Type: Sooty Moulds and Similar Fungi; Description: Subiculum superficial, vary variable, dark brown to black; on leaves and twigs; effuse and sparsely to densely velutinous on leaves; spongy and up to 7 mm wide when encircling twigs. Mycelium composed of repent and erect, pale brown to dark brown, septate, smooth to slightly warted moniliform hyphae, up to 23 μm wide, which taper toward their distal ends; erect hyphae 75 μm or more long, simple to frequently branched. Ascomata stromatic, scattered and few in number, partly immersed, brown to dark brown, subglobose to ellipsoidal with a short stalk-like base, 50–170 μm in diameter, ostiolate, bearing toward the apex numerous simple, pale brown to dark brown, moniliform, straight or slightly curved hyphal appendages up to 90 μm long. Asci fasciculate, broadly ellipsoidal to clavate, 45–90 × 15–30 μm. Ascospores ellipsoidal, slightly curved, 3-septate (rarely 3–5-septate), 16–26 × 7–11 μm, pale brown to dark brown. Capnobotrys synanamorph. Conidia ovoid to reniform, 1-septate, 11–16 × 8–11 μm, smooth, brown to very dark brown. Capnosporium synanamorph. Conidia ellipsoidal to obovate, 2–3-septate, 18–35 × 8–15 μm, smooth, brown to dark brown. Capnophialophora synanamorph. Conidia subglobose to ellipsoidal, 0-septate, 1.4 × 1.2 μm, hyaline.

Distribution: Northland, Auckland, Waikato, Taupo, Nelson, Buller, Westland.; 1st Record: Hughes (1981b).

Significance: This species is the commonest metacapnodiaceous sooty mould on leaves in New Zealand. All published records are from dicotyledonous plants and ferns.; Host(s): Aristotelia serrata, Beilschmiedia tawa, Blechnum discolor, B. procerum, Brachyglottis repanda, Coprosma areolata, C. grandifolia, C. lucida, C. tenuifolia, Cyathea dealbata, C. smithii, Dicksonia squarrosa, Elaeocarpus dentatus, E. hookerianus, Hedycarya arborea, Hymenophyllum sp., Leptospermum scoparium, Leucopogon fasciculatus, Melicytus ramiflorus, Myrsine australis, Nothofagus fusca, Olearia furfuracea, O. rani, Pittosporum ellipticum, Pseudowintera axillaris, Quintinia serrata, Raukaua edgerleyi, Rubus australis, R. cissoides, Sticherus cunninghamii, Weinmannia racemosa.

COLLECTIONS: (1) Aristotelia serrata, Wellington Province, Mt Ruapehu, Ohakune Track, 16.XII. 1966, J. M. Dingley, PDD 25874 (DAOM 117143b); (2) Beilschmiedia tawa twigs. North Auckland, Waipoua Forest, 18.Vi. 1963, J. M. D. & S. J. H., DAOM 96736b; (3) Blechnum discolor, Westland, Granville Forest, Orwell Creek, Ahaura, 3.IV.1963, S.J.H., PDD21418 (DAOM 105689);(4) Blechnum minor, Westland, Lower Poerua River, Harihari, 5.IV. 1963, J.M.D., DAOM 93414a; (5) Blechnum procerum, Auckland Prov., Titirangi, 22.1.1963, S.J.H., DAOM 93350b; (6) Coprosma areolata, Westland, Little Wanganui River, Harihari, 6.1V. 1963, S.J.H., DAOM 105682; (7) Coprosma australis, Auckland Prov., Upper Piha Valley, Waitakere Range, 30.XI. 1966, J.M.D., PDD 25878 (DAOM 117153b); (8) Coprosma lucida, Auckland Prov., Kitekite Stream, Piha Valley, 31,1.1963, S.J.H., DAOM 93348b; (9-12) Coprosma tenuifolia, (9) Auckland Prov., Pureora, 21.111.1963, S.J.H., DAOM 93420c; (10-12) Wellington Prov., (10) Tongariro National Park, Mahia Camp site, 5.111.1963, S.J.H., DAOM 93377a; (II) Mt Ruapehu, Ohakune Road (670 m), 16.XII. 1966, J.M.D., PDD 25876 (DAOM 117151b); (12) Wellington Prov., Mt Ruapehu (610 m), 16.XII. 1966, J.M.D., PDD 25871 (DAOM 117157b); (13,14) Cyathea dealbata, (13) Auckland Prov., Comwallis, 3.1.1963, S.J.H., DAOM 93366b; (14) North Auckland, Parahaki Mt, Whangarei, 21.VI.1963, S.J.H., DAOM 93444b; (15) Cyathodes fasciculata. North Auckland, Parahaki Mt, Whangarei, 21.VI. 1963, S.J.H., DAOM 105690; (16) Dicksonia squarrosa, Westland, Lower Poerua River, Harihari, 5.1V. 1963, S.J.H. ,PDD21385 (DAOM 105681); (17) Elaeocarpus dentatus, Wellington Prov., Tongariro National Park, Ohakune, 6.111.1963. S.J.H., DAOM 93386b: (18) Elaeocarpus hookerianus, Westland, Granville Forest, Orwell Creek, Ahaura, 2.1V. 1963, S.J.H., PDD 21413 (DAOM 105688); (19) Gleichenia cunninghami, Wellington Prov., Horopito, 8.111.1963, S.J.H., DAOM 93416d; (20) Hemitelia smithii, Wellington Prov., Tongariro National Park, Erua, 6.111.1963, S.J.H., PDD 21422 (DAOM 93419b); (21) Leptospermum scoparium, Auckland Prov., Pureora, 22.111.1963, S.J.H., PDD 20741 (DAOM 96438b); (22,23) Melicytus ramiflorus, (22) Nelson Prov., Eve's Bush, Waimea, 1.111.1967, J.M.D., PDD 25887 (DAOM 117321); (23) Auckland Prov., Piha, Waitakere Range, 31.V11.1963, S.J.H., DAOM 106808; (24) Myrsine australis, Auckland Prov., Comwallis, 3.1.1963, J.M.D. and F. J. Morton, PDD 20416 (DAOM 93371b); (25) Nothofagus fusca, Westland, Granville Forest, Orwell Creek, Ahaura, 3.IV. 1963, S.J.H., DAOM 105684a; (26) Olearia furfuracea, Auckland Prov., Kitekite Stream, Piha Valley, 31.1.1963, S.J.H., DAOM 93347b; (27) Olearia sp.. Nelson Prov., Eve's Bush, Waimea, 1.111.1967, J.M.D., PDD 25885 (DAOM 117345); (28) Pittosporum ellipticum, Auckland Prov., Upper Piha Valley, Waitakere Range, 30.XI.1966, J.M.D., PDD 25881 (DAOM 117322a); (29) Pseudopanax edgerleyi, Westland, Granville Forest. Orwell Creek, Ahaura, 3.1V. 1963, S.J.H., DAOM 105683; (30) Quintinia serrata, Westland, Swamp Forest, Harihari, 7.1V. 1963, S.J.H., DAOM 93415c; (31) Rubus cissoides, Westland, Little Wanganui River, Harihari, 6.1V. 1963, S.J.H., DAOM 93410a; (32,33) Weinmannia racemosa. Wellington Prov., (32) Tongariro National Park, Erua, 6.111.1963, S.J.H., DAOM 93480b; (33) Mt Ruapehu, Ohakune Track (610m), 16.XII. 1966 J.M.D. PDD 25889 (DAOM 117164c).
Six collections from New Zealand and two from Chile, including Juan Fernandez, preserved in Herb. K, S, and UPS, also bear Metacapnodium moniliforme.
New Zealand: "Antennaria Robinsonii B. & M. on living leaves of Brachyglottis repanda [scr. M. C. Cooke], b. 197. New Zealand, Rev. W. Colenso--Rec'd Dec., 1885". (K). Antennaria Robinsonii, New Zealand. M. C. Cooke, May '86 (scr. M. C. Cooke]. On Drimys axillaris, 294, near Wellington. 30.X1.85 T. Kirk. [scr. T. Kirk]". (K). "Swedish Botanical Australasia-Expedition. New Zealand, North Island, Ruahine, Cook Bot. Distr. York Bay, near Wellington . . . on adult leaves of Elaeocarpus dentatus, 7.XII.1926, No. 1012c. leg. G. Einar et Greta du Rietz". (UPS). "Antennaria Robinsonii Mont., New Zealand, [on Hymenophyllum] Colenso 2761 [scr. Berkeley]". (Herb. Berk. in K). "Antennaria Robinsonii. On living [leaves of] Olearia colorata. Dry Forests, [scr. M. C. Cooke], b. 383. New Zealand, Rev. W. Colenso.-Rec'd Dec., 1885". (K). "Antennaria Robinsonii, New Zealand. M. C. Cooke, May '86 [scr. M. C. Cooke]. On. Rubus australis, near Wellington. 30.XI, 85 T. Kirk. 293 [scr. T. Kirk]". (K). "Antennaria Robinsonii. New Zealand. M. C. Cooke May '86 [scr. M. C. Cooke]. Leaves of Drimys and Hedycarya infested with a Lecanium, the latter parasitized by a Chalcid or other insect, and both insects infested by a fungus. Near Wellington. T. Kirk. 298. [scr. T. Kirk]". (K).
Chile: Svenska Pacific expeditionen, 1916-17. "Limacinia (fernandesiana Neger) = scoriadea (Berk.) Keissler". On Myrceugenia fernandeziana. Juan Fernandez, Chile: Mas a Tierra. 250 m.s.m. 30.111.1917. Coil. Carl & Inga Skottsberg. Det. K. Keissler (1928). (S). "Flora Chilensis. Nr. 1995. [Chile] Llanquihue: Puerto Varas, Ensenada. In the low, open forest. 11 Jan. 1947. 60 m.s.m. Leg. Benkt Sparre". (UPS).

Subicula dark brown to black, superficial, very variable, effuse and thinly to densely velutinous on leaves to spongy and up to 7 mm wide when encircling twigs. Mycelium composed of repent and erect, pale brown to dark brown, anastomosing, septate, moniliform, smooth to slightly roughened hyphae which taper toward their distal end. Upright hyphae may be as short as 75 µm but are usually longer, simple to frequently branched with the branches scattered and arising singly (never in pairs) more or less at right angles to the parent hypha and then curving upwards. Hyphal cells are generally as broad as or broader than long, being broadly barrel-shaped and up to 23.5 µm wide; the younger distal cells are as narrow as 5 µm. Dense subicula on leaves and especially on twigs may bear compact fascicles of erect hyphae up to 800 µm tall.
Metacapnodium teleomorph. Ascostromata generally scanty, partly immersed in the subiculum, scattered, brown to dark brown, subglobose to ellipsoidal to broadly obovoid with a short stalk-like base, 90-230 µm high x 66-165 µm wide with numerous rooting hyphae at the base, ostiolate (sometimes on a short papilla) at maturity, glabrous or bearing toward the distal end up to 23 simple, subulate, pale brown to dark brown, straight or slightly curved hyphal appendages up to 90 nm long, up to 16 µm wide at the base and as narrow as 4.5 µm wide at the apex. Asci fasciculate, bitunicate, usually 8-spored, variable in size and shape and number in the ascostroma, broadly ellipsoidal to clavate, 45-90 x 15-30 µm. Ascospores crowded to biseriate to irregularly so, ellipsoidal and straight at First but finally inequilateral to slightly curved, pale brown to dark brown and slightly paler at the ends, usually 3-septate, sometimes 4-septate, uncommonly 2- or 5-septate, at first not constricted at the septa but moderately so especially after expulsion, with the convex wall thicker and appearing more deeply pigmented than the flattened or slightly concave wall: at maturity a circular paler and thinner area is found in the wall of the end cells just below the apex and above the base on the flattened or slightly concave side. Ascospores vary considerably in size and they may acquire a pigmented wall when they are as short as 15 µm long: within 8-spored asci the ascospores are commonly 16-26 µm long and 7.2-11 µm wide. Expelled ascospores, or those within asci in which only some of the initials have developed fully, or those' which have for some reason not been expelled, are often up to 35 µm long, or even up to 40 µm long and up to 12.5 µm wide. The fourth or fifth septum, when these are formed, are found in the terminal cells of the ascospore. Expelled ascospores germinate by producing a hypha or one or two Capnophialophora phialides terminally or laterally from one or both end cells. However, these structures do not develop, as one would expect, at the paler and thinner areas of the wall of the terminal cells. Over these paler areas of some expelled ascospores an outer wall layer appears to gelatinise to produce an irregular to more or less cylindrical appendage up to 7 µm long but usually shorter: perhaps such appendages have an adhesive function. An inner wall layer projects slightly beyond the contour of the original wall but no further development was seen at these sites.
Capnobotrys synanamorph. Distinctive conidiophores are lacking. Conidiogenous cells are borne in compact clusters or series on the lower or upper surface of slightly curved or abruptly bent ends of erect hyphae and of usually downwardly curved lateral branches. These fertile branches are 3-to 6-septate, usually occur singly and bear conidiogenous cells usually on the lower surface, directly on the distal cells and on 1- or 2-celled stalks on the proximal cells. Similar clusters of conidiogenous cells are borne on the lower surface of bent ends of separate erect hyphae. On fasciculate hyphae, which may be up to 800 µm long, conidiogenous cells are usually borne on the upper surface of the curved ends: such long fasciculate hyphae bear very few lateral fertile branches. Conidiogenous cells are subglobose to broadly ellipsoidal, smooth, pale brown to brown, becoming somewhat obpyriform by extension at the apex during successive conidium production: they are 5.5-9 µm wide and may reach a length of 10.8 µm with the distal end bearing up to 5 crowded denticulate scars after schizolytic conidium secession. Conidia are produced blastically on successive new growing points; initials are at first globose but soon elongate unilaterally so that mature conidia are borne at right angles or obliquely to the point of attachment. Maturing conidia are dry, ovoid, inequilateral, medially or supramedially 1-septate, smooth, brown to very dark brown with the proximal (basal) cell usually broader and slightly longer than the distal cell, with a minute circular and inconspicuous oblique or lateral scar: the distal cell has an apical, lateral or obliquely located, paler, thinner-walled area. Conidia usually secede in this condition or they may increase in size with both cells becoming more or less equal or the conidia may become reniform (in side view) with the convex wall markedly thicker than the flattened or concave wall which may be extended into two blunt projections, one bearing the conidium scar and the other the paler, thinner-walled area. Subsequent development of the larger conidia, usually on the surface of the host, results in a cracking of the outer wall so that the conidium appears irregularly warted. Uncommonly, the wall over the paler and thinner area shows some gelatinisation and an inner wall may project slightly. In the smooth, more or less ovoid condition conidia measure (9-) 11.5-16.2(-18) x (5-) 8-11 µm but the reniform conidia in side view can be up to 25 µm long and up to 16 µm wide (projections included).
Capnosporium synanamorph. Distinctive conidiophores are lacking. Conidiogenous cells are the terminal cells of otherwise undifferentiated erect hyphae. Solitary conidia are produced at an apparent pore at the apex of the conidiogenous cell: the larger conidia may produce another but smaller conidium at the apex. Occasionally two conidia may develop directly on a conidiogenous cell and less frequently the conidiogenous cell or a cell below will proliferate and produce another conidium on the 1- to 3-celled extension. Conidia are dry, straight, ellipsoidal to obclavate to obovoid, smooth, thick-walled, brown to dark brown, (l-)2- or 3(-5)-septate, constricted at the septa, readily seceding, rounded at the apex and at the base which bears a minute scar. They measure 13.5-18 x 10-10.8 µm (1-septate), (16.2-) 18-27 x 8.1-12.6(-14.4) µm (2-septate), (19-) 25-34(-36) x (9-)10-14.5 µm (3-septate), 39-45 x 12.6-14.5 µm (4-septate), and 46 x 14.5 µm (one 5-septate conidium).
Capnophialophora synanamorph. Distinctive conidiophores are lacking. Phialides are produced singly or in irregular clusters of 2 to 4 at the apex and laterally on otherwise undifferentiated erect hyphae and on their short branches. They have also been seen produced directly on expelled ascospores, on detached Capnobotrys conidia, less commonly on Capnosporium conidia or on hyphae derived from these. The venter is usually pale brown to brown, smooth or slightly roughened, subglobose, 3.9-5.5 (-6.5) µm wide with a single subhyaline to pale brown, well differentiated collarette 2.7-4.3 µm long and 2.8-3.2 µm wide which shows a deep constriction where it joins the venter. Phialides with two collarettes are rare. The venter of some terminal and adjacent phialides is much broader (up to 8.1 µm) than that of others and the collarette is somewhat narrower and shorter. Collarettes are at first ellipsoidal with a rounded apex, but the apex is opened presumably by lysis and the collarette finally appears funnel-shaped. Phialoconidia are hyaline, minute and apparently scanty. The first is produced enclosed within the lumen of the unopened collarette and is ellipsoidal and c. 2 x 1.4 µm, subsequent phialoconidia are subglobose to broadly ellipsoidal and c. 1.4 x 1.2 µm.

On living leaves of dicotyledons and ferns and less frequently on twigs

NOTES: Metacapnodium moniliforme is the commonest metacapnodiaceous sooty mould on leaves in New Zealand and is of particular interest because it produces three easily distinguishable synanamorphs. In the 33 recent collections cited above, Capnobotrys is accompanied by Capnophialophora in 30 collections and by Capnosporium in 27. The Metacapnodium teleomorph is present in 11 collections.
The identity of the teleomorph and synanamorphs is claimed not merely because of the frequency with which the morphs are closely associated but because hyphal connections have been traced several times from one morph to the other. Furthermore, Capnobotrys has been seen produced on a hyphal appendage of the teleomorph, and a short hyphal branch bearing Capnobotrys has been seen extending from a terminal conidiogenous cell bearing a Capnosporium conidium and also extending from other cells of erect hyphae, which bear Capnosporium. Ascospores and Capnobotrys conidia, and less frequently Capnosporium conidia may germinate to produce Capnophialophora phialides directly on these spores or on a short germination hypha.
Fraser (1935) included 16 collections in her account of Capnodium moniliforme: 15 of these are from Australia on various flowering plants and a fern and 1 from New Zealand on a conifer, Podocarpus hallii.
Parts of the type collection, on leaves of Backhousia myrtiflora, Salisbury, New South Wales, V. 1934 (DAR 19740, and BPI), have been examined (Fig. 11). Young ascostromata, with or without hyphal appendages were seen but no ascospores. However, the hyphae, Capnobotrys conidiogenous cells and conidia, and Capnophialophora phialides in the type are as described and illustrated above from the New Zealand collections. Also found in the type were some hyphae with an apparent pore at the apex, but no Capnosporium conidia were seen. Fraser described the ascospores as "averaging 21 x 9 µm., but varying considerably with age from 18 x 9 to 31.5 x 11 µm." Capnobotrys conidia "finally thick-walled and very dark" and "10-20 x 7-10 µm". The New Zealand collections fit well. Another collection cited by Fraser, on Polystichum aculeatum, Mt Wellington, Tasmania, 1.1933, coil. D. Martin (DAR 19733, and BPI), bears abundant Capnobotrys conidia ranging from ovoid to reniform and up to 21.5 x 14.5 µm (projection included); abundant Capnosporium (moniliforme Hughes) conidia in this collection are 21.6-27 x 10-13 µm (2-septate), 28.8-36 x 10.8-14.4 µm (3-septate) and a solitary 4-septate conidium is 43 x 14.4 µm. I have failed to find pyenidia in Metacapnodium moniliforme as reported by Fraser (1935) and Batista & Ciferri (1963).
Capnophialophora phialoconidia are minute and apparently not produced in abundance. They can be seen readily within the collarette by mounting material in a solution of haematoxylin in distilled water.
Of interest is the apparent preference of Metacapnodium moniliforme for non-coniferous hosts. All the collections cited above are on dicotyledonous flowering plants and ferns. I have not seen the collection upon which Fraser (1935) based the record of this species on Podocarpus hallii.
Unlike most of the cells of the subiculum the upper cells of fasciculate hyphae are usually longer than broad: in this feature they resemble the cells of synnemata in Capnocybe (Hughes 1966) and the distal cells of the longer Capnobotrys-bearing erect hyphae of C. laterivecta, C. paucispora, and of Metacapnodium dingleyae.