The Epiphyllous Lichens of Kitchener Park, Feilding, New Zealand.
[Read before the Philosophical Institute of Canterbury, 8th June, 1927; received by Editor, 20th April, 1928; issued separately, 10th August, 1928.]
A. Ecological (H. H. Allan.)
Hochstetter, in his classical sketch of New Zealand vegetation (1863, p. 417), was the first to emphasize the tropical or subtropical character of New Zealand forests. Made familiar by citation in Schimper's Pflanzen-Geographie, this view has gained wide currency, and subsequent research has only served to confirm it. The most recent treatment is that of Cockayne (1926, p. 7), who divides the New Zealand forests into the two great classes “subtropical rainforest” and subantarctic rain-forest.” In enumerating the special characteristics of these rain-forests he concludes with, “(13) small bryophytes and lichens frequently occur as epiphytes on leaves of trees.” This last feature has not hitherto received the attention its interest and importance merit. The only work so far done appears to be the well-known paper of Jennings (1896, p. 753), in which are described two species of Phycopeltis—P. nigra and P. expansa—and the association of the latter with a fungus to produce the lichen Strigula complanata is shown.
My own observations would lead me to suggest a modification of Cockayne's statement. I find epiphyllous bryophytes and algae occasional on the leaves of the shrub-layer, but rare on forest-trees. On shrubs in very humid forest the epiphyllous bryophytes may become frequent. I find epiphyllous lichens to be comparatively rare, but to occur on leaves of plants of all layers. These findings agree with those of Dr. G. E. Du Rietz, who recently made a lichenological survey of New Zealand.
The small remnant of forest, however, near Feilding, known as Kitchener Park and fortunately constituted a “reserve,” has epiphyllous lichens in abundance (algae and bryophytes are not wanting as epiphylls, but are of much less moment), and provides a convenient area for their study. Opportunity was therefore taken to secure the services of the doyen of lichenology, Hofrat Dr. A. Zahlbruckner, Director of the Department of Botany in the Natural History Museum of Vienna. The results of his work are given in section B. and provide a sure basis upon which ecological work can be built. In section C his colleague, Professor Dr. Karl Keissler, describes and discusses a new lichen-parasite of considerable interest that was discovered on certain specimens.
An attempt is made in this section to give a preliminary account of the ecological problems presented by the phenomena observed at
Kitchener Park. It will be realized, however, that at the present stage of our knowledge, only very general and introductory work can be expected. Nothing, e.g., of moment is known concerning the life-history of the leaves of any of our “evergreen” trees. This preliminary account, however, may have its value in drawing attention to a group of problems that apart from their intrinsic interest, must have an important bearing on general forest ecology.
2. The Problems Envisaged.
Ward (1884, p. 115) observed, “The whole group of ‘Epiphyllous Lichens’ would no doubt well repay prolonged and careful investigations.” The work so far accomplished on the ecology of epiphyllae has brought to light many of the problems concerned, but has not made much advance towards solving them. Ward's own contribution is (loc. cit., p. 87), “The organism to be described [Strigula complanata] occurs on the upper surface of the leaves of so many plants widely separated in affinity and origin, that one must regard the species of the supporting plant as an accident. One general feature, however, is common to all these leaves: they are invariably hard and persistent, differing moreover, in degree in this respect.” Schimper (1903, p. 328) merely remarks that epiphyllous forms of cryptogams are “apparently confined to the tropics,” and are “quite common features, particularly on ageing leaves, in very humid rain-forests.” The ground-breaking paper of Fitting (1910, p. 505) still remains the most important and suggestive work yet published. It is the only paper cited by Warming and Graebner (1918, p. 288) and Fünf-stück (1926, p. 11) in their brief treatments of the subject. Curiously it is not cited in the ecological chapter of Smith (1921, p. 356 et seq.). That author remarks (loc. cit., p. 363) concerning epiphyllous: lichens, “These grow on Ferns or on the coriaceous leaves of evergreens in the tropics.… Observations are lacking as to the associations or societies of these lichens whether they grow singly or in companies.”
Fitting (loc. Cit., p. 506) classes the crustaceous lichens of the upper surface of leaves in the Javanese forests studied by him, into three groups: (1) a small group the members of which penetrate more or less deeply into the leaf-tissues-causing local injury or death of the parts; (2) a large group the members of which cause the cuticle to peel off, and establish themselves on the outer walls of the epidermis; (3) a small group the members of which grow on the cuticle without injury to it. In this group may be included certain foliaceous lichens “in Gegenden mit sehr feuchter Atmosphaere.”
The influence on the leaf-tissues he considers comparatively small, the leaf reacting by way of the thickening of the outer walls of the epidermis or even of the outermost layer of palisade cells, and by the development of wound-cork. The damage done must be attributed to the alga as well as the fungus, and he concludes that the first, and to a less extent the second, group must be considered as parasitic, not merely epiphytic. He cites Busse's opinion, with modified approval, that epiphyllous lichens are confined to evergreen leaves, mainly those that are smooth and leathery. Very hairy leaves are free, but drip-tips and other adaptations favouring quick removal
of water from the leaf-surface have no effect in checking lichen growth on the surface. He thinks it not a general rule, as Busse considers, that leaves unprotected from heavy downpours and exposed to sun are only lichenized if a damp atmosphere persists during the dry period. Busse's statement that shading favours settlement is considered to be true only of group 3, the others requiring a certain intensity of light.
Fitting concludes by mentioning certain unsolved problems. It is very doubtful whether the qualitative and quantitative differences in the lichen-flora on leaves of different plants is solely to be attributed to different degrees of air-humidity, light-intensity, protection against the impact of rain-drops, wettability of leaves and so on. On what rests the different distribution of lichens on the different leaves of a plant? What determines the often very different lichen-floras of different species? Why do leaf-lichens avoid certain species?
Shreve (1914, p. 86) says “Jungner considers the ready drainage of water from the surface of leaves as of the greatest importance in cleansing them of the spores of epiphyllous hepatics, mosses, etc., and in preventing the presence on the surface of the leaf of a film of water which would be favourable to the growth of epiphyllae. In the Kamerun he has found the leaves with dripping points to be free of epiphyllae unless injured, and the leaves with blunt apices to be full of epiphyllae soon after reaching mature size.” After dealing (p. 87) with the epiphyllae of Jamaican rain-forest, Shreve concludes: “Wherever the humidity is high and constant, fog prevalent, and condensation of moisture common, those plants which are sheltered from breeze or subjected to drip from higher layers of foliage bear epiphyllae regardless of whether or not they possess dripping points. Plants of open situations, on the other hand, are devoid of epiphyllae regardless of their form.” It is to be noted that the epiphyllae in question were mainly Hepaticae of the genus Lejeunea. Shreve (p. 91) mentions that Stahl also rejects Jungner's view.
I am studying the questions raised by these citations, and certain preliminary observations are here offered.
3. Composition and Structure of the Kitchener Park Forest.
A brief account of the remnants, of which Kitchener Park is one, of the great rain-forest that once covered the drainage-area of the Manawatu River has already been published (Allan, 1923, p. 402). It will be sufficient here to indicate the main features of the Kitchener Park forest, in its present much-modified condition.
The dominant species are the tall trees Podocarpus spicatus and P. dacrydioides. In addition, there is in the upper layer much P. totara and Beilschmiedia tawa, the latter apparently in process of replacing the podocarps. Second-layer trees form a general roof-cover through which project the dominants, these latter having been to some extent felled, thus allowing of a more rapid development of Beilschmiedia tawa. The chief trees of this layer are Alectryon excelsum and Beilschmiedia tawa, with some Elaeocarpus dentatus, Olea Cunninghamii, and Plagianthus betulinus. Along shallow mud-floored water-channels, often dry for considerable periods, Cordyline australis in very large specimens is a true forest member.
There is an irregular third layer, consisting largely of Hoheria angustifolia, H. sexstylosa (these two mainly marginal, and there hybridizing), Melicytus ramiflorus, Myrtus bullata, Nothopanax arboreum, Paratrophis microphylla, Pennantia corymbosa, Pittosporum eugenioides, P. tenuifolium, Pseudopanax crassifolium. var. unifoliolatum, and Suttonia australis.
The chief members of the shrub-layer are Coprosma areolata, C. ratundifolia (and their hybrids), Melicope simplex, Melicytus micranthus, Myrtus obcordata (and M. bullata X obcordata in numerous forms), and Nothopanax anomalum. There are now very few tree-ferns, but Cyathea dealbata, Dicksonia fibrosa, and D. squarrosa were once common.
The floor-plants of moment are: Asplenium bulbiferum, Athyrium umbrosum, Carex dissita, Dryopteris decomposita, D. pennigera, Microlaena avenacea, Pellaea rotundifolia, Polystichum Richardi, and Uncinia uncinata.
The more common epiphytes are: Astelia solandri, Asplenium adiantoides (under the Astelia tussocks), Bulbophyllum tuber-culatum, Cyclophorus serpens, Dendrobium Cunninghamii, Earina mucronata, Pittosporum cornifolium.
Lianes are numerous, including Clematis foetida, C. hexasepala (and hybrids), Fuchsia perscandens, Metrosideros Colensoi, Parsonia capsularis, P. hetrophylla (and hybrids), and Rhipogonum scandens. On the forest-margins Muehlenbeckia australis, M. complexa, along with hybrid forms, are common.
The forest thus shows a general subtropical aspect, though the modifications due to partial milling, entry of stock, and other interference by man, have greatly lessened the density and changed the relative abundance of many species, especially in the undergrowth. The total florula is made up of 38 pteridophytes, 3 gymnosperms, 20 monocotyledons, and 79 dicotyledons. There are 16 groups of hybrids. The growth-forms comprise 29 trees, many also occurring as shrubs, 45 shrubs, 5 semi-woody plants, and 61 herbs. There are 11 epiphytes, 4 hemi-parasites, and 17 lianes.
4. The Distribution of Epiphyllous Lichens in Kitchener Park.
Epiphyllous lichens have been observed on the following species: Hymenophyllum demissum, H. scabrum, Dicksonia fibrosa, Cyathea dealbata, Polystichum vestitum, P. Richardi, Dryopteris decomposita, D. pennigera, Asplenium adiantoides, A. lucidum, A. bulbiferum, A. flaccidum, Blechnum procerum, Pellaea rotundifolia, Histiopteris incisa, Pteridium esculentum, Polypodium pustulatum, P. diversifolium, Cyclophorus serpens, Podocarpus totara, P. spicatus, P. dacrydioides, Astelia nervosa var. sylvestris, A. Solandri, Earina mucronata, Dendrobium. Cunninghamii, Loranthus micranthus, Beilschmeidia tawa, Melicope simplex (uncommon), Alectryon excelsum, Melicytus micranthus, Metrosideros Cobensoi, M. perforata, M. hypericifolia, Myrtus bullata, Nothopanax anomalum, Pseudopanax crassifolium, var. unifoliolatum, Olea Cunninghamii. A number of species, e.g., Rhipogonsum scandens, Myrtus obcordata, and various ferns, have been observed with Phycopeltis tropica, but with-
out lichens. Thus, of the 140 species found in the forest 38 are found to bear lichens. The following species usually bear them in abundance: Podocarpus totara, P. spicatus, P. dacrydioides, Earina mucronata, Beilschmiedia tawa, Alectryon excelsum, Metrosideros Colensoi, Myrtus bullata, and to these especial attention was directed, though many of the others, were often well covered. Theloschistes flavicans occasionally occurs on the leaves of Alectryon excelsum, Podocarpus spicatus and other species, but is not a typical epiphyllous lichen, and is not here further dealt with.
The following data concerning the species attacked are of interest—
(1) Habitat:—14 plants of the forest-floor are attacked, 21 of the shrub-layer, 15 of the tree-layer.
(2) Growth-form:—There are 17 ferns (5 small, 6 medium, 6 large); 2 tree-ferns; 14 shrubs; 2 semi-woody plants; 3 lianes; 7 trees (which may also be attacked in the shrub-stage).
(3) Leaf-size:—22 bear small leaves or leaflets, 11 medium, 5 moderately large.
(4) Leaf-texture:—2 are very thin, 13 thin, 18 rather thick, 5 thick.
(5) Upper leaf-surface:—26 are smooth, of which 5 are more or less glossy; 12 are very slightly rough. All are glabrous, or practically so, when mature.
(6) Leaf-apex:—None possess distinct “drip-tips.” In plants not affected only 1 can be said to possess a true “drip-tip.”
(7) Frequency of attack:—5 are rather rarely attacked, 14 occasionally, 9 frequently, 10 commonly. The most common species of lichen is Lopadium subcaerulescens, attacking 31 species. L. Allanii may be more abundant than is at present known. Not only are many species attacked, but many leaves may be almost completely covered by the lichen, especially in Beilschmiedia tawa and Alectryon excelsum. Strigula africana appears to be confined to B. tawa and A. excelsum, but on both may be very plentiful. Bacidia spp. appear to be confined to the podocarps, and Phylloporina spp. to the Myrtaceae.
The following tentative conclusions seem worthy of record as providing working hypotheses and a basis for further record:—
(1) The habitatal requirements of epiphyllous lichens must be considered in two groups— (a) the conditions offered by the leaves themselves, including their life-history, physical, anatomical and constitutional features: (b) the outer environmental factors, especially intensity of light, wind and humidity.
(2) The species of supporting plant is not purely accidental, though certain lichens have a wider range of hosts than others. This may not apply to Fitting's third group. The incidence of the species Physcia at Kitchener Park appears to depend on a moderate degree of humidity, moderate shade, and relatively persistent leaves on the host plant. But species otherwise favourable to lichen development may be free owing to an unfavourable outer environment, e.g., Myrtus bullata is free if the insolation is too strong. Certain genera with apparently suitable leaves do not appear to develop lichens, e.g., Hoheria, Griselinia, Pittosporum, Coprosma, Rhipogonum. The reason may lie in their constitutional characters.
(3) Leaves of relatively short-life may be attacked, but for full development of epiphylls a certain persistence is required. Leaves of Alectryon excelsum and Beilschmiedia tawa often show light attack by Strigula africana towards the end of the first growing season, this being as true of seedlings as of adult plants. The leaves are more and more covered till at leaf-fall, which does not appear to be hastened, the surface may be completely covered. The lichen may fructificate on dead, fallen leaves where the ground is dry. A considerable degree of humidity favours the early establishment of lichens. This is aparently especially important in filmy ferns, lichen-attacked leaves being first more or less covered with bryophytes.
(4) The lichens appear to inflict little real damage, as seedlings survive and older trees appear to be as vigorous as those free from attack.
(5) Great wettability of surface does not appear to be required, nor is a ready run-off of water of any moment.
(6) A certain optimum of light-intensity is required for full lichen-development. Leaf-lichens are very rare in the deepest shade, and in the fullest sun, with marked development in moderate shade. This is seen well in plants of the forest floor, and in leaves of the shrub-layer. As the tree is ascended the lichen development decreases to the vanishing point. Juvenile Nothapanax crassifolium is generally strongly attacked, while the adult is rarely so. There is in general a distinct belt of major attack.
(7) Considerable humidity favours attack. Margins of pools in the forest-interior show a more or less marked lichen girdle. Shade and humidity tend to be associated, but humidity cannot prevail over the effect of deep shade.
(8) Strong wind is apparently prejudicial to lichen attack, but its effect is not always easy to disentangle from that of insolation. As a result of both factors, marginal plants are less attacked than those more protected.
(9) The epiphyllous lichens form more or less distinct communities, this being aided apparently by the different environmental requirements of the lichens. Humidity favours Lopadium, and shade with less humidity Strigula. Leaves subject to much drip from above will be covered with Lopadium. Bacidia appears on juvenile leaves of podocarps in the shrub-layer, associated with Lopadium. L. sub-caerulescens is the most frequent lichen on Beilschmiedia tawa, and may form pure communities. Strigula africana may also form pure communities on B. tawa, but where the two lichens occur together S. africana is dominated and suppressed. On Alectryon excelsum Strigula africana is much the more common lichen, but here too it may be suppressed by L. subcaerulescens.
Allan, H. H., 1923. The Forest Remnants in the Neighbourhood of Feilding. Rep. Aust. Ass. Adv. Sci., vol. 16, p. 402 ff.
Cockayne, L., 1926. Monograph on the New Zealand Beech Forests. Part 1., Wellington.
Fitting, H., 1910. Über de Beziehung zwischen den epiphyllen Flechten und den von Ihnen bewohnten Blättern. Ann. du Jard. bot. De Buit., 3rd supp., 2nd pt., p. 505 ff.
Funfstück, M., 1926. Lichenes; Die Natürlichen Pflanzenfamilien. 8. Band, p. 11. Leipzig.
Hochstetter, F., von., 1863. New-Seeland. Stuttgart.
Jennings, A. V., 1896. On two new species of Phycopeltis from New Zealand. Proc. Roy. Irish Acad., 3rd ser., vol. 3, No. 5, p. 763 ff.
Schimper, A. F. W., 1903. Plant-Geography upon a Physiological Basis (Trans. W. R. Fisher). Oxford.
Shreve, F., 1914. The Direct Effects of Rainfall on Hygrophilous Vegetation. Journ. Ecol., vol. 2, No. 2, p. 82 ff.
Smith, A. L., 1921. Lichens. Cambridge.
Ward, H. M., 1884. Structure, Development and Life-History of a Tropical Epiphyllous Lichen (Strigula camplanata Fée.). Trans. Linn. Soc., vol. 2, pt. 6, 2nd ser., Bot., p. 87 ff.
Warming, E., and Graebner, P., 1918. Eug. Warming's Lehrbuch der okologischen Pflanzengeographic. Berlin.
B. Taxonomic (A. Zahlbruckner).
Phylloporina (sect. Segestrinula) rufula (Krmphbr.) Müll. Arg., Lich.. Epiphyll. Novi, 1890, p. 21, et in Flora, vol. 73, 1890, p. 196; A. Zahlbr., Catal. Lich., vol. 1, p. 532.—Verrucaria rufula Krmphbr. in Nouv. Giorn. Bot. Ital., vol. 7, 1875, p. 53.—Porina rufula Wain., Étud,. Lich. Brésil, vol. 2, 1890, p. 227.
On the leaves of Myrtus bullata.
2. Phylloporina (sect. Sagediastrum) cerina A. Zahblbr. nov. spec.
Thallus epiphloeodes, vix visibilis, maculas minores formans, valde tennis, substrato quasi suffusus et quoad colorem ab eo vix diversus, virens, opacus, continuus, in margine non bene limitatus, sorediis et isidiis nullis; gonidia phycopeltoidea. Apothecia sessilia, minuta, 0.2-0.5 mm., lata, dispersa, approximata vel confluentia, convexa, ad basin non constricta, ceracea-lutea, nitidula, poro haud conspicuo; excipulum dimidiatum, tenue, pallens (lutescens), a thallo tenuiter obductum, poro terminali, rotundo, 18-20 mmm. lato; nucleus decolor, purus, J. lutescens, imprimis asci; paraphyses increbrae, capillari-filiformes, simplices, eseptatae, ad apicem non latiores; asci oblongo-fusiformes, angusti, subrecti vel curvuli, ad apicem rotundati et membrana parum incrassata cincti, 8 spori; sporae in ascis biseriales, decolores, fusiformi-oblongae, utrinque rotundatae, rectae vel curvulae, 4 loculares, septis valde tenuibus, membrana tenui cinctae, 16-20 mmm. longae et 4-5 mmm. latae.
On the leaves of Metrosideros Colensoi.
3. Strignla africana Wain, in Catal. Welwitsch Afric. Plants, vol. 2, 1890, p. 461.
Frequent on the leaves of Beilschmiedia tawa, Alectryon excelsum.
4. Arthothelium vermiferum A. Zahlbr. nov spec.
Thallus epiphyllus, tenuissimus, substrato adhaerens, vulgo maculas parvas, ad 4 mm. latas, monocarpicas formans, apothecia annulatim cingens, demum rare approximatas, haud confluentes, virenti-albidus, opacus, KHO—, Ca Cl2O2—, continuus, laevigatus, sorediis et isidiis non praeditus, in margine obscurius non cinctus, sed bene limitatus; homoeomericus, ex hyphis tenuissimis, intricatis
et gonidiis ad Trentepohliam pertinentibus formatus, cellulis gonidiorum luteo-viridibus vel subaurantiacis, concatenatis et glomeratis, plus minusve oblongatis, 12-24 mmm. Iongis. Apothecia exigua, 0.1-0.18 mm. lata, obscure fusca vel nigricantia, opaca, rotunda vel rotundata, planiuscula, adpressa, emarginata; excipulum distinctum non evolutum; hymenium superne pallide cinnamomeo—vel umbrino—fuscescens, caeterum decolor et purum, J. vix lutescens; hypothecium angustum, decolor; paraphyses mox gelatinose confluentes, non distinctae; asci late ovales vel ellipsoideo-ovales vel subpanduriformes, superne latae rotundati et membrana bene incrassata cincti, 8 spori; sporae in ascis 4-6 seriales, plus minusve convolutae, vermiculari-subcylmdricae, utrinque rotundatae, arcuatae et demum semicirculares, murales, cellulis subcubicis, leptodermaticis, in seriebus superpositis ad 20, in seriebus horizontalibus vulgo 2, membrana tenui cinctae, J. parum lutescentes, 60-80 mmm. longae et ad 8 mmm. latae.
A very distinct species. On the leaves of Metrosideros Colensoi.
5. Bacidia (sect. Weitenwebera) perparva A. Zahlbr. nov. spec.
Thallus epiphyllus, crustaceus, uniformis, tenuis, submembranaceus, demum facile desquamescens, sordide cinerascena, opacus, KHO—, Ca Cl2O2—, continuus subleprosus vel subinaequalis, sorediis et isidiis nullis, in margine linea obscuriore non cinctus, fere homoeomericus, gonidiis cystococcoideis, globosis, laete viridibus, 9-10 mmm. latis (gonidiis aliis non ad lichenem pertinentibus hic inde intermixtis). Apothecia minima, ad 0.1 mm. lata, biatorina, fusco-nigra, opaca, rotunda, ad basin non constricta, e planiuscula modice convexa, dispersa vel approximata; excipulum dimidiatum, crassiusculum, aeruginoso-fusocscens, ex hyphis intricatis formatum, cum hypothecio aeruginoso-fuscescente (obseuriore) confluens; hymenium superne anguste umbrino-fuscum, non pulverulentum, caeterum decolor et purum, angustum, 30-35 mmm. altum, J. lutescens; paraphyses parum distinctae et increbrae, filiformes, simplices, eseptatae, ad apicem clavatae et obscuratae; asci crebri, ovali-clavati, superne bene rotundati et membrana modice incrassata cincti, 8 spori; sporae in ascis biseriales, decolores, oblongae, rectae, utrinque rotundatae, 4 loculares, septis valde tenuibus, membrana tenui cinctae, 11-12 mmm. longae et 3-5 mmm. latae.
On the leaves of Podocarpus spicatus, P. dacrydioides, P. totara.
6. Lopadium Allanii A. Zahlbr. nov. spec.
Thallus epiphyllus, sat late expansus, parum visibilis, quasi suffusus, maculas rotundatas et plus minusve confluentes formans, lutescenti-virens, opacus, KHO—, Ca Cl2O2—, continuus, submembranaceus, in margine linea obscuriore non cinctus, sorediis et isidiis destitutus, trichomatibus nigris superne non ornatus; fere homoeo mericus, ex hyphis intricatis, decoloribus, ad 2 mmm. crassis, leptodermaticis et ex gonidiis pleurococcoideis, globosis, lutescenti-viridibus, 6-8 mmm. latis formatus, hyphae thalli in ambitu thalli dissolutae, plus minusve radiantes, increbre ramosae et passim subreticulatae, non amyloideae. Apothecia biatorina, dispersa,
rotunda, parva, ad 0.3 mm. lata, sessilia, ad basin, levissime constricta, demum convexula, ceraceo-lutescentia, parum nitidula; discus angustus, thallo concolor, impressus; margo discum paulum superans, angustum persistens; excipulum angustum, decolor, 15-18 mmm. crassum, dimidiatum, ex hyphis subradiantibus et intricatis formatum, non paraplectenchymaticum; hymenium superne anguste et dilute umbrino-fuscescens, caeterum decolor et purum, J violaceo-coeruleum, 75-80 mmm. altum; paraphyses capillari-filiformes, densae, strictulae, simplices, eseptatae, ad apicem non latiores, gelatinose conglutinatae; asci oblongo-clavati, rotundati et membrana bene incrassata cincti, monospori; sporae decolores, oblongae, utrinque rotundatae, rectae vel subrectae, murales, cellulis numerosis, subcubicis et leptodermaticis, membrana tenui cinctae, 60-64 mmm. longae et 12-14 mmm. latae.
On the leaves of Metrosideros Colensoi, Alectryon excelsum.
7. Lopadium subcoerulescens A. Zahlbr. nov. spec.
Thallus epiphyllus, maculas minores, usque ad 5 mm. latas, rotundato-irregulares et passim confluentes formans, substratum arcte obducens, tenuissimus, albidus vel soridescenti-albidus, opacus, KHO-, Ca Cl2O2—, continuus, laevigatus, granulis non obsitus nec trichomatibus praeditus, sorediis et isidiis nullis, in margine linea obscuriore non cinctus. Apothecia parva, 0.1-0.3 mm. lata, fusco-nigra vel nigricantia, opaca, sessilia, rotunda, ad basin parum constrictula, e convexiusculo demum convexula; margo tenuis, integer, mox depressus, disco concolor, rare in juventute pallescens; recepta-culum extus nigricans; excipulum dimidiatum, molle, crassiusculum, subcoerulescenti-fuscescens, ex hyphis intricatis formatum, gonidia non includens; hymenium superne latiuscule sordido-coerulescens, non inspersum, caeterum decolor et purum, 70-80 mmm. altum, J coeruleum; hypothecium umbrino-fuscescens et demum nigricans, molle, sat angustum; paraphyses graciles, filiformes, ramosae eseptatae, ad apicem non latiores, laxiuscule conglutinatae; asci ellipsoideo-clavati, superne bene rotundati et membrana bene-incrassata cincti, monospori; sporae decolores, oblongae, utrinqne rotundatae vel in uno apice subangusto-rotundatae, subrectae, murales, cellulis numerosis, subcubicis, leptodermaticis, in seriebus superpositis 16-18, in seriebus horizontalibus 2-7, membrana tenui cinetae, 50-60 mmm. longae et 12-16 mmm. latae.
The most frequent of the epiphyllous lichens. On the leaves of Beilschmiedia tawa, Myrius bullata, Alectryon exculsum, Asplenium bulbiferum, polystichum vestitum.
8. Phyacia crispa Nyl.
On the leaves of Beilschmiedia tawa, Podocarpus spicatus.
9. Physcia tremens A. Zahlbr. nov. spec.
Thallus dichotome et sympodialiter laciniatus, radiatim crescens, tenuissimus submembranaceus, substrato arcte adpressus, albidus, KHO—, Ca Cl2O2—, laciniis linearibus, primum ad 0.2 mm. latis, planis, plus minusve discurrentibus, in margine subintegris vel leviter sinuato-dentatis, ad apicem parum latioribus, rotundatis vel
subemarginatis, demum latioribus, usque ad 1.5 mm. latis, plus minusve confluentibus et contiguis, superne paulum inaequalibus, sorediis et isidiis destitutus in margine non ciliatus, subtus pallidus, dilute et sordide fuscescens, rhizinis brevissimis et increbris; medulla alba, KHO—. Apothecia non visa.
On the leaves of Alectryon excelsum, Podocarpus spicatus.
C. A New Lichen-Parasite (Karl Keissler).
Chlorocyphella lichenicola Keissler nov. spec.
Recptaculis oblique cupularibus auricularibus, sessilibus, sub-membranaceis, obscure, aeruginascentibus demum expallescentibus, usque ad 1 mm. longis; hymenio concolore; basidiis?; basidiosporis hyalinis, filiformibus, apice incrassatis, dense septatis, primum sub-curvis deinde curvatis, saepe rectangulariter inflexis, non spinulosis, Ca. 25-50 × 2 mmm.
Hab. Neu-Seeland, in thallo sterili Lichenis ad folia Myrti bullatae, et in thallo Lopadii subcaerulescentis ad folia Polystichi vestiti et Alectryonis excelsi, Rain forest, Feilding, leg. H. H. Allan (herb. Mus. hist. nat. Vindob.)
Dieser merkwürdige Flechtenparasit, den ich übrigens auch schon aus China gesehen habe, macht in vieler Beziehung den Eindruck einer Cyphella, hat aber nicht kurze runde, ovale oder eifömige Sporen, wie sie bei dieser Gattung vorkommen, sondem solche von linealer Grestalt und bedeutender Länge. Nach diesem Merkmal passt dieser typus anachienend in die Gattung Chlorocyphella Speg.,* beschrieben in An. Mus. Nac. Buenos Aires T. 19, 1909, p. 279 (cfr. Saccardo, Syll. fungor., vol. 21, p. 424). Spegazzini hat dieses Genus als einen Vertreter der Hymenolichenen beschrieben, indem er der Meinung war, dass der Thallus auf dem die Cyphella-artigen Gehäuse sitzen, zu diesem dazu gehöre, das ganze also eine Flechte darstelle. Wenn man aber so wie hier bei Chl. lichenicola sieht, dass die Fruehtkörper bald auf sterilem Thallus, bald auf dem Thallus von Lopadium subcaerulescens aufsitzen, kommt man zur Veberzeugung, dass die Cyphella-artigen Gehäuse nicht als Fruktifikation zu dem Thallus anzusehen sind, sondern dass dieselben auf diesem als Parasiten auftreten. Es dürfte also Chlorocyphella Speg. als eine Gattung zu behandein sein, die nicht zu den Hymenolichenes sondern zu den Flechtenparasiten zu zählen ist. Die einzige von Spegazzini beschriebene Art ist Chl. subtropica, von Formosa angegeben. Die von mir aufgestellte Art unterscheidet sich von jener durch die grünlichen (nicht weisslichen oder rosagrau gefärbten) Fruchtbecher mit schiefer Müudung (ähnlich Cyphella capula), und durch die oft rechtwinklig eingebogenen dicht septierten, am ende kopforimg, verdickten Sporen.
In der Farbe und Gestalt der Becher erinnert die neu aufgestellte Art an C. aeruginascens Karst. (cfr. Sacc. Syll. fung., vol. 9, p. 247) von Wainio an Baumrinde in Minas Geraes (Sitio) gesammelt. Diese wird von Wainio, “Études Lich. Brésil,” in Act. Soc. Faun. Flor. Fernn., vol. 7, 1890, pars 2, p. 27, auch für Rio de Janeiro angeben und dort ausdrücklich als Flechtenparasit auf dem Thallus von Lecidia perpallida bezeiehnet. Karsten gibt kugelige
[Footnote] *Exemplare dieser Gattung habe ich keine in Händen gehabt, aber inzwichsen nach Buenos Aires darum geschrieben.
Sporen (allerdings mit Fragezeichen) an. An dem Originalexemplar, welches ich zum Vergleich von Herrn Direktor Elfing in Helsingfors erhielt, warem leider keine solchen zu sehen, was fast den Gedanken nahe legt, dass Karsten auch keine wahrgenommen habe tuid nur vermutete, dass sie dem Cyphella Typus entsprächen. Es wäre nicht undenkbar, dass die von mir aufgestellte Chl. lichenicola nur das mit Sporen versehene Stadium von Cyphella aeruginascens Karst. darstellt. Zu erwähnen ware noch C. foliicola Wain., “Lich. Phillipp. III.” in Ann. Acad. Scient. Fenn., vol. 15, nr. 6, 1921, p. 83, auf blattbewohnenden Thallus von Bilimbia polillensis, welche nach den langfädigen Sporen gewiss als Chlorocyphella (Chl. foliicola Keissl.) anzusprechen ist. Von dieser ist Chl. lichenicola durch die grüne, nicht blasse Farbe der Receptacula, souie durch das Fehlen der an der Innenseite der Sporenkrummung entwickelten, in einer Reihe stehenden Dornen verschieden.
Zum Schlusse sei noch bemerkt, falls die Receptacula der Chl. lichenicola mit einer gewissen Regelmassigkeit auf Lopadium-Thallus vorkommen sollten, man dieselben doch als eine Nebenfruktifikation der Flechten aussehen musste, namentlich dann, wenn es etwa gelingen sollte, Gonidien in den Receptacula nachzuweisen.
[It is thought advisable to add the following Abstract of Keissler's remarks on Chlorocyphella lichenicola Keissler nov. spec. This noteworthy lichen-parasite, which occurs also in China, has many resemblances to Cyphella, but falls into Chlorocyphella Speg. owing to its long linear spores. Spegazzini considered his genus to belong to the Hymenolichens, interpreting the cyphella-like structures as belonging to the thallus on which they occurred. When it is noted that the fruit bodies of Keissler's species occur now on a sterile thallus, now on that of Lopadium subcaerulescens it must be concluded that it is parasitic. The only species described by Spegazzini was Chl. subtropica, from Formosa. The present species differs in the greenish (not whitish or rosy-grey) fruit-cups with more oblique mouths (similar to those of Cyphella capula), and in the thickly septate spores, often bent at right angles, and with thickened head-shaped ends.
In the colour and structure of the cups the new species recalls C. aeruginascens Karst., collected by Wainio in Minas Geraes. Wainio records this also from Rio de Janiero, and expressly designates it as a lichen-parasite on the thallus of Lecidia perpallida. Karsten gives the spores as globular (with a query). No spores could be found by Keissler on the original specimen, and he thinks it probable that Karsten had not seen any, but had only conjectured that they were of Cyphella form.
It may be that Chl. lichenicola is a spore-bearing state of C. aeruginascens. C. foliicola Wain., on the leaf-dwelling thallus of Bilimbia pollinensis with long linear spores, is certainly a Chlorocyphella (Chl. foliicola Keissel.). C. lichenicola differs from this in its green, not pale, cups and the absence of a row of spinous processes on the inner side of the curvature of the spores.
If the receptacula of Chl. lichenicola occur with a certain regularity upon the thallus of Lopadium, they must be considered as a companion fructification (Nebenfruchtifikation) of the lichen, especially if gonidia are found therein..—H.H.A.]