Taxonomic studies on aquatic hyphomycetes: I. Lemonniera de Wildeman

Tram . Br. mycol. Soc. 69 (1) 89-109 (1977) Printed in Great Britain

TAXONOMIC STUDIES ON AQUATIC HYPHOMYCETES

I. LEMONNIERA DE WILDEMAN

By E. DESCALS C. AND J. WEBSTER

Department of Biological Sciences, University of Exeter

AND B.S.DYKO

Department of Botany, University of Tennessee, Knoxville, Tenn., U.S.A.

This first paper in a series of taxonomic revisions of the genera of freshwater aquatichyphomycetes describes general methods of isolation, culture and description. The sixaccepted species in Lemonniera de Wild. are illustrated. An extension of the generic conceptis proposed, an artificial key to the species is included, L. pseudofioscula sp.nov, is described,and nco-types are provided for L. aquatica, L. cornuta and L. filiformis.

De Wildeman (1893, 1894, 1895) drew attentionto the abundance of freshwater hyphomycetescolonizing submerged plant material. It was onlyafter Ingold (1942) published his paper that thisgroup has been fully appreciated. There is now aneed for taxonomic revision because severalgenera contain heterogeneous elements. The mostrecent comprehensive study is that of Dudka(1974) in Ukrainian, which was preceded byNilsson's in 1964.

Some members of the ' freshwater hyphomy-cetes " i.e. ' those with aquatic spores produced,liberated and normally dispersed underwater'(Ingold, 1942), have conidia which are morpho-logically similar to those of other groups, such assome terrestrial Sphaeropsidales, rhizopod-trap-ping fungi, aero-aquatic and dematiaceous hy-phomycetes, and marine and brackish-waterhyphomycetes. A few of them now are known toproduce ascomycete and basidiomycete perfectstates. Certain form species originally consideredaquatic are being consistently found in terrestrialhabitats (Bandoni, 1972; Park, 1972). Othersproduce their conidia readily on the surface ofwater or in atmospheres of high relative humidity.Therefore the term 'aquatic hyphomycete' mustbe interpreted in a rather loose sense. This seriesof papers will attempt to revise the taxonomy oftrue freshwater hyphomycetes as well as imper-fect states of other fungi isolated from aquatichabitats.

GENERAL METHODS

Collections are made of submerged decayingplant material and foam from freshwater. Sites arespecified in the individual species descriptions.

Foam is scooped up with clean plastic vials andbags and taken to the laboratory on the same day.The resultant suspensions are placed in glass

Petri dishes, and both the meniscus and the bottomof the dish scanned through transmitted light at100 or 200 power with a dissecting microscope.Spores on the bottom of the dish are picked upindividually with a Pasteur pipette drawn to avery fine tip, and placed in drops in plastic Petridishes on a dilute-nutrient medium which is usually0'1 % malt extract agar supplemented with Crys-tamycin 0'008 % (Glaxo Laboratories, Greenford,England). These are diluted with another drop ofsterile distilled water, which helps to reducecontamination by undetected smaller fungal spores.Floating spores can be easily picked up with aneyelash or flamed sewing needle, and placed on thesame medium, with or without previous washingsin drops of distilled water.

The isolated spores are allowed to germinate at15 °C. Germ-tubes are normally distinguishableafter 1-2 days. A small piece of agar containingthe spore is cut out with a flamed, spear-pointedneedle and transferred on to a clean area on thesame dish. There, the colony is allowed to growfor two to several days. A small piece from themargin is transferred to 15 ml slopes of 2 % maltextract agar or other suitable medium, in McCart-ney bottles, and stored at 15°.

Colonies still contaminated with bacteria arewashed in running tapwater for one to severaldays. They are rinsed in several changes of steriledistilled water and small pieces transferred on toclean isolation medium.

Plant material is collected in plastic bags andwashed several times in the laboratory in tapwater,Single leaves or pieces of stem are incubated indistilled water in glass Petri dishes or othercontainers for two to several days at 15 °. Single-spore isolates are established as above. Conidia-phores on margins of leaves and petioles, as well asmature detached spores, are mounted in water formeasurements, drawings and microphotography.

Taxonomic studies on aquatic hyphomycetes

For herbarium purposes, semi-permanent slidesare prepared in lactophenol cotton blue and sealedwith Glyceel (George T. Gurr, Searle ScientificServices, High Wycombe, Bucks., England).

When non-sporulating colonies have developedto at least 2 em diam, they are induced to sporu-late by submerging pieces in sterile distilled orstream water in Erlenmeyer flasks or Petri dishes.Erlenmeyer flasks can be aerated with filtered airat the optimum temperature. Isolates of severalspecies may require many days for appreciablequantities of spores to be detected, but manysporulate abundantly within two to three days.

For colony characterization and photography.glass Petri dishes with 20 ml nutrient agar areinoculated with plugs 5 mm diam and incubatedat optimum temperature under diffuse light.

LEMONNIERA DE WILDEMAN

This genus now includes six species with phialidicconidiogenous cells, and typically tetraradiateconidia with branches arising more or less simul-taneously from a globose primordium just abovethe collarette of the phialide.

Lemonniera was erected by De Wildeman (1894)in honour of Lemonnier, Professor of Botany atNancy (France). and based on L. aquatica, whichwas found on submerged leaves in the NancyBotanical Gardens. The species was describedfrom the natural substrate as producing tetraradi-ate as well as globular conidia. De Wildeman alsoreported the aquatic saprophytic nature of thefungus with 'umbellate' multiseptate conidio-phores, terminated by ' basidia-like' apical cellsbearing tetraradiate conidia. The genus wasplaced in the Mucedineae (Moniliaceae-Stauro-sporae). De Wildeman (1895) narrowed thespecies concept. excluding the globular conidiawhich were believed to be another fungus.

Ingold (1942) isolated spores of L. aquaticafrom submerged leaves of Alnus glutinosa in astream in Leicestershire, England. He redescribedthe species from a pure culture. Through carefulobservation of conidial development the typespecies was further characterized. Particularimportance was placed on the necessity of sub-mersion for sporulation. Petersen (1963) laterdetected conidia directly on agar but there is nodoubt that submersion greatly enhances sporula-tion.

Ingold also established the phialidic nature ofthe conidiogenous cells. with exogenous develop-ment of the primordium. late appearance of thebasal septum. followed by its cleavage at release.and successive basipetal proliferation. which waslater confirmed by Ranzoni (1953). Conidial arms

are always well developed and arise from theprimordium at the tip of the phialide. Mycelialcords and brown pigmentation were reported,but distinct sclerotia apparently were not formedin pure culture. They have never been reported innature but their presence in agar was later noticedby Ranzoni (1953). Tubaki (1957) and Petersen(1963). Tubaki (1957) further observed sclerotialarrangement in concentric rings. a feature whichmay occur in all six species. This commonconcentric arrangement of both sclerotia andpigmented bands can also be detected on culturesgrown in the dark.

Ranzoni (1953) described Lemonniera comutafrom California. U.S.A. It has cruciate conidiawith curved arms. So far this is the only specieslacking a distal conidial branch (i.e, one in line withthe longitudinal axis ofthe phialide).

Ranzoni also pointed out the genetic instabilityof sclerotial induction in his species, distinguishingcolony types on the basis of sclerotial presenceand distribution. colony pigmentation and type ofconidiophore supporting cells. The heterokaryoticnature of his fungus was assumed in view offrequent spontaneous sectoring.

Tubaki (1958) and Ingold (1958) separatelydescribed isolates of a saprophytic hyphomycetewith phialidic tetraradiate conidia which theyobtained from terrestrial and aquatic habitats re-spectively. Tubaki must have obviously consideredthe ontogenetic and morphological charactersmentioned above. as well as brown pigmenta-tion and presence of sclerotia. of more taxonomicvalue than its probable terrestrial origin (thefungus had been found sporulating aerially oncapsules of Jug/ans; on 0'1 % glucose Czapekagar and oatmeal agar aerial sporulation wasabundant). The species was named L. terrestris,Ingold isolated his fungus from submerged leavesbut did not detect conidia on malt extract agar.The fungus was narned L. brachycladia. Bothfungi are conspecific and Tubaki's epithet pre-vailed because of prior publication.

Conidia of L. terrestris were later isolated fromdiverse bodies of fresh water. either by incubatingleaves found submerged at the time of collectionor directly from foam (Table 2). Also Conway(1970) and Gonczol (1971) described unidentifiedconidia from aquatic sources which appeared tobe of L. terrestris. Profuse sporulation of L.terrestris from leaves incubated underwater in thislaboratory has been noticed on several occasions.On the other hand aerial conidia are quite readilyproduced even on very young cultures grown onthe isolation medium. The fungus. thus. is notstrictly aquatic.

Hyaline conidiophores and conidia are common

E. Descals C., J. Webster and B. S. Dyko 91

to all five species of Lemonniera, with only tworeports (Nilsson, 1962; Wolfe, 1971) indicatingoccasional brown pigmentation in L. terrestris.The conidia in this species may in fact be of adenser and more pearly protoplasmic appearance,and it has also been observed that conidia ofseveral other species frequently darken with age.The hyaline nature of the reproductive structuresis still retained for the genus.

Lemonniera filiformis Petersen [as 'filiforme']was first reported in 1963 from New York, U.S.A.Spores were isolated from submerged decayingleaves of Acer rubrum, and in pure culture wereproduced typically underwater. Although thedistinct basal isthmuses of the conidial armsresembled those of Articulospora tetracladia In-gold, the fungus produced phialides, with conidialarms radiating from the base. Agar cultures alsoproduced both sclerotia and the familiar brownpigmentation in concentric rings. Conidia be-lieved to be of the same species were recentlyisolated by Webster in N.W. Scotland and byDescals in North Wales. Taxonomic charactersmatch those of the original description except forconidial septation. This, however) is considered oflittle taxonomic value as distinctiveness andmorphology of septa change with conidial maturity.

Submerged decaying leaves of Fagus sylvaticacollected in Czechoslovakia by Marvanova (1968)yielded conidia of L. centrosphaera, which wasdescribed from cultures on malt extract agar.Conidia of this fungus were also collected byIngold (1968) from foam in Scotland. They havealso been isolated from aquatic habitats by Web-ster in N.W. Scotland and Descals in NorthWales. Their colony characteristics and conidialontogeny match those of the original description.

The genus as a whole shows varying degrees ofconidial branching, even under natural conditions.This is common among other freshwater hy-

phomycetes. In pure culture) up to 14 % tri- andpentaradiate conidia have been recorded for L.centrosphaera (Marvanova, 1968). High degreesof deformity can also be detected in conidia ofother species when cultures have been sporulatingfor several days in unchanged water. Supplemen-tary arms may appear on the main branches or onthe central body. The distal branch) though) isalways present in conidia of species that typicallyhave it.

Petersen (1963) and Marvanova (1968) ob-served secondary conidia in pure cultures of L.aquatica and L. terrestris respectively. Theybudded out from the tip of a conidial branch.Nevertheless) conidiogenous cells in Lemonnieraare typically phialidic, with exogenous develop-ment of the primordium as opposed to those formswith full conidial differentiation within the coni-diogenous cell.

The taxonomic characters discussed above formthe basis for the emended description of the genus.

LEMONNIERA de Wildeman emend. Descals &Webster

Saprophytic hyphomycete: hyphae initially hya-line, later with brown to reddish diffusible pig-mentation. Sporulation aerial or aquatic. Sclerotiapresent. Conidiophores hyaline, frequently arisingfrom dark-walled swollen cells on sclerotia,macronematous, mononematous or caespitose,erect, rnultiseptate, simple or more frequently withan irregularly branched head; conidiogenous cellsphialidic, hyaline, apical, single or penicillate,arising in succession) ampulliform to cylindrical.Phialoconidia hyaline, always arising from aglobose primordium directly above the collarette,which develops into a more or less distinct swollencentral body with typically four (sometimes threeor five) branches of approximately equal length)straight or curved) arising more or less simul-

(A)(AA)

Artificial key to species of Lemonniera*

Mature detached conidia with all arms distinctly curved.All arms straight or slightly sinuous.(B) Central body distinct, round or tetrahedral.

(C) All arms with basal isthmus.(CC) Basal isthmus always lacking; arms very fine.

(BB) Central body not well-defined.(D) Lateral arms with basal isthmus; distal arm lacking basal isthmus.(DD) Basal isthmus always lacking.']

(E) Arms parallel-walled, typically 60 pm long.(EE) Arms tapering towards the apex; typically 35pm long.

L. cornuta

L. pseudoflosculaL. centrosphaera

L. filiformis

L. aquatica

L. terrestris

* Growth and sporulation under conditions specified in the text.t Conidial size and morphology overlap in the remaining two species; their identification may require pure-

culturing.

Species

L . aquatica

L. centrosphaera

L. cornuta

L. filiformis

L . pseudofloscula

L. terrestris

\0l,)

Table 1. Taxonomic dimensions 0/ Lemonniera spp. (pm)*Conidia

, "-- --.Source Conidiophores Phialidest Central body Armst

De Wildeman, 1894 ? x5-7 ? ?-70 x ?-?Ingold, 1942 100-500 x ?- ? (stipe) ? 20-70 x 3-4Ranzoni, 1953 100-400 x 4-6 20-30 x 5-7 50-120 x ?Tubaki, 1957:1: 100-500 x 3'5-5 '5 15-20(-30) x 3"5 30-80 x 2'5-5

~Dudka, 1962, 1974 60-400 x 3-8 20-30 x 4-7 32-110 X 2-4Wolfe, 1971§ ? 10'5-30(-46) x 2"5-4'5 25 '5-88 x 2-4 <::>

5Our data~

From nature 125- 300 x 4-7 17'5-27'5 x 5-6'5 45-85 x 4-4'5 (:;.F rom cultures§ 75-150 x 3-6 17'5-27'5 x 6-6'5 40-105 x 4-4 '5 ...

Marvanova, 1968 ? 30 x5 6-10 66-142 x 3- 3'5 (apex 3'5-4'5) [DUdka, 1973, 1974 60-300 x 2'5-3 '5 15-30 x 3- 5 6'5-10'5 66-142 x 3-3'5 (apex 3'5-4'5) ~ .

M iura, 1971 60-300 x 2'5-3 '5 15-32 x 3-4 6'5-8'5 60-82 x 1'5-2'5 (at centre) <::>Our data ;:l

F rom nature 200-750 x 5-5'5 17'5-40 x 4-7 6'5-7'5 60-100 x 2' 5- 4 (apex 4-4'5) ]F rom culture§ 125-200 x 3-4'5 14'25 x 4-4'5 6'5-7'5 (35)-78 x 2-3 (apex 3'5-4)

Ranzoni, 1953 50-600 x 5-6 20-30 x 5-7 30-45 x 3-5 (apex 2'5)...(:;.

M iura, 1967 200-450 x 2'5-4 20-25 x 3'5-5 30-55 x 4-4'5 (apex 2-2'5)~Iqbal, 1972 (a, b) 100-500 x 5-6 18-30 x 5 '5 30-52'5 x 3'5 -s

Our data (from cultures)§ 125-175 x 3-5 15- 30 x 5-5'5 12'5-57 x 3-4 5"Petersen, 1963 100-200 x 3- 4'5 ? 60-105 x 3- 4 (distal arm) ~Our data (from cultures)§ 150-300 x 3'5-4 15-17'5 x 5-6 (12'5)-100 x 4-5 ~Our data (from cultures) 200-480 x 3-4 15-25 x 4-7 5-6 40-70 x 4-6 ~Tubaki, 1958 20- 50 x 4-5 '5 15-20 x 6-6'5 20-22 x 6-9 (distal arm)Ingold, 1958 ? ? 15-50 x 4-6 (apex 3)Our data (from cultures)§ 50-125 x 4-5 10-22'5 x 5-7 '5 20-47'5 x 4-5

* Rounded off to the nearest 0'5 pm,t Width measured at the widest point unless stated ,:I: Figures in brackets indicate extreme values recorded infrequently.§ Figures compounded from more than one description.

E. Descals C.,J. Webster and B. S. Dyko 93

Table 2.

( -, Information lacking; +, fungus either described or pure-cultured.)

Country Author(s) Habitat Substrate Description Culture

Lemonniera aquatica

Australia (E.) Cowling, 1963 Streams FoamCowling & Waid, 1963 Subtropical Leaves

streams

Berlin Baumann & Poelt, 1970 Streams Dicot. leaves Partial

Canada (W.) Baerlocher & Kendrick, River Dicot. petioles1974

Canada (E.) Ingold,1960 Streams Scum, dicot. leaves

Czechoslovakia Marvanova Se Marvan, 1963 River Dicot. leavesMarvanova, 1968 Stream Dicot. leavesSladeckova, 1963 Stream Baits (wood, plastics)

England Ingold,1942 Stream Dicot. leaves + +Ingold, 1943(a)Ingold,1943(b) Streams Dicot. leavesIngold, 1944-5Ingold,1958 Streams ScumIqbal, 1972(a) River Dicot, leavesIqbal & Webster, 1973 Rivers Filtered waterJones, 1965 FoamJones & Oliver, 1964 Brackish water Beech wood baitNewton, 1971 River Dicot.Ieaf baitsWilloughby & Archer, 1973 Streams Dicot. leaf (wood)

baitsWilloughby, 1974 Streams Wood baits

France Durrieu, 1970 Streamde Wildeman, 1894 Pond (deep) Dicot. leaves Partialde Wildeman, 1895 Pond (deep) Dicot. leaves

German Dem. Rep. Arnold,1967 Streams FoamArnold,1968 Streams FoamArnold,1970(b) Spring Pseudotsuga twigs,

needlesCasper, 1965 Lakes (9 m deep) Dicot, leavesCasper, 1966 Streams Cyperaceae, dicot.

leavesGreenland (Denm.) Kobayasi et al. 1971 Soil with mosses Cellophane bait Partial

Hungary Banhegyiv retiz River Foam +Gonczol,1971 Stream Foam

Ireland (N.) Fenton, 1950

Ireland (S.) Ingold,1943(a) Stream Alnus leavesItaly Ciferri,1955 Aquatic Dicot. leaves +

Ciferri,1959 Various Leaves (Salicaceae)

Japan Tubaki, 1957 Aquatic Dicot. leaves + +Nimura& Suzuki, 1962Miura, 1967Miura, 1973 ScumMiura, 1974 Streams Scum

Norway Nilsson, 1964

Poland Nilsson, 1964

Romania Toth,1973 Streams Scum

Scotland Ingold, 1973(a) River FoamNilsson, 1964

South Africa Greathead,1961 Streams Dicot. leaves Partial

94 Taxonomic studies on aquatic hyphomycetes

Table 2 (cont.)


Lemonniera aquatica (cont.)

Spain Nilsson, 1960 Streams Scum

Sweden Nilsson, 1964 Stagnant, Scum, leaves +running waters

Willen, 1958, 1959 Brackish water VariousWillen, 1961 Lake Water sample

Switzerland Ingold,1949 Rivers Dicot, leaves

U.S.S.R. Dudka, 1974 Various Various PartialAstrakhan Mil'ko & Belyakov, 1968 River WaterCrimea Dudka & Logvinenko, 1968 StreamsEstonia Dudka,1968 Streams Dicot. leavesRussia Dudka & Florinskaya, 1971 Ponds Partial

Dudka, 1962 Streams PartialDudka, 1964Mil'ko, 1965 River Water sampleDudka, 1967 Radon springsArnold, 1970a

U.S.A.Calif. Ranzoni,1953 Dicot. debris. + +Fla. Conway, 1969 Streams Dicot. leaves PartialHawaii Anastasiou, 1964 Streams, ditches Foam, leavesIowa Dyko & Tiffany, 1972 River Dicot. leavesMass. Nilsson, 1964N.B. Nilsson, 1964N.Hamp. Nilsson, 1964N.J. Petersen, 1963N.Y. Conway, 1970 Streams Dicot. leaves

Petersen, "6']N. Carol. Petersen, 1963Penn. Petersen, 1963 (Same as for N.J.)S. Carol. Petersen, 1963Tenn. Petersen, 1963Va. Crane, 1968 Streams Foam, dicot. leaves +

Scott & Umphlett, 1963Umphlett, 1959Wolfe, 1971 Lake, streams Foam, dicot. leaves + +

W.Va. Crane, 1968 Streams Foam, dicot. leaves +Wise. Woelkerling & Baxter, 1968 Alkaline water Dicot. leavesWyo. Baxter, 1960 Streams Dieot. leaves

Wales Ingold, 1943(a)

Review paper Ingold,1973(b)Rabenhorst, 1907 PartialSaccardo & Sydow, 1899 PartialIngold, 1959(a)Ingold,1967Ingold,1959(b)

Lemonniera centrosphaera

Czechoslovakia Marvanova,1968 Stream Dicot. leaves + +Japan Miura, 1971 Stream Dicot, leaves + +

Miura, 1974 Temperate Scumstreams

Scotland Ingold, 1968 River Foam PartialIngold, 1973(a) River Foam

U.S.S.R. (Ukraine) Dudka,1973 +Dudka, 1974 +


Table 2 (cont.)


Lemonniera cornuta

England Iqbal,1972(a) Rivers Dicot. leaves + +Iqbal,197z(b) River Dicot, leaves + +Iqbal & Webster, 1973 Rivers Water (filtered)

Japan Miura, 1967 +Miura, 1973 ScumMiura, 1974 Temperate Scum

streamU.S.S.R. (Ukraine) Dudka,1973

Dudka,1974U.S.A.

Calif. Ranzoni,1953 River Dicot. leaves, twigs + +Petersen, 1963 Partial

W.Va. Crane, 1968 Stream Dicot. leaves, foam +Lemonniera filiformis

German Dem. Rep. Arnold,1967 Stream FoamArnold,1968 Stream Foam

U.S.S.R. (Russia) Arnold,1970(a) +Dudka, 1974 +

U.S.A. Petersen, 1963 Stream Dicot. leaves + +Lemonniera terrestris

Australia Tubaki, 1965 Streams Foam,scumCanada (E.) Ingold,1960 Streams Scum, dicot. leavesCuba Marvanova & Marvan, 1969 Streams Dicot. leavesCzechoslovakia Marvanova & Marvan, 1963 River Dicot. leavesEngland Ingold,1958 Streams Scum Partial +

Iqbal,1972(a) River Dicot. leavesIqbal & Webster, 1973 River Filtered waterJones, 1965 Foam

France de Wildeman, 1893 Pond Dicot. leavesWilloughby & Archer, 1973 Streams Leaf baits

German Dem, Rep. Arnold,1967 Streams FoamArnold,1968 Streams Foam

Hungary G5ncz51, 1971 (tentative) Stream FoamJapan Miura, 1967

Miura, 1973 ScumMiura, 1974 Temperate Scum

streamTubaki, 1958 Terrestrial Juglans capsule + +

South Africa Greathead. tqtir Streams Dicot. leavesSweden Nilsson, 1962 Streams Dicot. leaves Partial +

Nilsson, 1964 Terrestrial, Dicot. leavessemi-aquaticor aquatic

U.S.S.R.Ukraine Dudka,1974Russia Arnold,1970(a) Partial

U.S.A.N.B. Nilsson, 1964Wyo. Baxter, 1960 Streams Dicot. leaves

Reviewpaper Ingold,1959(a)Ingold,1967Ingold,1959(b)

Lemonniera pseudofloscu/a + +U.S.A.

North Carolina Streams Dicot, leaves + +Tennessee


taneously and radiating in a cruciate or tetrahedralpattern.

Type species L. aquatica de WildemanIn the following account references under

, Illustrations' followed by an asterisk containfigures showing conidial development.

LEMONNIERA AQUATICA de Wildeman, Ann. Soc.beige Microscop. 18: 135-161 (1894). (Figs. 1, 2.)

Pure culture: single-conidium isolate from sub-merged decaying leaf of Quercus in Devon,England.

Growth conditions. Two per cent malt extractagar, 20°, under diffuse light. Colony radialgrowth: 9 em in 50 days. Mycelium compact,superficial, initially white, turning brown withage; reverse brown, extracellular pigment absentor reddish brown. Submerged hyphae hyaline topale brown, branching irregularly, with conspicu-ous septa, 2'5-5 pm diam. Sclerotia: presence,time of appearance and morphology variable, withcultures frequently producing non-sclerotial sec-tors; scattered or in concentric rings, sometimesconfluent, superficial or partly submerged, c. 1mm diam, brownish to black. Mycelial cords inolder areas. Aerial mycelium scanty and near thecentre of the colony, low and white. Sporulationon agar not detected, but when pieces of agarcolony are submerged in unchanged distilled orstream water, with or without aeration, underdiffuse light, they produce abundant conidia.Conidiophore supporting cells usually dark, inter-calary and swollen, up to 20 pm diam, caespitoseon sclerotia or scattered; conidiophores hyaline,septate, erect, simple or more frequently withirregularly branched heads starting 50-150 pmabove the base; 75-150 x 3-6 pm at the base, thebranches arising at acute angles immediatelybelow septa; secondary branching may be present.Conidiogenous cells phialidic, hyaline, single or inpenicilli of up to eight, appearing successively,ampulliform, 17'5-27'5 x 6-6'5 pm at the widestpoint. Conidial primordia immediately branchingdirectly above the tip of the phialide into typicallyfour arms which diverge tetrahedrally, one armalways distal, and the rest growing slightly back-wards; all arms developing typically at the samerate or the distal branch slightly faster; septadistinct only after release. The basal septum maybe distinct at maturity. Conidia released bycleavage at the basal septum. Mature detachedconidia hyaline, with the central body indistinct orslightly globose, 5-6'5 pm diam ; attachmentdenticle sometimes visible; 4-(5) branches, cylin-drical or tapering very slightly towards the apex;40-105 x 4-4'5 pm slightly constricted at the base;

apex rounded; 2-3 septate. Germ-tubes andappressoria arising apically or laterally on any arm.

Habitat, substrates and distribution (table 2).Illustrations. Banhegyi, 1962 (fig. 4), Baumann

& Poelt, 1970 (figs. 6B, C), Casper, 1965* (fig. 8),Conway, 1969* (fig. 2F), Cowling, 1963 (fig. 3E),Dudka, 1962 (fig. 6), Dudka, 1967 (fig. 1), Dudka,1968 (fig. 5), Dudka, 1974* (fig. 100), Dudka &Florinskaya, 1971* (fig. 8), Durrieu, 1970 (fig. 7),Dyko & Tiffany, 1972* (fig. 2), Gonczol, 1971(table 1, fig. 20), Greathead, 1961* (plate 28, fig.1; fig. 13, a-j), Ingold, 1942* (text-figs. 1-4;pl. XII, fig. 1; plate XVI, fig. 3), Ingold, 1943(a)*(fig. 1.22), Ingold, 1944-1945 (fig. B, p. 44), In-gold, 1949* (fig. 1.1), Ingold, 1958* (pI. 20),Ingold, 1959(b)* (figs. 1, 11), Ingold, 1959(a)*(fig. 4), Ingold, 1967 (fig. 1), Ingold, 1973(b) (fig.10), Miura, 1974 (fig. 12), Nilsson, 1964* (fig.12i), Petersen, 1963* (figs. 1D, D, Rabenhorst,1907* (figs. 1,2, p. 541), Ranzoni, 1953* (fig. 10),Scott & Umphlett, 1963* (pl. II, fig. 3), Sladec-kova, 1963 (pl. I, fig. 8), Tubaki, 1957* (fig. 9),de Wildeman, 1894* (pI. v, figs. 1-22).

Holotype : not designated. Neorype : IMI 201899,Isoneotypes: University of Exeter Mycol Herb. no.3200. Other collections: IMI 48567a, 45711.

To our knowledge isolates of this fungus havebeen adequately described only from England,Japan, Sweden and the U.S.A. (Table 2). Innature, and especially when grown in pure culture,its conidiophores, phialides and conidia show greatvariations in size (Table 1). It is known to sporu-late on agar, although very sparsely (Petersen,1963). Colony colour can mutate readily (petersen,1963). The perfect state is not known, but Web-ster (1961) has detected presumed ascogonia inpure culture.

Detached conidia may be confused with thoseof other water-borne hyphomycetes w hen ob-served under low power, and their isolation istherefore recommended for proper identification.Conidia of what is believed to be L. aquatica havebeen detected in samples from 21 countries in allfive continents, but they have not been seen in theTropics. This relatively wide distribution ascompared to that of the other four species, couldbe partly explained by the adaptability of thefungus to various substrates as well as waterqualities (Table 2).

LEMONNIERA CENTROSPHAERA Marvanova, Trans.Br. Mycol. Soc. 51 : 613-616 (1968). (Figs. 1B, 3.)

Pure culture: single-conidium isolate from foamin Dundonnell, N.W. Scotland.

Growth conditions: as for L. aquatica (excepttemp. 16-17°). Colony radial growth 9 em diam in

E. Descals C.,]. Webster and B. S. Dyko 97

Scale C : 50JImScale A: 50JImII -

Scale B: 50JIm

Fig . 1. Conidial development in Lemonniera (from pure cultures). (A) L. aquatica (scale A). (B) L.centrosphaera (scale A). (C) L. comura (scale C) . (D) L. filiformis (scale A) (l M l 201901 ) . (E) L.terrestris (scale B).

4 MYC 69


Fig. 2. Lemonniera aquatica. Pure culture. (A-D) Conidium ontogeny; (A) conidiophore arising fromdark swollen cell; (E-H) mature, detached conidia; (E, F) germinating conidia.

LEMONNIERA CORNUTA Ranzoni, Farlowia 4 : 378-379 (1953)· (Figs. 1C, 4.)

Pure culture: single-conidium isolate from foamin Devon, England.

Growth conditions: as for L. aquatica. Colonyradial growth 8 em diam in 40 days. Myceliumcompact, superficial, initially hyaline, later reddishbrown, extracellular pigment brown, sectoringreadily. Submerged hyphae up to 4'5 pm diam,branching irregularly. Mycelial cords present.Sclerotia appearing in older cultures, usually inconcentric rings, dark brown to black, similar tothose of L. aquatica. Non-sclerotial sectors arefrequent. Aerial mycelium in the centre of thecolony or scattered, less than 1 mm high, white togreyish. Sporulation may occur on 2 % MEA(under NUV) but very sparsely; slices of agarcolony submerged for 24 h form abundantconidia. Conidiophore supporting cells intercalary,frequently dark-walled, swollen, scattered or onsclerotia. Conidiophores hyaline, erect, branchingirregularly or with an unbranched stipe, up to125-175 x 3-5 tun, usually septate; branches aris-ing at acute angles below the septa. Conidiogenouscells phialidic, hyaline, single or frequently inpenicilli of up to 3 per branch, ampulliform tocylindrical, 15-30 x 5-5'5 pm. Conidial primordiainitially spherical, later appearing as invertedtetrahedrons, the four arms arising simultaneouslyor in pairs from the equator; the distal branch israrely present; diverging at first laterally, thentwo opposite branches curve acropetally and thealternate pair basipetally ; septa indistinct beforerelease. Mature detached conidia hyaline, tetra-radiate, with an indistinct central body; armscurved of approximately equal length, 12'5-57 x3-4 pm at the base, tapering to 2'5-3 pm at therounded apex; basal constrictions very slight;up to several septa per arm. The attachmentdenticle is visible when conidia are observedlaterally. Germination by germ tubes and ap-pressoria arising from the apical cells of the arms.

Habitat, substrates and distribution (Table 2).Illustrations. Dudka, 1973 (fig. 14), Dudka,

E. Descals C., J. Webster and B. S. Dyko 99

51 days. Mycelium compact, superficial, reddish probably the U.S.S.R. Although Marvanovabrown, in concentric rings or scattered patches; (1968) reported conidial arm length up to 142 pm,reverse brown; extracellular pigments isabel- material studied by Miura (1971) and by us neverline brown; submerged hyphae l'5-3'5 pm diam, reached those dimensions, but there is no doubtwith conspicuous septa. Mycelial cords scattered that all these isolates are conspecific.and superficial, pale brown. Sclerotia scattered, Aerial sporulation has neither been detected onc. 2 mm diam, brown to black. Aerial mycelium Czapek-Dox agar (Miura, 1971), YGA (Miura,low, patchy, mainly in the centre of the colony, 1971) nor MEA, and the fungus has only beenwhite to greyish. Sporulation does not occur seen in water. This is probably its true habitat,directly on malt extract agar, but pieces of culture but more data will have to be recorded.on the same medium submerged in unchangeddistilled or tapwater for 2-4 days sporulateabundantly. Conidiophore supporting cells inter-calary, dark-walled and swollen, c. 17 pm diam.Conidiophores aquatic, hyaline, septate, erect,simple or more commonly irregularly branched atthe upper part, with branches arising at acuteangles immediately below a septum; 125-200 x3-4'5 pm diam, distinctly septate. Conidiogenouscells phialidic, hyaline, single or in penicilli of2-3, ampulliform to cylindrical, 14-25 x 4-4'5 pmat the widest point. Conidial primordia initiallyspherical, 3 pm diam; the first branch alwaysdistal, and immediately succeeded by the simul-taneous appearance of 3(-4) lateral arms aroundthe base of the primordium, diverging slightlybackwards; septa indistinct or one per branch;release by cleavage at the basal septum. Maturedetached conidia hyaline, with a very distinctspherical central body 6-10'5 pm diam, with theattachment denticle usually visible; (3)-4-(5)arms, the distal arm sometimes longer than theothers, 35-78 x 2-3 pm at the base, parallel-walled or more typically expanding to 3'5-4 pmat the rounded apex; basal constrictions absent;1-3 septa per arm, distinct after release. Germina-tion by germ tubes and appressoria arising fromthe apical cells of the arms.

Habitat, substrates and distribution (Table 2).Illustrations. Dudka, 1973 (fig. 13), Dudka,

1974* (fig. 97), Ingold, 1968 (pl. VIII, fig. D; text-fig. 2), Marvanova, 1968* (fig. 1), Miura, 1971(fig. 2), Miura, 1974 (fig. 50).

Holotype: CCM No. F 149 in BRNU (n.v.). Othercollections: Miura No. 1028 (Japan); University ofExeter MycoI. Herb. No. 3201 (authenticated).

Type material was not available but a permanentmount with seven conidia from Sweden, sent andauthenticated by Marvanova, was examined. Thecentral body of the conidia measured 6'25-8"75pm diam, and the four arms were 72-102 x 2'5-5pm at the base, slightly tapering towards the apex.The apiculus was visible. These data are withinthe limits of the original description.

L. centrosphaera has only been isolated fromCentral Europe, the British Isles, Japan and


Fig. 3. Lemonniera centrosphaera. Pure culture. (A-F) Conidium ontogeny; (E) conidiophore arisingfrom dark cell; (G-F) mature, detached conidia. (Arrows point at detachment pegs.)

V>o1::3

1


Fig. 4. Lemonniera cornuta, Pure culture. (A- H) Conidium ontogeny j (A, C) primordia developingatypically ; (F, H) conidiophore supporting cells, dark and swollen. A-D drawn at larger scale.

LEMONNIERA TERRESTRIS Tubaki, J. Hatt. Bot.Club 20 : 165-166 (1958). (Figs. 1E, 6.)

L. brachycladia Ingold, Trans. Br. mycol. Soc.41, 365-372 (1958).Pure culture : single-conidium isolate from sub-merged decaying leaf in Devon, England.

Growth conditions : as for L. aquatica. Colonyradial growth approximately 9 em diam in 80days. Mycelium very compact, superficial, initiallywhite, later pale cream, with frequent pale-brownto chestnut concentric zonations; hyaline margin;

Neotype: IMI 208222. Isoneotype: University ofExeter Mycol. Herb. No. 3237. Other collections:IMI 2019°1 and Exeter MycoJ. Herb. No. 3203.

This taxon was first described by Petersen(1963) from New York, U.S.A. No type wasdesignated at that time and no original materialcould be found which would be suitable for typi-fication. Records from other countries appeardoubtful. Conid ia of what probably is the samefungus were twice isolated by us from the BritishIsles and by Wolfe (1971) in the U.S.A.

L. filijormis can be easily distinguished by thefine attachments of only the lateral arms to theindistinct central body. This characteristic ispresent through the entire conidial development.Both the lateral and the distal arms of conidia ofL. aquatica, L. terrestris and L. com uta, on theother hand, are always broad-based, although aslight swelling near the base can be frequentlyobserved. The term basal isthmus should belimited to the type of arm constriction charac-terizing conidia of L. filiformis and L. pseudo-floscula. It is also typical of Dendrospora Ingold,Triscelophorus Ingold and Tricladium Ingold,among others .

Lemonniera filiformis R. H . Petersen exDyko (Figs. 1D, 5.)

Lemonniera filijormis R. H. Petersen (as 'fili-forme'), Mycologia ss: 574-576 (1963), notvalidly published.

Pure culture :single-conidiumisolate from Virginia,U.S.A.

Growth conditions as for L. aquatica. Colonyradial growth 9 em diam in 51 days. Myceliumcompact, superficial, first white, later pink to darkchestnut; reverse hyaline to brown ; extracellularpigments dark brown. Submerged hyphae hyaline,1'5-6 pm diam, branching irregularly. Sclerotia inconcentric rings, confluent in the central region,appressed to rounded, up to 3 mm diam, super-ficial or erumpent, dark brown to black. Aerialmycelium scanty, visible over the sclerotial masses,white to greyish, low. Sporulation on agar notdetected, but abundant and typical when pieces ofagar culture are submerged in water. Conidio-phore supporting cellsusually dark-walled, swollen,c. 15 p,m diam, in chains, scattered or on sclerotia.Conidiophores hyaline, septate, erect, simple orwith an irregularly branched head, with branchesarising at acute angles immediately below thesepta, 62-300 x 3'5-4 pm; septa distinct andwithout constrictions. Conidiogenous cells phiali-die, hyaline, ampulliform to cylindrical, single orin penicilli of 2-3 per branch, appearing in succes-sion, 15-17"5 x 5-6 pm at the widest point.Conidial primordia initially spherical, with thedistal arm always developing first, but soon threelateral branches bud out from near the base and


1974* (fig. 96), Iqbal, 1972(a)* (fig. 3), Iqbal, radiate slightly backwards and at the same rate;1972(b) (fig. 34), Miura, 1967* (fig. 1, O-P), septation ind istinct initially ; release by cleavageMiura, 1974 (fig. 51), Petersen, 1963* (fig. 1C), at the basal septum. Mature detached conidiaRanzoni, 1953* (figs. 1S, 14). hyaline, with an indistinct main body which

Holotype:notdesignated. Neotype :IMI 201900 (au- merges with the distal arm; beaklike attachmentthenticated), Isoneotypes : Universityof ExeterMycoJ. denticle sometimes visible; typically tetrahedral,Herb. NO.3202. Other collections: deposited by Iqbal with the distal arm (12'5)-100 x 4-5 pm at the(1972b) at CBS and 1M!. widest point; lateral arms slightly smaller, basal

No holotype was designated, and lectotypes isthmus very fine ; all arms tapering at both ends;could not be selected, but conidia of this fungus apex rounded; septa distinct, up to six per arm.are very characteristic and isolates from Japan and and sometimes slightly constricted. Germinationthe British Isles match Ranzoni 's illustrated des- by germ-tubes arising from the apical cells on thecriptions from material in North America. arms.

Little is known about this apparently aquatic Habitat, substrates and distribution (see Tablefungus. It is interesting, though, that the same 2).general arm arrangement has been depicted for an Illustrations. Arnold, 1967 (table 1, fig. 4),undescribed propagule (Arnaud, 1952, p. 211; Arnold, 1970(a) (fig. 2.4), Dudka, 1974 (fig. 98),Miura, 1974), and it is also known to occur in Petersen, 1963* (figs. 3a-e).ascospores of the marine fungus Antennosporaquadricornuta, as well as in propagules of variousmembers of the Chlorococcales (Nilsson, 1964,P·46).

E. Descals C.,J. Webster andB. S. Dyko

/7,

- ;'" ' ..<

~

'..,?.;G

81::3

1°3

Fig. 5. Lemonniera filiformis. Pure culture. (A-H) Conidium ontogeny; (B) conidiophore supportingcells dark and swollen; (I-L) mature, detached conidia; (I) atypical, 5-armed conidium.

1°4 Taxonomic studies onaquatic hyphomycetes

Fig. 6. Lemonniera terrestris. Pure culture. (A-I) Conidium ontogeny; (B) drawn at larger scale; (D-H)conidiophore supporting cells, dark and swollen; (J-P) mature, detached conidia; (M, 0, P) atypicalforms.

E. Descals C.,J. Webster and B. S. Dyko 1°5

Fig. 7. Lemonniera pseudofloscula. Pure culture. (A-C) Conidium ontogeny ; (D-E) mature, detachedconidia; (F) dark cells supporting conidiophores ; (G) photomicrographs showing conidium ontogeny atdifferent stages.

Growth conditions at 10° on malt yeast soytoneagar: colony radial growth 5-5 '5 em diam in 18days; mycelium compressed, superficial, firsthyaline, later tan brown; colony reverse same;extra-cellular pigments tan brown. Hyphae 1-6 pmdiam, branched, sometimes in ropes. Sclerotia inconcentric rings, confluent in the central region,flattened, rounded, up to 3 mm diam, superficial orerumpent, black. Aerial mycelium scant, visibleover sclerotia and colony surface, hyaline, low.Sporulation on agar not detected, but moderateand typical on MYS and YpSs agars when stripsof culture are submerged in distilled water andaerated. Conidiophore: hyaline, septate, erect,simple or often with an irregularly branched head,with branches arising immediately below the septa,200-480 x 3-4/tm . Conidiogenous cells phialidic,hyaline, ampulliform to cylindrical, single or inpenicilli of up to 3 per branch, 15-30 x 4-7 pm.Conidial primordia initially spherical to ovate be-coming tetrahedral, with distal arm developingmore rapidly than laterals; 3-4 lateral arms bud-ding from the central body and quickly becomingof the same length as the distal arm; constrictionsappearing early at the base of all arms; septa

Lemonniera pseudofloscula Dyko, sp.nov,(Fig. 7.)

Fungi Imperfecti, Hyphomycetes. Aquaticus sub-mersus, mycelio septato, ramoso, repente, hyalino.Conidiophora hyalina, septata, ramosa, 200-480 pmlongitudine, 3-4 pm latitudine, ad apices 1 vel 3phialides gerentia, Phialides a forma cylindrica usquead ampullaceam, 15-30 pm longitudine, 4-7 pm lati-tudine maxima. Phialosporae hyalinae cum partecentraliglobosa, 4 vel7 pm diamcum phialideconnexa,cum uno axe et 3 vel 4 ramis, divergentibus. Rami etaxis constricti ad basim, 34-70 psn, longitudine,4-7 pm latitudine maxima, 2 vel 5 septati,

Habitat : in foliis putrescentibus in rivo CarolinaeSeptentrionalis in montibus Appalachianis, U.S.A.Holotypus: IMI 208455, Tom Henry Branch, High-lands, N.C., U.S.A., 1 Jan. 76, B. J. Dyko. Isotypi:N.Y., TENN, No. 40056; Univ. of Exeter Mycol.Herb No. 3225 .

Holotype: Nagao Institute, Kitashinagawa, Tokyo(Japan) (n.v.) , Other collections ; University of ExeterMycol. Herb. no. 3204, IMI 70810.

Cultures and permanent mounts deposited atthe latter institution have been recently examinedby Ingold.

In L. terrestris conidial dimensions frequentlyoverlap those of L. aquatica, but in the former theprotoplasm is pearlier (probably due to glycogen),the distal arm can be slightly longer than the rest(Nilsson, 1962; Tubaki, 1958) and, most impor-tant of all, conidia appear as sticky masses on theagar colony, especially in older cultures (Tubaki,1958). Sporulation cannot be considered as typi-cally aerial because the sticky conidia are probablyproduced only in very humid air. They are cer-tainly small enough to develop within minute


reverse brown. Extracellular pigments brown. condensation droplets. Thirdly, their efficientSubmerged hyphae 3'5 pm diam, branching dispersal would require submersion in runningirregularly. Sclerotia in concentric rings, confluent, water.similar to those of L. aquatica. Aerial mycelium This fungus has been isolated from Japan, theabundant in the centre, dark; elsewhere less than British Isles and Sweden although records from0'5 mm high. Conidiophore supporting cells other countries seem to indicate a similar distribu-intercalary, hyaline or swollen and brown; tion to that of L. aquatica, with which it appearsscattered or on sclerotia, up to 35 pm long. frequently associated on the same substrate andConidiophores hyaline, septate, erect ; protoplasm habitat. There is one record from the tropicsslightly vacuolate; simple or with a penicillate (Marvanova & Marvan, 1969). Conidia illus-head with branches arising irregularly at acute trated by Arnold (1967,1968, 1970a) are probablyangles below the septa, 50-125 x 4-5 pm. Coni- those of another organism .diogenous cells phialidic, hyaline, single or inpenicilli of up to five, ampulliform, cylindrical orlageniform, 10-22'5 x 5-7'5 pm at the widestpoint. Conidial primordia initially spherical, butsoon branching out into typically four arms whichdiverge tetrahedrally, i.e. one branch alwaysdistal, the others radiating slightly backwards andat the same rate; basal septa very distinct; othersepta on the arms may appear after release. Maturedetached conidia hyaline; protoplasm pearly;central body more or less distinct, 4-7 pm diam;branches (3)- 4- (5), of equal size, (7)- 35-(67) x(4)-6-(9) pm at the base tapering more or lessdistinctly towards the rounded apex; broad-basedor with a very slight constriction, 0-3 septate.Germ-tubes and appressoria developing vigor-ously from the apical cells of the arms.

Habitat, substrates and distribution (Table 2).Illustrations. Arnold, 1967 (table 1, fig. 6),

Arnold, 1968 (fig. II, 5), Arnold, 1970(a)* (fig.1.2), Dudka, 1974* (fig. 99), Gonczol, 1971 (table1; figs. 18, 19; pl. 3, fig. 4), Greathead, 1961*(pI. 28, fig. D, Ingold, 1958* (text-fig . 1, pl. 20),Ingold, 1959 (b)* (fig. 1), Ingold, 1959(a)* (fig. 4),Ingold, 1967 (fig. 1), Miura, 1974 (fig. 49), Nils-son, 1962* (fig. 12), Nilsson, 1964* (fig. 12F),Tubaki, 1958* (fig. 21).

REFERENCES

ANASTASIOU, C. J. (1964). Some aquatic fungi imper-fecti from Hawaii. Pacific S cience 18, 202-206.

ARNAUD, G. (1952). Mycologie concrete. Genera 1.Bulletin trimestriel de la Societe mycologique de laFrance 68, 181-223.

ARNOLD, G. R W. (1967). Sporen aquatischer Hypho-myzeten aus dem 'Paradies' bei Weimar. W est-faellische Pilzbr. 6, 156-159.

ARNOLD, G. R W. (1968). Sporen aquatischer Hypho-myzeten aus Gewaessern im Bezirk Halle. Wissen-schaftliche Zeitschrift der Friedrich-Schiller Unioer-sitaet fena (Mathematik-Naturwissenschaft Reihe) 17,369-373.

ARNOLD, G. R W. (1970a). Hyphomycetes aquatici exaquariis regionis Leningradensis isolat. NovostiSistematik niz Kast Akademiya Nauk. SSSRBotanicnyi Institut 6, 176-189.

ARNOLD, G. R W. (1970b). Aquatische Hyphomyzetenauf Koniferen. Limnologica 7, 381-382.

BAERLOCHER, F. & KENDRICK, W. B. (1974). Dynamicsof the fungal population on leaves in a stream.Journal of Ecology 62, 761-791.

BANDONI, R J. (1972). Terrestrial occurrence of someaquatic hyphomycetes. Canadian Journal of BotanyS0, 2283-2288.


initially indistinct; conidial release by cleavage at BANHEGYI, J. (1962). Danubialia hungarica 18. AnnaIesthe basal septum. Mature detached conidia hyaline, Universitatis scientarum budapestiensis de Rolandotetrahedral, with distinct subspherical central Eotoos nominatae. Budapest (Sectio biologica).

BAUMANN, L. H. & POELT, J. (1970). Wasserbe-body 4-7 pm diam, a truncate scar of attachment wohnende Hyphomyzeten im Raum von Berlin.sometimes present, with arms 34-70 x 4-7 pm with Verhandlungen der B otanischen Vereins der Provinzthe distal arm often slightly longer than the laterals; Brandenburg 107, 65-79.basal isthmus 1-5 pm diam; arm apices tapered, BAXTER, J. W. (1960). Aquatic hyphomycetes fromrounded; septa distinct, up to 7 per arm. Germina- Wyoming. Mycolog ia 52, 654-655.tion by germ-tubes arising from the apical cells on CASPER, S. J. (1965). Hyphomyzeten-Studien. I. Diethe arms. Suesswasser Hyphomyzeten des Stechlinsee-

This taxon is very similar to L. filiformis but an Gebietes. Limnologica 3, 257-270.obvious difference is the presence of a basal isthmus CASPER, S. J. (1966). Hyphomyzeten-Studien. II.on the distal arm. This isthmus, with those of the Suesswasser-Hyphomyzeten aus dem Thueringer

Wald, dem Erzgebirge und dem Riesengebirge.lateral arms, defines a central, subspherical body. Limnologica 4, 471-481.The isthmi tend to be longer and the arms shorter ClFERRI, R. (1955). Nuovi reperti italiani di funghi deland slightly wider in this taxon than in those found suolo ed idrofili. Allionia 2, (2), xxi-xxiv (appendix).in L. filiformis . A central body is also found in CIFERRI, R. (1959). Osservazioni ecologiche su ifomi-conidia of L. centrosphaera but it is slightly larger ceti aquatici. Omagiu lui Traian Savulescu cu prilejuiand its narrower arms are not constricted at their implinirii a 70 de ani, pp. 157-162. Academiabases. Republicii populare Romine. Editura Academici

Eight isolates of this fungus were made from the Republicii Populare Romine.same collection of material. All isolates required CONWAY, K. E. (1969)· Some aquatic hyphomycetes of

Florida. Quart erly Journal of the Florida A cademy ofthe use of YpSs agar for good sporulation. This is S ciences 32, 210-20.quite unlike the other isolates of Lemonniera, CONWAY, K. E. (1970). The aquatic hyphomycetes ofwhich sporulate readily on MEA 2 %. Central New York. My cologia 62, 516-530.

We are very grateful to Professor C. T. Ingold COWLING, S. W. (1963). The aquatic hyphomycetes ofEastern Australia. B.Sc. dissertation, Botany De-

for examining cultures and herbarium material, to partment, University of New England, ArmidaleMr T. Learmouth, of the University of Exeter (New South Wales), Australia. 93 pp. +7 figs.Library, for Ukrainian and Russian translations, COWLING, S. W. & WAID, J. S. (1963). Aquatic hypho-to Mr R. A. Davey for competent technical assist- mycetes in Australia. Australian Journal of Scienceance, and to Professor F. W. Clayton, University 26, 122-123.of Exeter, for the Latin translation. CRANE, J. L. (1968). Freshwater hyphomycetes of the

Northern Appalachian highland including NewEngland and three Coastal Plain States. AmericanJ ournal of Botany 55, 996-1002.

DUDKA, I. A. (1962). [Aquatic hyphornycete speciesnew for the Ukrainian SSR.] Ukrainbskyi BotanicnyiZhurnaI19,66-71.

DUDKA, I. A. (1964). [On the ecology and seasonaldynamics of aquatic hyphomycetes of the Southernpart of the Kiev forest area.]Journal of Botany of theA cademy of Sciences of the Ukraine 21, 50-57.

DUDKA, I. A. (1967). [Aquatic hyphomycetes of radonsprings in forest-steppe on the right bank of theDnieper in the Ukrainian SSR] Mikologija iFitopatologija 1,441-446.

DUDKA, I. A. (1968). [Aquatic hyphomycetes of north-eastern Estonia.] Aesti NSV Tead Akademiya Toim .Biological Series 17, 394-403.

DUDKA, I. A. (1973). [Aquatic hyphomycetes of theUkraine. 1. New species for the mycoflora of theSoviet Union.] Ukra inskii Zhurnal30, 449-456.

DUDKA, I. A. (1974). Ukrainian aquatic hyphomycetes.Academy of Sciences of the Ukrainian RS.RM.G.Holodny Botanical Institute. Publications ' NaukovaDumka ', Kiev.

DUDKA, I. A. & FLORINSKAYA, A. A. (1971). [New andrare species of aquatic fungi in fish ponds in theLeningrad district .] Mikologi]« i Fitopatologija 5,431-438.

108 Taxonomic studies onaquatic hypJwmycetes

DUDKA, I. A. & LOGVINENKO, L. I. (1968). [The IQBAL, S. H. (1972b). Some observations on aquaticmycoflora of mountain rivers and streams of the hyphomycetes. Ph.D. Thesis, University of Exeter,Crimean State reservation.] Mikologija i Fitopatolo- England.gija 2, 269-274. IQBAL, S. H . & WEBSTER, J. (1973). Aquatic hypho-

DURRIEU, G. (1970). Hyphomycetes aquatiques de la mycete spora of the River Exe and its tributaries.region Toulousaine et des Pyrenees. Bulletin de la Transactions of the British Mycological Society 61,Societe d' Histoire Naturelle de Toulouse 106, 331-346.112-116. JONES, E . B. G. (1965). Some aquatic hyphomycetes

DYKo, B. J. & TIFFANY, L. H. (1972). A preliminary collected in Yorkshire. The Naturalist, Leeds 893,study of aquatic hyphomycetes in Iowa. Proceedings 57-60.of the Iowa Academy of Sciences78, 72-73. JONES, E. B. G. & OLIVER, A. C. (1964). Occurrence of

FENTON, A. F . (1950). Aquatic hyphomycetes of aquatic hyphomycetes on wood submerged in freshNorthern Ireland. Irish Naturalist's Journal 10, and brackish water. Transactions of the British22-23· Mycowgical Society 47, 45-48.

GONCZOL, J. (1971). Aquatic hyphomycetes from the KOBAYASI, Y., HIRATSUKA, N., OTANI, Y., TUBAKI, K,Mountains Boerzsoeny. Annals of Natural History of UDAGAWA, S., SUGIYAMA, J. & KONNO, K. (1971).the National Museum of Hungary 63, 57-75. Mycological studies of the Angmagssalik Region of

GREATHEAD, S. K . (1961). Some aquatic hyphomycetes Greenland. Bulletin of the National Science Museumin South Africa . Journal of South African Botany 27, 14, 1-96.195-228. MARVANovA, L. (1968). Lemonniera centrosphaera

INGOLD, C. T . (1942). Aquatic hyphomycetes of sp.nov. Transactions of the British Mycologicaldecaying alder leaves. Transactions of the British Society 51, 613-616.Mycological Society 25, 339-417. MARVANovA, L. & MARVAN, P. (1963). Einige Hypho-

INGOLD, C. T . (1943a). On the distribution of aquatic myzeten aus den F1iessenden Gewassern des Hrubyhyphomycetes saprophytic on submerged decaying [esenik, Acta Musei Silesiae. Opaoa (C.S.S.R.)leaves. New Phytologist 42, 139-142. (ser.A) 12, 101-108.

INGOLD, C. T. (1943b). Further observations on MARVANovA, L. & MARVAN, P. (1969). Aquaticaquatic hyphornycetes of decaying leaves. Transac- hyphomycetes in Cuba. Geskd Mykologie23, 135-140.tions of the British Mycological Society 26, MIL'KO,O. O. (1965). [Fungi isolated from water of the104-115. Soviet part of the Danube.] Mykrobiolohichniyi

INGOLD, C. T. (1944-5). The tetraradiate spores of Zhurnal zq, 38-49.certain aquatic hyphomycetes and the propagules in MIL'KO, O. O. & BELYAKOV, L. A. (1968). [Speciessome species of Sphacelaria. Proceedings of the composition of fungi of the River Volga .] Mikrobio-Linnean Society of London 157, 43-45. logiya 37, 944-956. (Translation by Institute of

INGOLD, C. T . (1949). Aquatic hyphomycetes from Microbiology, Academy of Sciences of the USSR,Switzerland. Transactions of the British Mycological Moscow. UDC 582.28 :577.472, pp. 795-797.)Society 32, 341-345. MIURA, K (1967). An annotated list of aquatic

INGOLD, C. T. (1958). New aquatic hyphomycetes: hyphomycetes of Sugadaira, Japan. Transactions ofLemonniera brachycladia, Anguillospora crassa and the Mycological Society of Japan 8, 23-27.Fluminispora oualis. Transactions of the British MIURA, K (1971). Notes on filamentous fungi fromMycological Society 41, 365-372. Japan. Nos. 5-6. Journal of Japanese Botany 46,

INGOLD, C. T . (1959a). Fungi. In Vistas in botany (ed. 193-198.W. B. Turrill), pp. 348-386. London: Pergamon MIURA, K (1973). Stream spora, Proceedings of thePress. Japanese Society of Plant Taxonomists 3, 1-4·

INGOLD, C. T . (1959b). Submerged aquatic hypho- MIURA, K (1974). Stream spora of Japan. Transactionsmycetes. Journal of the Queckett Microscopical Club, of the Mycological Society of Japan 15, 289-308.ser, 4, 5, 115-130. NEWTON, J. (1971). A mycological study of decay in the

INGOLD, C. T . (1960). Aquatic hyphomycetes from leaves of deciduous trees on the bed of a river. Ph.D.Canada. Canadian Journal of Botany 38,803-806. thesis, University of Salford, England.

INGOLD, C. T . (1967). Spores from foam. Bulletin of NILSSON, S. (1960). Aquatic hyphomycetes fromthe British Mycological Society 1, 60-63. Northern Spain. Svensk botanisk Tidskrift 54, 530-

INGOLD, C. T. (1968). More spores from rivers and 532.streams. Transactions of the British Mycological NILSSON, S. (1962). Second note on Swedish freshwaterSociety 51, 138-143. hyphomycetes. Botaniska Notiser 115, 73-86.

INGOLD, C. T. (1973a). Aquatic hyphomycete spores NILSSON, S. (1964). Freshwater hyphomycetes. Taxo-from West Scotland. Transactions of the British nomy, morphology and ecology. Symbolae botanicaeMycological Society 61, 251-255 . upsalienses 18, 1-130.

INGOLD, C. T . (1973b). Branched aquatic propagules NIMURA, H . & SUZUKI, S. (1962). A list of the aquaticwith four or five divergent arms. Acta Botanica hyphomycetes in Japan. Journal of Japanese BotanyIndica 1, 69-72. 37, 3°-32.

IQBAL, S. H . (1972 a). New aquatic hyphomycetes. PARK, D. (1972). On the ecology of heterotrophicTransactions of the British Mycological Society 59, microorganisms in freshwater. Transactions of the301-307. British Mycological Society 58, 291-299 .

E. Descals C.,J. Webster and B. S. Dyko 1°9PETERSEN, R. H. (1963). Aquatic hyphomycetes from

North America. III. Phialosporae and miscellaneousspecies. Mycologia 55, 570-581.

RABENHORST, G. L. (1907). Kryptogamenflora vonDeutschland, Oesterreich und der Schweiz, Bd. 1 (8),pp. 537-546. Leipzig, Kummer, 1884-1907.

RANZONI, F. V. (1953). The aquatic hyphomycetesof California. Farlowia 4, 353-398.

SACCARDO, P. A. & SYDOW, P. (1899). Sylloge Fungorumomnium hucusque cognitorum, vol. XIV, pp. 1067-1068.

SCOTT, W. W. & UMPHLETT, C. J. (1963). Some newand unusual fungi from Virginia II. Aquatic hypho-mycetes, Virginia Journal of Science 14,47-64.

SLADECKOVA, A. (1962). Lymnological study of thereservoir Sledice near Zeliv. XX. Periphytonstratification during the second year-long period(August 1958-June 1969). Sbornik oysoke skolychemicko-technologiske v. Prase. Praha 6, 221-291.

SLADECKOVA, A. (1963). Aquatic deuteromycetes asindicators of starch campaign pollution. Institut Rev.ges. Hydrobiologie 48, 35-42.

TOTH, S. (1973). Aquatic hyphomycetes from Transyl-vania. Botanikai Koezlemenyek 60, 153-156.

TUBAKI, K. (1957). Studies on the Japanese hypho-mycetes, III. Aquatic group. Bulletin of the NationalScience Museum Tokyo 41, 249-268.

TUBAKI, K. (1958). Studies on the Japanese hypho-mycetes. V. Leaf and stem group with a discussionof the classification of hyphomycetes and theirperfect stages. Journal of the Hattori Botanical Club20, 142-244.

TuBAKI, K. (1960). On the Japanese aquatic hypho-mycetes. Scum and foam group, referring to thepreliminary survey of the snow group. Nagaoa 7,15-29.

TUBAKI, K. (1965). Contribution towards the fungusflora of Australia and New Zealand. Annual Report ofthe Institute of Fermentation, Osaka 2, 39-62.

UMPHLETT, C. J. (1957). Preliminary survey of aquatichyphomycetes on angiosperm debris. VirginiaJournalof Science (N.S.) 8, 291.

UMPHLETT, C. J. (1959). Heliscus tentaculus, n.sp., anaquatic hyphomycete. Virginia Journal of Science 10,

27-32.WEBSTER, J. (1961). The Mollisia perfect state of

Anguillospora crassa Ingold. Transactions of theBritish Mycological Society 44, 559-564.

WILDEMAN, E. DE (1893). Notes mycologiques. Fascicle2. Annales de la Societe belge de Microscopie Bruxelles17, 35-68.

WILDEMAN, E. DE (1894). Notes mycologiques. Fascicle3. Annales de la Societe belge de Microscopie, Bruxelles18, 135-161.

WILDEMAN, E. DE (1895). Notes mycologiques, Fascicle6. Annales de la Societe beige de Microscopie, Bruxelles19, 193-206.

WILLEN, T. (1958). Conidia of aquatic hyphomycetesamongst plankton algae. Botaniske Notiser 111,

431-35.WILLEN, T. (1959). The phytoplankton of Gorvaln, a

bay of Lake Malaren, Oikos 10, 241-274.WILLEN, T. (1961). The phytoplankton of Osbysion,

Djursholm. I. Seasonal and vertical distribution ofspecies. Oikos 12, 36-69.

WILLOUGHBY, L. G. (1974). In Biology of plant litterdecomposition, vol. II (ed. C. H. Dickinson andG. F. J. Pugh), pp. 659-682. London, N.Y.:Academic Press.

WILLOUGHBY, L. G. & ARCHER, J. F. (1973). The fungalspora of a freshwater stream and its colonizationpattern on wood. Freshwater Biology 3, 219-239.

WOELKERLING, W. J. & BAXTER, J. W. (1968). Aquatichyphomycetes of Wisconsin: distribution andecology. Mycopathologia et Mycologia Applicate 35,33-36.

WOLFE, C. C. (1971). Taxonomic considerations in thefreshwater hyphomycetes. Ph.D. thesis, VirginiaPolytechnic Institute and State University, 120 pp.

(Accepted for publication 12 March 1976)

Documents

Taxonomic studies on aquatic hyphomycetes: I. Lemonniera de Wildeman