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Volume 84, 1956-57
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Australasian Cyttariaceae

[Received by the Editor, October 27, 1955.]

Abstract

This paper describes two new species of Cyttaria, parasitic on Nothofagus menziesii (Hook. f.) Oerst., in New Zealand, as C. nigra and C. pallida. C. gunnii Berk, is redescribed and notes are given on the Australian species C. septentrionalis Herbert.

The origin and taxonomic importance of certain anatomical features are discussed, particular attention being focused on elevations surrounding openings in the surface of the stroma, here termed “papillae”. A table of host relations for the 10 species of Cyttaria and 11 species of Nothofagus attacked is provided. Stress is laid on gall characters in distinguishing species. Special points of interest in the New Zealand species are noted, and economic considerations in connection with Cyttaria in forests are enumerated.

I. Introduction

In the course of an investigation, carried out in 1946, into the fungi associated with Nothofagus menziesii (Hook. f.) Oerst. it was noted that the trees were parasitised by three Ascomycetes belonging to the family Cyttariaceae. Only one species of this family having been recorded previously from New Zealand, the identity of the other two became a matter of considerable interest. From a study of the relevant literature it became evident that these species were undescribed. The Cyttariaceae, a family of Inoperculate Discomycetes, contains a single genus, Cyttaria, all species of which form galls on members of the genus Nothofagus.

The genus Cyttaria was erected by Berkeley, in 1841, to contain C. darwinii and C. berteroi, two species collected in South America by Charles Darwin in 1834. Berkeley also described C. hookeri in 1847 and C. gunnii in 1848. Fischer separated C. harioti from C. darwinii with which species it appears to have been included under Berkeley's description of the material collected by Darwin. C. espinosae was described by Lloyd in 1917; C. septentrionalis by Herbert in 1930 and C. johowii by Espinosa in 1940.

Palm (1932) cited Skottsberg as the authority for the description of three forms of C. darwinii, f. vulgaris, f. bridgesii, and f. onea; Santesson (1945) considered f. onea to be synonymous with C. harioti Fisch. Marchionatto (1940) described three forms of C. hookeri. f. typica, f. moroidea and f. Candida. The f. Candida he considered to be near to, if not identical with, C. intermedia Palm, which is itself a synonym for C. hookeri.

In 1932 Palm erected the genus Cyttariella to contain forms of which only the conidial stage was known, and included two species, C. deformans and C. skottsbergii; Santesson (1945) considered both these to be young forms of Cyttaria hookeri.

In spite of the confusion between the different species there has been comparatively little synonymy. C. reichii Henning is a synonym for C. berteroi Berk., C. alba Philippi (1869) is a nomen nudum, as is C. purdi [ unclear: ] i Buchanan (1885); the host record of Nothofagus fusca for the latter thus becomes inadmissible. C. disciformis Lev. is not a Cyttaria and C. intermedia Palm is a synonym for C. hookeri Berk.

Distribution and Host Relations

With the exceptions of C. gunnii in Tasmania, Victoria and New Zealand and C. septentrionalis in Queensland and New South Wales, all the described species are from South America.

Of the species of Nothofagus in the southern hemisphere, eight South American species are attacked by one or more species of Cyttaria, but only three of the Australasian species are known to be susceptible.

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Recorded host-parasite relations are given in the following table:—

[The section below cannot be correctly rendered as it contains complex formatting. See the image of the page for a more accurate rendering.]

Table of Host-parasite Relations.
Cyttaria Nothofagus
1. C. darwinii Berk. N. betuloides (Mirb.)
Oerst.
N. antarctica (Forst.)
Oerst.
N. pumilio (Poepp. et Endl.)
Krasser
2. C.harioti Fisch. N. betuloides
N. antarctica
N. pumilio
N. dombeyi (Miro.) Oerst.
N. nitida (phil.) Krasser
3. C. hookeri Berk. N. betuloides
N. antarctica
N. pumilio
N. dombeyi
4. C. johowii Espinosa N. dombeyi
5. C. berteroi Berk. N. obligua (Mirb.) Oerst.
N. glauca (phil.) Krasser
6. C. espinosae Lloyd. N. glauca
N. obliqua
N. procera (Poepp. et Endl.)
Oerst.
7. C. gunnii Berk. N. menziesii (Hook. f.)
Oerst.
N. cunninghamii (Hook. f.)
Oerst.
8. C. nigra n. sp. N. menziessii
9. C. pallida n. sp. N. menziesii
10. C. septentrionalis Herb. N. moorei (F. Muell.)
Krasser
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No Cyttaria has been recorded on N. fusca (Hook. f.) Oerst., N. truncata Cock., N. cliffortioides (Hook. f.) Oerst., and N. solandri (Hook. f.) Oerst. in New Zealand, or on N. gunnii (Hook. f.) Oerst. in Tasmania. N. glauca is probably merely a variety of N. obliqua. Other varieties of N. obliqua sometimes referred to as species are var. leoni Espinosa and var. macrocarpa D.C.

N. alessandrii Espinosa and N. montagnei Reiche are of uncertain status and are possibly hybrids.

Economic Considerations

In dense natural forest Cyttaria attack does little damage, trees attacked in the bole die through suppression, and galls on small branches are shed with the branches before the fungus becomes properly established. N. menziesii has the ability to cast small branches through the formation of an abscission layer.

When forests are put under management the elimination of infected trees becomes an important consideration in marking for thinning. Too heavy thinning could favour the retention of branches and of cankered trees and so allow the fungus to develop and spread, with a corresponding increase in the proportion of malformed stems.

The greatest incidence of Cyttaria is on heavily branched trees growing on the borders of clearings or in open stands. On such trees the branches are not shed to the same extent as in dense stands, and trees weakened by attack are not subject to suppression, consequently the galls persist indefinitely. Difference in susceptibility is most marked; one tree may be heavily attacked while the surrounding trees, with interlacing branches, are quite free of the disease. In New Zealand two species may occur on the same branch, but all three have not yet been observed on the same tree.

The galls and stromata (fruit bodies) are of commercial value in South America, the galls for making ornaments and the stromata were used for food by the Indians. No such use has been made of Cyttaria in this country, although the stromata are edible. C. gunnii has a pleasant flavour, C. nigra is somewhat bitter and smells of dried apricots or peaches, C. pallida is tasteless and is very unappetizing in appearance. The stromata are eaten by opossums and pigeons.

II. General Characters

1.Gall formations. (Plate 11.)

The method of infection of the host by Cyttaria is not known, but it appears that hyphae from a germinating spore invade the tissues of tender new growth. According to Janet M. Wilson (1937) hyphae become intracellular and control the growth of the host tissues.

All species of Cyttaria produce galls on species of Nothofagus; the galls are perennial and from them arises the annual crop of stromata.

Two types of gall are produced, globose galls and longitudinal galls. With globose galls, such as are produced by C. [ unclear: ] arwinii, C. harioti, C. espinosae, C. gunnii and C. nigra, the mycelium does not travel any great distance in the host and consequently only local malformations are produced.

With longitudinal galls, on the other hand, the mycelium travels down the stem or branches and the malformation increases steadily in length for many years. Longitudinal galls are produced by C. hookeri, C. berteroi, C. septentrionalis and C. pallida.

The characteristic formation of galls produced by the four Australasian species enables identification to be made in the absence of fructifications. The characters used are: gall type, relative size of gall to branch, and the shape of spiny protrusions from the woody tissue beneath the bark.

While insufficient material has been obtained to place C. septentrionalis exactly, the following key will enable the New Zealand species to be differentiated.

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Key Based on Gall Characters
1.Gall of globose type 2
Gall of longitudinal type 3
2.Height greater than diameter, spines short, few, on dome-shaped projections C. nigra
Height less than diameter, spines long, numerous, not on projections C. gunnii
3.On N. menziesii C. pallida
On N. moorei C. septentrionalis

From the information available, the genus can be divided into the following two groups:—Species with pycnidia are marked (p) and the most primitive are placed first.

Group 1. Longitudinal Galls

  • 1.C. hookeri (p).

  • 2.C. berteroi.

  • 3.C. pallida.

  • 4.(?) C. septentrionalis (p?)

Group 2. Globose Galls

  • 5.C. darwinii (p).

  • 6.(?) C. joho [ unclear: ] ii (p) (galls not described).

  • 7.C. nigra (p?).

  • 8.C. espinosae.

  • 9.C. harioti.

  • 10.C. gunnii.

2.The stroma or fructification

The stroma is compound and is apparently formed through the fusion of numbers of simple stromata each bearing one apothecium.

The stromata arise from the spines produced in the wood of the host by the action of the parasite. These spines, consisting of fungal hyphae and woody tissue, project from the hypertrophied woody growths and extend through the bark nearly to the exterior. As the stroma enlarges it assumes a more or less spherical form with an oval apical portion and conical basal portion, it is jelly-like or rubbery in consistency and is covered by a tough skin or cortical layer. Apothecia are formed beneath this layer, which becomes tightly stretched and ruptures at maturity. The medulla or central portion of the stroma is a mass of hyphae through which cartilaginous fibres run from the point of attachment and ramify towards the cortex.

The main features in the structure of the stroma, many of which are useful for purposes of identification, are as follows:—

(a) The External Cortical Layer.

The stroma is covered by a tough elastic skin or pseudoperidium which may be dry (as in C. gunnii) or hygroscopic, (as in C. pallida) or covered with black incrustations of carbonaceous material (as in C. nigra and C. septentrionalis). It may be perforated, between the apothecia by openings raised on various types of papillae. At maturity the cortex covering the mouths of the apothecia becomes membranous, tightly stretched, and finally ruptures, rolling back in a characteristic manner. The cortex of the basal portion of the stroma is usually striated, fluted or flattened, but not perforated by apothecia.

(b) The Apothecia.

The hymenial layer arises in hollows which form within the stroma beneath the cortex, and between the papillae and internal fibres in species in which these occur. During their formation the apothecia are filled with a very clear, highly refractive, mucilaginous substance which disappears some time before the spores are ripe. The hymenium consists of asci and paraphyses arising from a sub-hymenial layer of

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swollen, septate hyphae, and lines the whole of the apothecium except for the part beneath the cortex.

The ascus is eight spored, and behind each spore there is a plug or wad. When ripe the spores occupy about half of the ascus and, if fresh material is mounted in water, the ripe asci explode violently and discharge the spores. In water the spores are liable to burst. The spores are dark, sub-globose, and measure about 12μ in diameter. They are of little value, at present, for purposes of identification because they show little difference between species, but records show great variation in size, probably depending on the method of mounting and state of maturity. For example, spores of C. hookeri are variously given as 10 × 15; 12–14 × 10–13; and 16 × 19μ; similar discrepancies occur in descriptions of other species. Paraphyses are septate, branched, usually expanded at the tip and often brightly coloured. Spores are discharged in visible clouds from ripe asci. Spore discharge may be demonstrated readily

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Fig. 1.—L.S. of young stromata. 1. Cyttaria gunnii. 2. Cyttaria nigra. 3. Cyttaria pollida. 4. Cyttaria moorei. M, medulla; P, papilla; C, cortical layer; A, apothecium; F, fibres; T. cartilaginous tube; I, incrustations.

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by breathing on the ripe fruit bodies; the spore print is black in all New Zealand species.

(c) Medulla.

In the young stroma the central part is occupied by an interwoven gelatinous mass of hyphae. The hyphae become directional near the internal fibres and around the apothecia. At maturity, in most species, the central mass disintegrates and the space beneath the apothecia becomes filled with air. C. berteroi, C. espinosae and C. hookeri are said to remain solid and have no central cavity.

(d) The Internal Fibres.

Characteristic internal structures have been variously termed “nerves” or “fibres”. It is here considered that they represent remnants of the cortex of the individual apothecia enclosed during the fusion into a compound stroma. The term “fibres” is retained.

Running up from the point of attachment through the short basal portion is a tube of cartilaginous mycelium from which branches radiate and pass between the apothecia to the cortex. In New Zealand species the tube enlarges to a mushroom-shaped head in the centre of the stroma and the fibres radiate from this. In other species the fibres anastomose; in C. septentrionalis they appear to be flattened and anastomosing. In other species again the fibres appear to be lacking. The central tube may pull out when the stroma drops from the tree; this frequently happens with C. nigra. The fibres may be white and shining or variously coloured; they end just before reaching the cortex, where they expand to form cavities, coinciding, approximately, with similar cavities formed beneath the papillae.

(e) The Papillae.

Papillae, which provide a feature of considerable diagnostic value, arise between the apothecia (but before these are formed) and at, or near, the ends of the internal fibres. They surround a pore opening to the surface from a cavity beneath the cortex.

In the few examples of C. septentrionalis available for study, papillae are absent, but where the fibres reach the cortex small simple holes may be seen.

In C. gunnii the papillae are situated in shallow depressions and barely reach to the level of the general surface. In C. pallida papillae form very pronounced elevations in young specimens and are sharply conical, while in C. nigra they are large and dome-shaped.

The presence and diagnostic value of papillae appears to have been largely overlooked, although Fischer (1888) records that the fibres (nerves) “break through the skin” in C. darwinii. He also records similar “perforations” in C. hookeri.

It is possible to identify the New Zealand species from the papillae alone even in very small young specimens. The function, if any, of papillae is not clear, although they may serve to admit air through the cortex to the fibres during the period of rapid increase in size, just prior to maturity and may by the same means, prevent collapse during dry periods. Another function may be the production of the black substance in young stromata of C. nigra or of slime in C. pallida.

(f) Pycnidia.

Berkeley (1841) recorded the presence of black granules on the base of C. darwinii stromata which he described as “granulated like shagreen”. Fischer (1888) interpreted these granules as being ‘spermogonia’ while others have referred to them as pycnidia or pycnoconidangia. They are small, black spherical bodies measuring from 95 × 90μ in C. harioti to 180 × 264μ in C. johowii; they are superficial or immersed in the pseudoperidium. They are present in C. hookeri but absent from C. berteroi. C. espinosae is covered with a black substance when young but is said not to produce pycnidia. Similarly C. septentrionalis and C. nigra are covered by black granulations when young; these are taken to be pycnidia, but the condition

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Cyttaria gunnii on Nothofagus menziesii. (Photo by the late W. C. Davies, of the Cawthron Institute.)

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A. Cyttaria pallida on branch of Notholagus menziesii. (Photo A. R. Teixeira.) B. Cyttaria nigra, immature stromata to show papillae and gall on branch of Nothofagus menziesii. (Photo A. R. Teixeira.) C. Mature stromata of Cyttaria nigra. (Photo N. Z. F. S.)

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Decorticated Cyttaria
galls on Nothofagus menziesii. A. Cyttaria nigra. B. Cyttaria pallida. C. Cyttaria gunnii.

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A. Spindle-shaped galls and Cyttaria stromata on twigs of Nothofagus moorei from Mt. Hobwee, Queensland.
B. Gall of Cyttaria pallida on Nothofagus menziesii sapling. C. Galls of Cyttaria gunnii on Nothofagus menziesii sapling. One gall decorticated. (Photo N. Z. F. S.)

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may prove to be similar to that of C. espinosae and the correct interpretation will require a critical comparison with the South American species.

In C. gunnii and C. pallida no black incrustation occurs and pycnidia are considered to be definitely absent.

No satisfactory expalanation for the presence or function of pycnidia in Cyttaria has yet been offered.

III. Description of the Species
Genus Cyttaria

Cyttaria

Berkeley, in Trans. Linn. Soc. London, Vol. 19, p. 40, 1841.

Berkeley erected the genus Cyttaria on the following characters:—

Receptacula carnoso-gelatinosa in stroma commune subglobosum, epiderme crassiuscula vestitum, aggregato; basi stipitiforma granulata. Cupula peripherica, primo clausa, gelatina distenta, demum epidermide rupta aperta. Hymenium, margine excepto, separabile. Asci ampli, distincti, demum liberati, paraphysibus immixtis. Velum persistens demum ruptum, margine plus minus reflexo. Sporidia pallida.”

Massee (1910) also defines Cyttaria.

Berkeley's diagnosis of Cyttaria gunnii, in Hooker's London Journal of Botany, 1848, p. 576, reads as follows: “Globoso-piriformis, demum cava, basi attenuata, nec stipitiformis, nec scabra, loculis parvis, poro amplo saepe irregulari dehiscentibus; ascis cylindraceis breve stipitatis; sporidiis late ellipsoideis, monostichis, octonis.

Hab. in nodis ramorum viventium, Fagi cunninghamii, in Tasmania.

Ascomata dense gregaria, 1.5–2 cm cr. sicca flaccida; loculi 3 mm lat., crebri. Hymenium cito evanidum.

As Berkeley's description is somewhat inadequate for comparison with species described later, a revised description has been compiled from a large series of New Zealand specimens and also some Tasmanian specimens collected for me by J. de B. G. Groome. The New Zealand and Tasmanian specimens were found to be identical, even to the extent of the inclusion among the Tasmanian collection of specimens having red coloration round the borders of the opened apothecia, a common feature in New Zealand and thought to be due to disease. The first record from New Zealand appears to be Lloyd, G. G. (1916).

The Latin descriptions for C. nigra and C. pallida have been compiled for me by Mr. C. M. Smith, and his assistance is gratefully acknowledged.

1. Cyttaria gunnii Berk., in Hooker's London Journal of Botany, Vol. 7, p. 576, 1848.

Cyttaria gunnii Berk. Revised description. Plate 9.

Stromata solitary or in dense clusters, up to 2 cm in diameter, pear-shaped, smooth and dry but becoming slimy if wetted, fawn or tan above, light coloured below, bright yellow when mature except for a sterile base which remains white. Hollow at maturity, internal fibres white. Apothecia very numerous, up to 200, bright yellow.

Papillae small, scattered between apothecia, appearing as white flecks on young stroma, arising as minute elevations within shallow depressions, not raised above the surface of the stroma Pycnidia absent Asci 145–165 × 14μ, ascospores sub-globose 12–12.5 × 6.5–12μ, dark coloured, spore print black. Paraphyses as long as asci, 2μ wide, septate, branched, swollen at ends.

An obligate parasite forming globose galls on Nothofagus menziesii (Hook f.) Oerst., in New Zealand and Nothofagus cunninghami (Hook. f.) Oerst., in Tasmania and Victoria. Range in New Zealand as for host.

The sporophores of this species first appear at the time when the buds of the host are beginning to swell; in Southland, in 1946, when most of the investigation was carried out, this occurred during the first week of September. Spore shedding starts early in October and reaches a peak towards the end of November. The first fructifications seen were growing from small twigs showing little or no sign of gall formation; large galls produce fructifications later.

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The galls are small in proportion to the diameter of the branch or stem, as compared with those of C. nigra; they usually occupy about two-thirds of the circumference of the branch and are flattened slightly at the point of origin. When a branch is completely girdled the upper part may die, in which case the death of the gall may follow.

In contrast to C. nigra, galls on the main stem take the form of rope-like encircling bands; for example one sapling of 1½ in d.b.h. had three galls, at 3ft, 4ft 2in. and 5ft 6in from the ground. The lowest was 2in high and 12in in circumference, leaving 1in of bark unaffected, the middle gall was 1½ in high and 10in in circumference, and the highest was 1½ in high and 11in in circumference. The last two galls completely encircled the stem. See Plate 12 C.

The surface of the gall appears smooth, but if the bark is removed the wood is found to be very uneven, being contorted and produced into sharp spines, the tips of which split into fibres which penetrate through the bark. It is from these points that the stromata develop. The scar left on the gall when the fructification falls is a white circular spot, with, in many cases, the remains of the central tube of fibres which pulls out of the stroma.

2. Cyttaria nigra sp. nov. Plate 10 B, C.

Fungus in Nothofago menziesii parasiticus.

Stromata solitaria vel dense aggregata, ad 2 cm dimetro, pyriformia, pusulata, robiginosa, sicca, atra vel interdum aurescentia sed mature adusta. Basis sterilis, adusta, striata, plicata compressa; demum cava; nervis internis flavis saepe contorta. Apothecia permulta usque ad CC, alba. Papillae magnae, exstantes, inter apothecia dispersae, fornicatae, super stromata elatae. Pycnidia (?) in tota superficia densa. Pycnidiosporas non vidi. Asci 132–150μ longi 15μ lati. Ascosporae 9.5–12.5μ longae 8–12μ latae caducae in chartam albam atrae. Paraphyses ascos longitudine aequantes, 2μ lati septati ramulosi extremis tumidulis.

Stromata solitary or in dense clusters, up to 2 cm diam, pear shaped, shagreened with black incrustations, dry, black or black and gold, becoming pallid nigger brown. Sterile base undifferentiated, nigger brown, striate, pleated and compressed. Hollow at maturity, internal fibres yellow to orange, often contorted. Apothecia very numerous, up to 200 or more, pallid white. Papillae large, prominent, scattered between apothecia, dome shaped, elevated above surface of stroma. Pycnidia (?) profuse over entire surface, pycnospores not seen (the identity of bodies termed pycnidia is doubtful). Asci 132–150 × 15μ, ascospores subglobose 9.5–12.5 × 8–12μ dark coloured in mass, spore print black. Paraphyses as long as asci, 2μ wide, septate, freely branched, swollen at ends.

An obligate parasite on Nothofagus menziesii (Hook. f.) Oerst, in New Zealand, causing large globose galls.

Range, recorded from Southland to Lake Taupo in north. Type collection from Woodlaw State Forest, October, 1946, in Forest Research Institute herbarium; specimens at Plant Diseases Division of D.S.I.R. Auckland.

C. nigra comes nearest to C. espinosae Lloyd, but differs from it in colour, internal fibres and shape of apothecia. Both species have a smell resembling apricots. Galls of C. nigra resemble those of C. harioti. Sporophores of C. nigra occur about the same time as those of C. gunnii; the first shedding of spores in 1946 was noted on October 10. The galls are distinguished from those of C. gunnii by their large size in relation to the diameter of the host stem, and their greater height in proportion to diameter. One sapling of 3in d.b.h. had a gall at 8ft from the ground, 30in in circumference and 7in high, completely encircling the stem. This was separated by only 2in from another, 26in in circumference and encircling the stem except for a space of lin.

Relatively larger galls are found on larger trees; galls of 3ft or more in diameter are frequent. Where the upper part of the attacked tree or branch dies, globose galls of a foot or more in diameter are common.

When the bark is removed the gall is seen to differ from C. gunnii in that the spines are very short and less numerous, they arise from bluntly rounded projections on the surface of the gall. The scar left when the fructification falls is orange in colour and may retain the orange cartilaginous tube from the stroma.

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3. Cyttaria pallida

sp. nov. Plate 10 A.

Fungus in Nothofago menziesii parasiticus, ramos caulesque ulceribus fusiformibus deformans. Stromata nonnunquam solitaria saepius duo in ordines coacta vel in unicum ordinem, ad 1.5 cm lata subglobosa, paululum complanata, levia, omnino leucochracea simillimaque argillae, valde viscida, basi curta, occulta, amorpha; matura cava; nervis internis albis basim versus aurantiacis. Apothecia pauca ad L, subaurantiaca. Papillae inter apothecia ordinatim dispositae coniformes supra stromatis curvaturam exstantes. Pycnidia desunt. Asci 150μ–167μ longi 14μ lati. Ascosporae subglobosae 7.5–12.5μ longae, 7.5–9.5μ crassae; acervatae suffuscae; in chartam albam caducae atrae. Paraphyses ascis breviores 100μ longi 2μ lati septati ramulosi.

Stromata solitary, or in single or double rows, up to 1.5 cm diam. sub-globose, somewhat flattened, smooth, uniform dirty white (clay coloured), extremely viscid. Base short, hidden, undifferentiated. Hollow at maturity, internal fibres white, orange at base. Apothecia few, up to 50, dull orange yellow. Papillae uniformly distributed between apothecia, conical, elevated above surface of stroma. Pycnidia absent. Asci 150–167 × 14μ ascospores sub-globose, 7.5–12.5 × 7.5–9.5μ, dark-coloured in mass, spore print black. Paraphyses shorter than asci, 100 × 2μ, septate, branched.

An obligate parasite on Northofagus menziesii (Hook. f.) Oerst., in New Zealand causing large spindle shaped galls. Range, Southland northward to Ohakune. Type collection in Forest Research Institute herbarium, collected at Woodlaw State Forest, October, 1946; specimens at Plant Diseases Division, D.S.I.R., Auckland.

C. pallida appear to be nearest to the South American species C. berteroi Berk., but differs from it in colour, size, hollow mature stroma, internal fibres and length of paraphyses.

The first sporophores were seen in Southland in 1946, somewhat later than those of C. gunnii; spore discharge was noted on October 24.

The galls are of an entirely different type from those of C. gunnii or C. nigra. On twigs they appear as longitudinal rows of pimples erupting through the bark, and from these the sporophores arise singly. These galls spread and cause twisting of the branches, which become swollen and brittle; many branches die, others suffer from breakage due to wind. In large trees the crown becomes thin and the branches distorted.

The most striking deformity results from the fungus attacking a small shoot on the trunk of a sapling; in this case the fungus spreads down the twig and attacks the trunk. At first a ridge is formed beneath the bark, as growth proceeds the ridge divides into two and the final result is a paddle-shaped gall. As an example, a sapling of 1½ in d b.h. had a gall extending 1ft above and 3ft below the source of infection, which was a small twig on the trunk. The canker was 3in wide and the stem three-quarters encircled. The same stem had several such cankers of larger or smaller size and the upper part had been killed by the attack. See Plate 12 B.

In these paddle-shaped galls the central area may be dead, covered by pimples, or may appear to be normal bark. Most of the sporophores occupy the margins of the galls.

4. Cyttaria septentrionalis Herbert. Proc. Roy. Soc. Queensland, Vol. 41, No. 13, p. 158–161, 1930.

Herbert (1930) described C. septentrionalis as follows:—

“Stromata gregaria, globosa, 5–7 cm diam, demum cava, superficie ubique locellata, flava; locellis 3 mm latis; ascis cylindraceis 170–210μ longis, 17μ latis, octosporis, non stipitatis; sporidiis globosis. 15μ diam, paraphysibus filiformibus; species edulis. Hab in summo monte Hobwee, ad ramos vivos Nothofagi moorei, September, 1929.”

To this description it may be added that the young stromata are shagreened with a black pitchy substance (pycnidia?). Simple pores open to the exterior between the apothecia and raised papillae are absent. The internal fibres are well developed; they are plate-like and irregularly anastomosing. The galls are spindle-shaped in the specimens kindly collected for me on Mt. Hobwee by J. de B. G. Groome. Janet M. Wilson (1937) states that the galls are long and narrow or short and round, so that two species on N. moorei may be involved.

Herbert (1930) does not describe the galls but conveys the impression that they are globose; his material was collected from the MacPherson Ranges. Material used

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by Janet M. Wilson was collected from the Mt. Royal Ranges; her illustrations show either galls of two species or of one species intermediate between the spherical and longitudinal types. The above notes refer to immature stromata on small, spindle-shaped galls. Whether these specimens belong to Herbert's species or to a new species cannot be decided on the material available.

References

Berkeley, M. J., 1841. On an edible Fungus from Tierra del Fuego, and an allied Chilian species. Trans. Linn. Soc. 19: 37–42.

—— 1846. Flora Antarctica. 1: 452, 1844–47.

—— 1848. Hooker's London Journal of Botany. Vol. 7: 576.

Espinosa, M. R., 1926. Los hongos chilenos del genero Cyttaria. Revist. Chil. Hist. Nat. 30: 206–256.

—— 1930. Cyttaria harioti en Colchagua y clave de las especies chilenas del genero Ibid. 34: 140–142.

—— 1940. Un hongo nuevo del genero Cyttaria Berk. Bol. Mus. Noc. Chile. 18: 23–26.

Fischer, E. D., 1888. Zur Kenntnis der Pilzgattung Cyttaria. Bot. Zeit., Bol. 46. Leipzig.

Herbert, D. A., 1930. Cyttaria septentrionalis, a new fungus attacking Nothofagus moorei in Queensland and New South Wales. Proc. Roy. Soc. Queensland. 41: 158–161.

Lloyd, C. G., 1916. Cyttaria gunnii. Myc. Not. 42, Lloyd's Myc. Writ.

—— 1917. The genus Cyttaria. Myc. Not. 48. Ibid.

—— 1918. Cyttaria gunnii. Myc. Not. 53. Ibid.

Marchionatto, J. B., 1940. Las especies de Cyttaria y Cyttariella en la Argentina. Reve. Inst. Bot. Darwinion de Buenos Aires.

Massee, G., 1910. Diseases of Cultivated Plants and Trees.

Palm, B. T., 1932. On Cyttaria Berk. and Cyttariella n.gen. Annales Mycol. Bol. 30. Berlin.

Santesson, R., 1945. Cyttaria, a genus of Inoperculate Discomycetes. Svensk Bot. Tidscrift, Bd. 39: 4. Uppsala.

Spegazzini, C., 1887a. Fungi Patagonici. Bol. Acad. Mac. Ciene. (a) Cordoba, Vol. 11, p. 286. Buenos Aires.

—— 1887b. Fungi Fuegiani. Ibid. p. 135.

—— 1922. Criptogamae nonullas fuegianae. Anal. Soc. Cient. Argent., Vol. 94, Buenos Aires.

—— 1923. Algumos hongos de Tierra del Fuego. Physic. 7: 7–23. No. 10.

Wilson, Janet M., 1937. Structure of Galls formed by Cyttaria septentrionalis on Nothofagus moorei. Proc. Linn. Soc. N.S.W. 62: 1–8.

G. B. Rawlings

,
Forest Research Institute,
P.B., Whaka.