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Johnston, P.R. 1990: Hypohelion gen. nov. (Rhytismataceae). Mycotaxon 39: 219-227.

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Johnston, P.R. 1990: Hypohelion gen. nov. (Rhytismataceae). Mycotaxon 39: 219-227.
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An ascomycete fungus belonging to the family Rhytismataceae. Ascomata subcuticular, upper wall uniform in internal structure. In opened ascomata upper wall uniform in thickness, and without differentiated cells lining the ascomatal opening. Ascomata lacking a darkened lower wall. Paraphyses swollen, clavate.
Fungus ascophorus ad familiam Rhytismatacearum pertinens. Ascomata subcuticularia, paries superior structure interna indutus uniformi. Ascomatibus ruptis paries superior crassitudinis respectu uniformis, sine celluis dissimilibus in superficie exposita. Ascomata sin parie inferiors fusco. Paraphyses tumidae, clavatae.

Etymology: Hypo-; refers to Hypoderma, the genus from which H. scirpinum has been segregated. -helion = swamp or marsh dwelling; refers to the habitat of the species included in this genus.

Johnston (1988a, 1988b, 1989, 1990) discussed the unsatisfactory nature of currently accepted generic limits in the Rhytismataceae, and showed that, at least for nonconiferous species, an alternative is possible. Rather than using single characters chosen a priori as important, genera may be based on sets of several characters, which in this case are mostly associated with ascomatal structure and development.

The characters used to define Hypohelion and to distinguish it from other genera in the Rhytismataceae are:
1. Subcuticular ascomata.
2. The upper wall of unopened ascomata is homogeneous in structure, comprising dark brown, thick-walled, angular cells.
3. In opened ascomata the upper wall is more or less uniform in thickness.
4. The edge of the ascomatal opening is not lined with differentiated cells.
5. The ascomata lack a darkened lower wall.
6. The paraphyses are swollen at the apex.

Hypoderma, in the sense accepted by Johnston (1990), possesses only the first of the characters listed above, and the upper wall of the ascomata has thickened labia and a characteristic internal differentiation. Powell (1974) considered that vestiges of thickened labia, in the form of a medial keel, could be seen in the upper wall of the ascomata in many collections of H. scirpinum. My observations showed that although a keel-like ridge is often seen on unopened ascomata, this is due to a folding, rather than a thickening, of the ascomatal wall. It cannot be compared with the thickened labia characteristic of Hypoderma rubi and other species of Hypoderma.

Hypohelion scirpinum and H. parvum are morphologically and anatomically similar, and are also similar with respect to host preference. Both are found on Cyperaceae growing in swampy habitats. The same kind of relationship between ascomatal structure and host substrate was demonstrated for a group of Lophodermium, Lophomerum, and Meloderma spp. found on the closely related families Epacridaceae, Ericaceae, and Pyrolaceae (Johnston 1988b). This correlation between morphological and biological characters suggests that the character set used in this paper, and by Johnston (1990) for Hypoderma, may define rhytismataceous genera in a way which reflects natural relationships amongst these fungi.

Species typica: Hypohelion scirpinum (de Candolle:Fries) Johnston.
PDD 48879 - holotype.
Ascomata developing on dead leaves, not associated with paler areas on leaf, but sometimes with individual ascomata surrounded by narrow, black zone lines. Not associated with conidiomata. In surface view ascomata 0.4-0.7 x 0.2-0.3 mm, more or less elliptic or irregular in outline. Unopened ascomata with uniformly black walls. Ascomata opening by an irregular split either elongate or radiate, depending on ascomatal shape. No differentiated zone visible along edge of opening split.

Ascomata subcuticular. In vertical section upper wall of unopened ascomata up to 20 µm thick, comprising brown to dark brown, thick-walled, angular cells. In opened ascomata upper wall 15-20 µm thick, becoming gradually narrower toward outer edge, comprising dark brown, thick-walled, angular cells. No differentiated layer associated with edge of the ascomatal opening. Darkened lower wall absent.

Paraphyses 1.5-2 µm diam., swelling up to 5 µm diam. at the clavate apex, extending 5-10 µm beyond asci. Asci 65-90 x 9-10 µm, clavate, tapering gradually to rounded apex, wall undifferentiated at apex, 8 spored, spores extending almost to base, maturation sequential. Ascospores 23-32 x 3-3.5 µm apex rounded, tapering to more or less acute base, 1 septate, surrounded by narrow gelatinous sheath.

New Zealand: Westland.
Dead leaves of Carex.
Ab H. scirpinum ascocarpi 0.4-0.7 x 0.2-0.3 mm, ascis 65-90 x 9-10 µm, ascosporis 23-32 x 3-3.5 µm differens.
Etymology: parvum = small; refers to small size of ascomata, asci and ascospores compared to H. scirpinum, the type species of the genus.
New Zealand, Westland, Fox Glacier, Lake Gault on Carex sp., coll. P.R. Johnston (R174), 10.iv.1983 (PDD 48879).
CANADA: Ontario, on Scirpus occidentalis, coll. H. S. Jackson et al., 4.x.1933 (NY ex TRTC 4760). GERMANY: coll. Winter, v.1874, ex Herb. Thumen (NY); Bavaria, on Scirpus lacustris, coll. A. Vill, iv.1909, Sydow, Myc. Germ. No. 984 (PDD); Berlin, on Scirpus lacustris, coll. P. Sydow, ix.1886 (NY); on Scirpus lacustris, ex Herb. Barbey-Boissier No. 975 (NY); on Scirpus lacustris, coll. G. Winter, vi.1874, de Thumen Myc. Universalis No. 178 (NY); on Scirpus lacustris, coll. J. Kunze, iv.1879, Kunze, Fungi selecti exsiccati No. 277 (NY); on Scirpus lacustris, coll. Krieger, xi. 1911, Rehm Ascomycetes No. 1953 (NY). SWEDEN. Fries, Fungi Sueci No. 170 (K). USA: Washington, Seattle, on Scirpus lacustris var. occidentalis, coll. A. M. Parker, x.1892, Ellis & Everhart, North American Fungi No. 3038 (PDD, NY); Dakota, on Scirpus sp., coll. J. F. Brenkle, 9.x.1910, Fungi Dakotenses No.142 (NY); North Dakota, on Scirpus lacustris, coll. J. F. Brenkle, v.1908 (NY); California, Lake County, Bogg's Lake, on Scirpus acutus, coll. L. Bonar, 24.viii.1945, Californian Fungi No. 644 (NY); Washington, Seattle, on Scirpus lacustris var. occidentalis, coll. C. V. Piper, x.1892, Ellis & Everhart, Fungi Columbiani No. 328 (NY); Massachussets, Essex, on Scirpus lacustris, viii.1890 (NY).
Ascomata developing in slightly paler areas of the leaves, sometimes surrounded by narrow, black zone lines. Associated with anamorph conidiomata.

In surface view ascomata 0.8-3(-7) x 0.5-0.8 mm, elliptic to broad-elliptic in outline, with rounded ends. In unopened ascomata the wall is uniformly black, usually developing a fold-like ridge along the long axis of the ascomata. Ascomata opening by an elongate split which usually forms more or less along the fold-like ridge. No differentiated zone visible along the edge of the opening split. Parts of the upper wall adjacent to the opening split often break off and fall away, leaving the hymenium partially exposed.

Ascomata develop between the host cuticle and epidermis, although when ascomata develop over stomata in the host leaf, hyphae invade and break down a few host cells immediately below the stomata. In vertical section, at an early stage of development, the ascomatal initial comprises 2-3 layers of hyaline plectenchyma, covered with a single layer of brown, thick-walled, angular cells, which comprises the first stage in the development of the upper wall (Fig. 2A). As the ascomata develop the upper wall thickens to 20-30 µm, comprising thick-walled, brown, angular cells, and a cavity develops within the hyaline plectenchyma, with paraphyses growing upward from the base of this cavity (Fig. 2B). From this stage the upper wall does not become thicker, but it darkens and becomes distorted and folded by the pressure exerted from the developing hymenium. The upper wall shows no internal differentiation, comprising uniformly thickened and darkened, angular cells, but shows a distinct upward fold near the centre of the ascomata (Fig. 2C). The upper wall splits open in the region of this fold (Fig. 2D). No layer of differentiated cells is present along the edge of the opening split. A darkened lower wall does not develop, the hyaline tissue below the hymenium resting directly on the more or less intact epidermal cells of the host.

Paraphyses 1.5 µm diam., swelling more or less abruptly to 4-6 µm diam. at the clavate to knob-like apex. Asci 130-160 x 15-18 µm, clavate, tapering gradually to rounded apex, wall at apex sometimes slightly thickened with indistinct central pore, 8 spored, spores extending two-thirds down the ascus, maturation sequential. Ascospores 40-75 x 4.5-6.5 µm, tapering slightly to both ends, 0-1 septate, surrounded by narrow gelatinous sheath.

Conidiomata 0.2-0.3 mm diam., round in outline, dark brown to black, pustulate, subcuticular. In vertical section lenticular in shape, upper wall 3-5 µm thick, comprising dark brown material without obvious cellular structure. Darkened lower wall absent, the conidiogenous layer developing on 2-3 rows of thin-walled, hyahne, angular cells. Conidiogenous cells 9-15 x 2-3 µm, cylindric, solitary, forming a palisade-like layer, with sympodial proliferation, often with more than one developing conidium held at the apex. Conidia 3-4.5 x 1 µm, oblong-elliptic with rounded ends, or short-clavate with a broadly rounded apex and tapering to a truncate base, hyahne, 0 septate.

Northern Europe and northern North America (Powell 1974).
Dead leaves of Scirpus.

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18 March 2001
24 February 2005
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