70-100 µm wide. It comprises a 10-30 µm wide layer of brown, thin walled hyphae, 3-5 µm diam., and a 60 µm wide layer of crystals. Periphyses becoming more or less obscured by the crystals, except near the base of the wall.
Paraphyses 1.5-2 µm diam., sometimes swelling to 3 µm diam. near apex, often with short side branches, not extending beyond asci. Asci 63-100 x 6-8.5 µm, cylindric with a short, broad basal stalk, acute apex, uniformly thin walled, non-amyloid, 8-spored. Ascospores filiform, not tapering, 46-84 x 1.5-2.2 µm, 1-septate, with a gelatinous sheath.
CHARACTERISTICS IN CULTURE: Ascospores germinated on agar plates within 24 h. After 4 weeks on oatmeal agar, colonies 3.5-4 cm diam., mycelium white to pale grey, dense, cottony to felted; agar surface undulate, toward centre of colony covered with numerous fruiting bodies of the teleomorph. Ascocarps globose, dark walled, opening by irregular splits to expose the grey, pruinose hymenium. Dimensions of hymenial elements same as those found in nature. Anamorph not formed.
Sherwood's (1977) description of this fungus does not mention that the ascospores are 1-septate, and have a gelatinous sheath. These characters are found on both the type specimen, and the New Zealand collections on Metrosideros. .
Several Coccomyces and Lophodermium species are commonly found on M. fulgens, but rarely are other Rhytismataceous species found on M. robusta or M. excelsa.
New Zealand: NORTHLAND: Omahuta S.F., on Agathis australis, P. R. Johnston, G. J. Samuels, 14.IV.1982 (PDD 44690). BULLER: Buller Gorge, 11 km S. of Murchison, Fern Flat Rd, on Weinmannia racemosa, P. R. Johnston, G. J. Samuels, 16.IV. 1983 (PDD 43971).
Central America: CUBA: Cuba Fungi No. 712, Wright 1857 (isotype of Stictis foliicola - FH). PANAMA; Prov. Cocle, vic. La Mesa, on unidentified leaf, K. P. Dumont, S. E. & S. M. Carpenter, S. A. Mori, 14.VI.1975 (NY - PA315).
South America: COLOMBIA: Dpto. Cauca, El Tambo, Vereda Guazabarita, on unidentified leaf, K. P. Dumont, J. H. Haines, J. M. Idrobo, L. F. Velasquez, 15.VII.1974 (NY-CO1405). VENEZUELA: boundary Dpto. Federal and Edo. Miranda, La Silla, on Clusia sp., K. P. Dumont, J. H. Haines, B. Manara, 18.VI.1971 (NY-VE106).
Ascocarps subepidermal. In vertical section the marginal tissue comprising a few layers of hyaline, thin walled hyphae, and a poorly developed layer of periphyses. The periphyses becoming intermixed with crystals, which form a broad, wedge-shaped layer near top of margin. Subhymenium rests directly on disintegrating host tissue.
Paraphyses 1-2 µm diam., undifferentiated to slightly and irregularly swollen near the apices, often with short side branches, intermixed with scattered crystals, not extending beyond asci. Asci 72-101 x 6.5-9.5 µm, cylindric, apex broadly rounded to acute, uniformly thin walled, non-amyloid, , 8-spored. Ascospores 53-72 x 1.8-2.5 µm, not tapering, 0-1 septate, gelatinous sheath present.
CHARACTERISTICS IN CULTURE: Ascospores of PDD 43971 and PDD 44691 germinated on agar plates. Colonies on oatmeal agar 2-3 cm after 12 weeks; aerial mycelium white to slightly pink, cottony to felted; agar brownish near centre of colony. Remaining sterile.
Identification keys
Coccomyces
Propolis
Cited scientific names
- Agathis australis (D.Don) Lindl.
- Asplenium bulbiferum G.Forst.
- Astelia solandri A.Cunn.
- Beilschmiedia tarairi (A.Cunn.) Kirk
- Beilschmiedia tawa (A.Cunn.) Kirk
- Blechnum
- Blechnum discolor (G.Forst.) Keyserl.
- Clematis paniculata J.F.Gmel.
- Coccomyces clavatus P.R. Johnst. 1986
- Coccomyces crystalligerus Sherwood 1980
- Coccomyces cupressini P.R. Johnst. 1986
- Coccomyces globosus P.R. Johnst. 1986
- Coccomyces lauraceus P.R. Johnst. 1986
- Coccomyces libocedri P.R. Johnst. 1986
- Coccomyces limitatus (Berk. & M.A. Curtis) Sacc. 1889
- Coccomyces longwoodicus P.R. Johnst. 1986
- Coccomyces phyllocladi P.R. Johnst. 1986
- Coccomyces radiatus Sherwood 1980
- Cordyline
- Dacrydium cupressinum Lamb.
- Dendrobium cunninghamii Lindl.
- Dicksonia squarrosa (G.Forst.) Sw.
- Dracophyllum
- Dracophyllum latifolium A.Cunn.
- Dracophyllum longifolium (J.R.Forst. & G.Forst.) R.Br.
- Elaeocarpus dentatus (J.R.Forst. & G.Forst.) Vahl
- Halocarpus biformis (Hook.) Quinn
- Knightia excelsa R.Br.
- Leptospermum scoparium J.R.Forst. & G.Forst.
- Libocedrus bidwillii Hook.f.
- Litsea calicaris (A.Cunn.) Kirk
- Metrosideros
- Metrosideros excelsa Gaertn.
- Metrosideros fulgens Gaertn.
- Metrosideros robusta A.Cunn.
- Nestegis lanceolata (Hook.f.) L.A.S.Johnson
- Nothofagus menziesii (Hook.f.) Oerst.
- Phyllocladus
- Phyllocladus alpinus Hook.f.
- Polystichum
- Propolis dendrobii P.R. Johnst. 1986
- Propolis dracophylli P.R. Johnst. 1986
- Propolis emarginata (Cooke & Massee) Sherwood 1977
- Propolis quadrifida (Lév.) Mont. 1854
- Pseudopanax
- Rubus cissoides A.Cunn.
- Weinmannia racemosa L.f.