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Seasonal observations on ‘aquatic’ hyphomycetes on oak leaves on the ground
Notes and brief articles
108
the maximum concentration (1 '0 em") of the fumigant after 15 days. These species formed the dominant group of recolonizers of the soil, after 60 days, at the higher concentrations of formalin. In addition, A. fiavus, A. fumigatus, Penicillium citrinum, P. rubrum and A. terreus also exhibited tolerance to formalin.The fumigant had a depressing effect on A. niger. Rhizopus nigricans, A. nidulans and A. luchuensis appeared to be completely intolerant. Trichoderma oiride, which has been recorded as dominant recolonizer of the fumigated soil by several investigators, was hardly recorded from treated and untreated soils after 60 days. The direct or indirect effect of formalin on Aspergillus and Penicillium leading to an increase in soil has also been observed by Tam & Clark (1943), and Moubasher & Mazen (1971) noted the tolerance of A. fumigatus, A. fiavus and P. notatum to formalin in various doses in Eygptian alkaline soil. The increase in fungal colonies at 0'1 and o·5 em" concentrations after 15 days was due to an increase ofunaffected white and brown sterile mycelia at the former concentration and A. sydowii and A. humicola at the latter one. The authors thank the Head of the Department of Botany, Banaras Hindu University for laboratory facilities. REFERENCES
BLISS, E. D. (1951). The destruction of Armillaria mellea in citrus soil. Phytopathology 41, 665-683. EVANS, E. (1955). Survival and recolonization by fungi in soil treated with formalin or carbon disulphide.
Transactions of the British Mycological Society 38, 335-348. GARRETT, S. D. (1957). Effect of a soil microflora
selected by carbon disulphide fumigationon survival of Armillaria mellea in woody host tissue. Canadian
Journal of Microbiology 3, 135-149.
GARRETT, S. D. (1965). Towards biological control of soil borne plant pathogens. In Ecology of soil-borne plant pathogens (ed. K. F. Baker and W. C. Snyder), pp. 4-16. University of California Press. LILY, K. (1961). Ecological studies on soilfungi. Ph.D. Thesis, University of Saugar, India. MARTIN, J. P. (1950). Effect of fumigation and other soil treatments in the green house on the fungal population of old citrus soil. Soil Science69, 107-122. MOLLISON, J. E. (1953). Effect of partial sterilization and acidification of soil on fungal population.
Transactions of the British Mycological Society 36, 215-228. MOUBASHER, A. H. (1963). Selectiveeffects of fumiga-
tion with carbon disulphide on the fungus flora.
Transactions of the British Mycological Society 46, 338-344. MOUBASHER, A. H. & MAZEN A. B. (1971). Selective
effects of three fumigants on Egyptian soil fungi.
Transactions of the British Mycological Society 57, 447-454· MUGHOGHO, L. K. (1968). The fungus flora of fumigated soils. Transactions of the British Mycological Society 51,441-459. OVERMAN, A. J. & BURGIs, D. S. (1956). Allyl alcohol as soil fungicide. Phytopathology 46, 532-635. SAKSENA, S. B. (1960). Effect of carbon disulphide fumigation on Trichoderma viride and other soil fungi. Transactions of the British Mycological Society 43,111-116. TAM, R. K. & CLARK, H. E. (1943). Effect of chloro-
picrin and other soil disinfectants on the nitrogen nutrition of the pineapple plant. Soil Science 56,
245-26t.
WAKSMAN, S. A. & STARKEY, R. L. (1923). Partial sterilization of soil microbiological activities and soil fertility. I, II, III. Soil Science16, 137-156; 247-268; 343-357·
WARCUP, J. H. (1951). Effect of partial sterilization by steam or formalin on the fungus of an old forest nursery soil. Transactions of the British Mycological Society 34, 519-53 2.
SEASONAL OBSERVATIONS ON 'AQUATIC' HYPHOMYCETES ON OAK LEAVES ON THE GROUND
J. WEBSTER Department of Biological Sciences, University of Exeter
Although certain hyphomycetes, notably those with branched or sigmoid spores are abundant and cosmopolitan on leaves and twigs in fresh waters and their spores are present in stream and river foam (Ingold, 1975), evidence is accumulating (Bandoni, 1972, 1974), Park (1974a, b) that some of these fungi are not exclusively aquatic, but their spores may be found on leaves and other substrata Trans. Br. mycol. Soc. 68 (1), (1977).
well away from contact with flowing or standing water. The purpose of this paper is to record monthly observations on the occurrence of spores of such Hyphomycetes during one year. The site chosen for study was a steeply sloping south-facing oak-wood overlooking the River Teign at Steps Bridge, Devon, England, Map reference SX 802884. The site was about 100 m upstream
Printed in Great Britain
Notes and brief articles
1°9 9 I
40
30
10
10m
Fig. 1. Elevation of valley slope at Steps Bridge, showing the position of the nine sampling points. from a weir extending across the river. Marked pegs were driven into the ground at 10 m intervals from the bank of the river . N ine pegs were placed and their heights above river level were determined. Peg 1 was placed at the edge of the river, and peg 9 was sited some 38 m above river level. (Fig. 1) , Pegs 1, 2 and possibly 3 were at heights which were probably occasionally inundated when the river was in flood, as indicated by the accumulation of flotsam on branches of adjacent trees. The woodland was dominated by coppiced oak (Quercus petraea) with a few trees of birch (Betula verrucosa) holly (flex aquijolium) and beech (Fagus sylvatica). The ground flora was dominated by Holcus mollis, Agrostis setacea and Luzula syloatica. Usually, five oak leaves were collected from the surface of the ground as near as possible to each peg and placed in new polythene bags. Sampling began on 13 December 1974, and continued at approximately monthly intervals until November 1975. For a few months early in the sampling period five leaves of the 1974 season and also five of the 1973 season were collected. Later, it became difficult to collect 1973 leaves. (The leaves were collected in new polythene bags, marked with the peg number, and the season of leaf fall.) In the laboratory each leaf was placed in a separate Petri dish to which 20 em" distilled water were added, The dishes were incubated for 4 days at 10 °C. The dishes were then carefully scanned
Trans. Br. mycol. Soc. 68 (1), (1977).
Table 1. Rainfall data for the period December 1974 to No vember 1975 (Wooston Castle Rainfall Station, South West Water Authority) Rainfall (mm) 1914 1975
December January February March April May June July August September October November December
92 '9
138'1 36'S 72'8 57'8 29'8 8'0 16'3 24'1 88'0 33'4 11'0 31'4
under a low-power binocular microscope with a magnification of x 50, using a combination of transmitted and reflected light. The meniscus, the leaf margins and the bottom of the dish were searched for hyphomycete spores, and where possible spores or conidiophores were transferred to slides for confirmation of identity. Careful note was made of the occurrence of single or isolated conidia, numerous conidia, and especially of the presence of conidiophores attached to the :leaf. In some cases it was not possible to determine with confidence the identity of certain spores.
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Notes and brief articles
110
Table 2. Records of aquatic hyphomycete spores at various levels above the River Teign at Steps Bridge on monthly samples of oak leaves
Peg no. Fungus species
1
2
3
4
5
,-~
~
~
~
~
1973
Anguillospora longissima Articulospora tetracladia Camposporium pellucidum Clavariopsis aquatica Dendrospora erecta Dendrospora sp. Flagellospora curvula Heliscus stellatus Isthmotricladia sp. Lemonniera aquatica Pleuripedium tricladioides Tetrachaetum elegans Tetraploa aristata Tricladium chaetocladium Tricladium splendens Triposporium elegans Varicosporium elodeae
1974
1973
1974
1973
1/20 5/20* 12/60 1/60 1/20* 1/60
3/20
1/60
1/20
6/20
2/60*
Anguillospora longissima Articulospora tetracladia Camposporium pellucidum Clavariopsis aquatica Dendrospora erecta Dendrospora sp. Flagellospora curvula Heliscus stellatus Isthmotricladia sp. Lemonniera aquatica Pleuripedium tricladioides Tetrachaetum elegans Tetraploa aristata Tricladium chaetocladium Tricladium splendens Triposporium elegans Varicosporium elodeae
*
1973
1974
1/60
1973
1/60
1/20*
1/20 3/20* 2/20*
2/60 2/60 1/60 1/60 1/60 15/60
2/20 8/20
1/60
2/20
1/20
2/20 6/20
6/60 4/ 60
6
7 ~
1974
1/20
1/60
2/60 1/60
2/20
1/60
1/60
~
1973
1974
2/60
3/20
1/20
Peg no.
1974
1973
1/20
1/20 8
1974
3/60
,----A-..,
1/20 9
~
1973
1974
1973
1974
1/20
1/60
5/ 20
1/60 1/60
1/20 2/20
1/20
1/20
3/ 60
1/20
1/20
1/20
2/60
Denotes presence of conidiophores on certain leaves at this level.
Meteorological records for the period from the Wooston Castle Rainfall station some 4 km northwest of Steps Bridge are presented in Table 1. The summer of 1975 was unusually warm and dry. Results of the observations are presented in Tables 2 and 3. In deciding which spores to record some subjective judgement was inevitable. In general, those recognizable spores which had previously been reported from aquatic habits were included, whereas those which are usually associated with Trans. Br. mycol, Soc. 68 (1), (1977).
terrestrial substrata, such as species of Cladosporium, Alternaria, Botrytis, basidiospores of agarics, or urediospores of rust fungi were not recorded. Fructifications of Protostelida were seen frequently floating on the meniscus, but these were not recorded. Although it is difficult to summarize observations on 560individual leaves,certain trends areapparent, (1) More spores were observed on leaves which had been on the ground for several months, than
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Notes and briefarticles
111
Table 3. Total records of all species of aquatic hyphomycetes (The denominator indicates the number of leaves sampled) Peg no. Sampling date 13. xii. 74 17.i.75 14. ii.75 14. iii. 75 19. iv. 75 23· v.75 23. vi. 75 18. vii. 75 15. viii. 75 26. ix. 75 20. x. 75 28. xi. 75 Totals
1 3/ 10 17/10 13/10 8/10 3/5 6/5 2/5 2/5 1/5 4/5 3/5 62/80
2
3
11/10 9/10 9/10
1/10 2/10 1/10 1/10
4
5
6
7
1/10
2/10
2/10
1/10 1/10
5/5
1/5
36/80
6/80
1/5 2/80
4/5 2/5 8/80
3/80
9
1/10 1/10
1/10 7/ 10 1/10
1/5
1/5
2/5
8
1/5 3/80
2/5 1/5 1/5 3/5
3/80
16/80
N.B. Zero results have not been included in the Table. those which had been there for shorter periods. It is not clear whether this reflects the time taken to colonize leaves, or longer opportunity to accumulate spores, or whether the moister conditions beneath the surface layers are more conducive to spore production, survival and dispersal. (2) The number of records near pegs 1-2 is greater than at other levels. In view of the possibility of occasional flooding bringing leaves from the river to these levels, this is not surprising. (3) Conidia attached to conidiophores were only found near pegs 1-3. However, the frequency was very low, condiophores being detected in only 11 out of a sample of 560 oak leaves (1'96 'Yo). This is a much lower frequency ofconidiophore production than would be expected from leaves taken from the river. It would obviously be of interest to determine the survival of hyphomycetes on leaves removed from rivers. The origin of the conidia found at the higher levels is, at present, unknown. Whilst it is possible that these fungi have a terrestrial existence in the mycelial state at these levels, the mere existence of spores attached to leaves, is, of course, no proof of this. The numbers of conidia found floating on the meniscus were often very low, sometimes only one
Trans. Br, mycol. Soc. 68 (1), (1977).
or two per dish, and this again contrasts with the large numbers of spores which are found, after incubation in water, on leaves removed from streams.
I am indebted to the South West Water Authority for the rainfall data, and to Mr R. A. Davey for competent technical assistance. REFERENCES
BANDONI, R. J. (1972). Terrestrial occurrence of some aquatic Hyphomycetes. Canadian Journal of Botany S0, 2283-2288. BANDONI, R. J. (1974). Mycological observations on the aqueous films covering decaying leaves and other litter. Transactions of the Mycological Society of Japan 15, 309-315.
INGOLD, C. T. (1975). An illustrated guide to aquatic water-borne Hyphomycetes (Fungi Imperfecti) with notes on their biology. Freshwater Biological Association Scientific Publication No. 30, 96 pp. PARK, D. (1974a). Tricladium terrestre sp.nov. Transactions of the British Mycological Society 63, 179-183. PARK, D. (1974b). Aquatic Hyphomycetes in nonaquatic habitats. Transactions of the British Mycological Society 63, 183-187.
Printed in Great Britain
108
the maximum concentration (1 '0 em") of the fumigant after 15 days. These species formed the dominant group of recolonizers of the soil, after 60 days, at the higher concentrations of formalin. In addition, A. fiavus, A. fumigatus, Penicillium citrinum, P. rubrum and A. terreus also exhibited tolerance to formalin.The fumigant had a depressing effect on A. niger. Rhizopus nigricans, A. nidulans and A. luchuensis appeared to be completely intolerant. Trichoderma oiride, which has been recorded as dominant recolonizer of the fumigated soil by several investigators, was hardly recorded from treated and untreated soils after 60 days. The direct or indirect effect of formalin on Aspergillus and Penicillium leading to an increase in soil has also been observed by Tam & Clark (1943), and Moubasher & Mazen (1971) noted the tolerance of A. fumigatus, A. fiavus and P. notatum to formalin in various doses in Eygptian alkaline soil. The increase in fungal colonies at 0'1 and o·5 em" concentrations after 15 days was due to an increase ofunaffected white and brown sterile mycelia at the former concentration and A. sydowii and A. humicola at the latter one. The authors thank the Head of the Department of Botany, Banaras Hindu University for laboratory facilities. REFERENCES
BLISS, E. D. (1951). The destruction of Armillaria mellea in citrus soil. Phytopathology 41, 665-683. EVANS, E. (1955). Survival and recolonization by fungi in soil treated with formalin or carbon disulphide.
Transactions of the British Mycological Society 38, 335-348. GARRETT, S. D. (1957). Effect of a soil microflora
selected by carbon disulphide fumigationon survival of Armillaria mellea in woody host tissue. Canadian
Journal of Microbiology 3, 135-149.
GARRETT, S. D. (1965). Towards biological control of soil borne plant pathogens. In Ecology of soil-borne plant pathogens (ed. K. F. Baker and W. C. Snyder), pp. 4-16. University of California Press. LILY, K. (1961). Ecological studies on soilfungi. Ph.D. Thesis, University of Saugar, India. MARTIN, J. P. (1950). Effect of fumigation and other soil treatments in the green house on the fungal population of old citrus soil. Soil Science69, 107-122. MOLLISON, J. E. (1953). Effect of partial sterilization and acidification of soil on fungal population.
Transactions of the British Mycological Society 36, 215-228. MOUBASHER, A. H. (1963). Selectiveeffects of fumiga-
tion with carbon disulphide on the fungus flora.
Transactions of the British Mycological Society 46, 338-344. MOUBASHER, A. H. & MAZEN A. B. (1971). Selective
effects of three fumigants on Egyptian soil fungi.
Transactions of the British Mycological Society 57, 447-454· MUGHOGHO, L. K. (1968). The fungus flora of fumigated soils. Transactions of the British Mycological Society 51,441-459. OVERMAN, A. J. & BURGIs, D. S. (1956). Allyl alcohol as soil fungicide. Phytopathology 46, 532-635. SAKSENA, S. B. (1960). Effect of carbon disulphide fumigation on Trichoderma viride and other soil fungi. Transactions of the British Mycological Society 43,111-116. TAM, R. K. & CLARK, H. E. (1943). Effect of chloro-
picrin and other soil disinfectants on the nitrogen nutrition of the pineapple plant. Soil Science 56,
245-26t.
WAKSMAN, S. A. & STARKEY, R. L. (1923). Partial sterilization of soil microbiological activities and soil fertility. I, II, III. Soil Science16, 137-156; 247-268; 343-357·
WARCUP, J. H. (1951). Effect of partial sterilization by steam or formalin on the fungus of an old forest nursery soil. Transactions of the British Mycological Society 34, 519-53 2.
SEASONAL OBSERVATIONS ON 'AQUATIC' HYPHOMYCETES ON OAK LEAVES ON THE GROUND
J. WEBSTER Department of Biological Sciences, University of Exeter
Although certain hyphomycetes, notably those with branched or sigmoid spores are abundant and cosmopolitan on leaves and twigs in fresh waters and their spores are present in stream and river foam (Ingold, 1975), evidence is accumulating (Bandoni, 1972, 1974), Park (1974a, b) that some of these fungi are not exclusively aquatic, but their spores may be found on leaves and other substrata Trans. Br. mycol. Soc. 68 (1), (1977).
well away from contact with flowing or standing water. The purpose of this paper is to record monthly observations on the occurrence of spores of such Hyphomycetes during one year. The site chosen for study was a steeply sloping south-facing oak-wood overlooking the River Teign at Steps Bridge, Devon, England, Map reference SX 802884. The site was about 100 m upstream
Printed in Great Britain
Notes and brief articles
1°9 9 I
40
30
10
10m
Fig. 1. Elevation of valley slope at Steps Bridge, showing the position of the nine sampling points. from a weir extending across the river. Marked pegs were driven into the ground at 10 m intervals from the bank of the river . N ine pegs were placed and their heights above river level were determined. Peg 1 was placed at the edge of the river, and peg 9 was sited some 38 m above river level. (Fig. 1) , Pegs 1, 2 and possibly 3 were at heights which were probably occasionally inundated when the river was in flood, as indicated by the accumulation of flotsam on branches of adjacent trees. The woodland was dominated by coppiced oak (Quercus petraea) with a few trees of birch (Betula verrucosa) holly (flex aquijolium) and beech (Fagus sylvatica). The ground flora was dominated by Holcus mollis, Agrostis setacea and Luzula syloatica. Usually, five oak leaves were collected from the surface of the ground as near as possible to each peg and placed in new polythene bags. Sampling began on 13 December 1974, and continued at approximately monthly intervals until November 1975. For a few months early in the sampling period five leaves of the 1974 season and also five of the 1973 season were collected. Later, it became difficult to collect 1973 leaves. (The leaves were collected in new polythene bags, marked with the peg number, and the season of leaf fall.) In the laboratory each leaf was placed in a separate Petri dish to which 20 em" distilled water were added, The dishes were incubated for 4 days at 10 °C. The dishes were then carefully scanned
Trans. Br. mycol. Soc. 68 (1), (1977).
Table 1. Rainfall data for the period December 1974 to No vember 1975 (Wooston Castle Rainfall Station, South West Water Authority) Rainfall (mm) 1914 1975
December January February March April May June July August September October November December
92 '9
138'1 36'S 72'8 57'8 29'8 8'0 16'3 24'1 88'0 33'4 11'0 31'4
under a low-power binocular microscope with a magnification of x 50, using a combination of transmitted and reflected light. The meniscus, the leaf margins and the bottom of the dish were searched for hyphomycete spores, and where possible spores or conidiophores were transferred to slides for confirmation of identity. Careful note was made of the occurrence of single or isolated conidia, numerous conidia, and especially of the presence of conidiophores attached to the :leaf. In some cases it was not possible to determine with confidence the identity of certain spores.
Printed in Great Britain
Notes and brief articles
110
Table 2. Records of aquatic hyphomycete spores at various levels above the River Teign at Steps Bridge on monthly samples of oak leaves
Peg no. Fungus species
1
2
3
4
5
,-~
~
~
~
~
1973
Anguillospora longissima Articulospora tetracladia Camposporium pellucidum Clavariopsis aquatica Dendrospora erecta Dendrospora sp. Flagellospora curvula Heliscus stellatus Isthmotricladia sp. Lemonniera aquatica Pleuripedium tricladioides Tetrachaetum elegans Tetraploa aristata Tricladium chaetocladium Tricladium splendens Triposporium elegans Varicosporium elodeae
1974
1973
1974
1973
1/20 5/20* 12/60 1/60 1/20* 1/60
3/20
1/60
1/20
6/20
2/60*
Anguillospora longissima Articulospora tetracladia Camposporium pellucidum Clavariopsis aquatica Dendrospora erecta Dendrospora sp. Flagellospora curvula Heliscus stellatus Isthmotricladia sp. Lemonniera aquatica Pleuripedium tricladioides Tetrachaetum elegans Tetraploa aristata Tricladium chaetocladium Tricladium splendens Triposporium elegans Varicosporium elodeae
*
1973
1974
1/60
1973
1/60
1/20*
1/20 3/20* 2/20*
2/60 2/60 1/60 1/60 1/60 15/60
2/20 8/20
1/60
2/20
1/20
2/20 6/20
6/60 4/ 60
6
7 ~
1974
1/20
1/60
2/60 1/60
2/20
1/60
1/60
~
1973
1974
2/60
3/20
1/20
Peg no.
1974
1973
1/20
1/20 8
1974
3/60
,----A-..,
1/20 9
~
1973
1974
1973
1974
1/20
1/60
5/ 20
1/60 1/60
1/20 2/20
1/20
1/20
3/ 60
1/20
1/20
1/20
2/60
Denotes presence of conidiophores on certain leaves at this level.
Meteorological records for the period from the Wooston Castle Rainfall station some 4 km northwest of Steps Bridge are presented in Table 1. The summer of 1975 was unusually warm and dry. Results of the observations are presented in Tables 2 and 3. In deciding which spores to record some subjective judgement was inevitable. In general, those recognizable spores which had previously been reported from aquatic habits were included, whereas those which are usually associated with Trans. Br. mycol, Soc. 68 (1), (1977).
terrestrial substrata, such as species of Cladosporium, Alternaria, Botrytis, basidiospores of agarics, or urediospores of rust fungi were not recorded. Fructifications of Protostelida were seen frequently floating on the meniscus, but these were not recorded. Although it is difficult to summarize observations on 560individual leaves,certain trends areapparent, (1) More spores were observed on leaves which had been on the ground for several months, than
Printed in Great Britain
Notes and briefarticles
111
Table 3. Total records of all species of aquatic hyphomycetes (The denominator indicates the number of leaves sampled) Peg no. Sampling date 13. xii. 74 17.i.75 14. ii.75 14. iii. 75 19. iv. 75 23· v.75 23. vi. 75 18. vii. 75 15. viii. 75 26. ix. 75 20. x. 75 28. xi. 75 Totals
1 3/ 10 17/10 13/10 8/10 3/5 6/5 2/5 2/5 1/5 4/5 3/5 62/80
2
3
11/10 9/10 9/10
1/10 2/10 1/10 1/10
4
5
6
7
1/10
2/10
2/10
1/10 1/10
5/5
1/5
36/80
6/80
1/5 2/80
4/5 2/5 8/80
3/80
9
1/10 1/10
1/10 7/ 10 1/10
1/5
1/5
2/5
8
1/5 3/80
2/5 1/5 1/5 3/5
3/80
16/80
N.B. Zero results have not been included in the Table. those which had been there for shorter periods. It is not clear whether this reflects the time taken to colonize leaves, or longer opportunity to accumulate spores, or whether the moister conditions beneath the surface layers are more conducive to spore production, survival and dispersal. (2) The number of records near pegs 1-2 is greater than at other levels. In view of the possibility of occasional flooding bringing leaves from the river to these levels, this is not surprising. (3) Conidia attached to conidiophores were only found near pegs 1-3. However, the frequency was very low, condiophores being detected in only 11 out of a sample of 560 oak leaves (1'96 'Yo). This is a much lower frequency ofconidiophore production than would be expected from leaves taken from the river. It would obviously be of interest to determine the survival of hyphomycetes on leaves removed from rivers. The origin of the conidia found at the higher levels is, at present, unknown. Whilst it is possible that these fungi have a terrestrial existence in the mycelial state at these levels, the mere existence of spores attached to leaves, is, of course, no proof of this. The numbers of conidia found floating on the meniscus were often very low, sometimes only one
Trans. Br, mycol. Soc. 68 (1), (1977).
or two per dish, and this again contrasts with the large numbers of spores which are found, after incubation in water, on leaves removed from streams.
I am indebted to the South West Water Authority for the rainfall data, and to Mr R. A. Davey for competent technical assistance. REFERENCES
BANDONI, R. J. (1972). Terrestrial occurrence of some aquatic Hyphomycetes. Canadian Journal of Botany S0, 2283-2288. BANDONI, R. J. (1974). Mycological observations on the aqueous films covering decaying leaves and other litter. Transactions of the Mycological Society of Japan 15, 309-315.
INGOLD, C. T. (1975). An illustrated guide to aquatic water-borne Hyphomycetes (Fungi Imperfecti) with notes on their biology. Freshwater Biological Association Scientific Publication No. 30, 96 pp. PARK, D. (1974a). Tricladium terrestre sp.nov. Transactions of the British Mycological Society 63, 179-183. PARK, D. (1974b). Aquatic Hyphomycetes in nonaquatic habitats. Transactions of the British Mycological Society 63, 183-187.
Printed in Great Britain