[R ead 12th N ovem ber, 1936; issued separately 19th -July ... · The present writer gives an...

206 K ath leen M-: C rooks:

A r t . X IV .— S tu d ies on A ustra lian A qua tic Phycom ycetes.

By K A T H L E E N M . C R O O K S.

[R ea d 12th N o v em b er , 1936 ; issu ed se p a ra te ly 19th -July, 1937.]

I n t r o d u c t io n .

In recent years, investigations have been carried out in d ifferent countries, notably G reat B rita in , A m erica, G erm any, and D enm ark, to ascertain the natu re of the aquatic fungus flora. T he ra rer m embers, B lastocladia, Saprom yces, and R hipid ium w ere not recorded until late in the n ineteenth century,' w hen they w ere described by C ornu (1871), Reinsch (1878 ), and T h ax te r (1894-1896), although investigations in the Saprolegniaceae had commenced much earlier. E arly in the tw entieth century accounts of aquatic fungi by V on M inden (1912) in Germ any, and Petersen (1910) in D enm ark were published, but only occasional records appeared from then until 1922, since when, this, group, particu larly the ra re r fam ilies, B lastocladiaceae and Lepto- m itaceae, has d raw n the attention of a num ber of w orkers. Recently, L und (1934) has contribu ted to the records of the aquatic fungi in D enm ark, and A pinis (1929) has investigated the Saprolegniaceae in Latvia. Coker (1922-1926) and his w orkers thoroughly studied the group in A m erica, followed by o ther papers— K anouse (1925, 1927, 1 9 3 2 ) ,-Couch (1926, 1927, 1932), H arvey (1930), and S parrow (1932, 1936). In G reat B rita in , the earliest records of aquatic fungi w ere by M assee (1891) and Ram sbottom (1914-1916), bu t since 1932 other mycologists have contributed to th is group, nam ely Barnes and M elville (1932), Cook and Forbes (1932), Forbes (1935), and ju s t before the com pletion of th is, paper Sparrow (1936) published the results of an extensive investigation. H itherto , in A ustra lia , the attention of w orkers has. been confined to the terrestria l fungus flora, w ith the exception of records by Rodw ay (1897), H ard y (1 9 1 0 ) ,. and Johnston (1917, 1921) ofSaprolegnia fe ra x associated w ith a disease in fish.

T he present w riter gives an account o f some m em bers o f the B lastocladiales and Saprolegniales and com pares them w ith those recorded in o ther -countries. N one of the fung i described in th is paper, w ith the exception of Sapro legnia fe ra x , has p re viously been described from A ustra lia.

T he w rite r was particu larly in terested in the ra re r B lastocladiales and Leptom itaceae, and, w hen collecting, efforts were m ade to secure these form s ra th e r th an the filam entous fungi—

[ P &�� S �� 4 �� z , 49 ( N . S . ) , P t . I I . , 1937.]

Saprolegniaceae or Pythiaoeae, although the la tte r w ere also obtained. T he fungi have been collected chiefly from artificial ponds, and, only in th ree instances, have they been collected from natu ra l w aters— Y arra R iver a t R udder Grange, B arham R iver a t Apollo Bay, and M cCallum ’s Creek near M aryborough.

S tu d ies on A ustra lian A quatic P hycom ycetes. 207

M e th o d s o f C o l le c t io n .

T he m ethods used w ere those employed by o ther w orkers on this group— i.e., the B lastocladiales and Leptom itaceae— the setting of traps containing suitable baits-in a likely locality.. T he baits used w ere tw igs, rose hips, apples, oranges, bananas, grapes, prunes, Cotoneaster berries, Crataegus berries, S o lanum pseudo-, capsicum 'berries, and Japonica fru its. They w ere pu t into fine w ire-m esh baskets, which w ere placed in the pond or stream , anchored to the shore by a long w ire, and subm erged fo r one or two m onths. W hen the baits were brought into the laboratory, they were thoroughly .washed in runn ing w ater fo r about a day, and then transferred to sterile distilled w ater, in glass dishes w ith loosely fitting covers, so as not to exclude the air. E xam ination of the baits was commenced immediately. In m ost cases, the pustules form ed by Blastocl.adia, G'onapodya, Saprom yces, and R h ip id iu m w ere visible when brought in and som etimes the fru its w ere covered thickly w ith these whitish patches. In fection of the f r u i ts 'a n d fu r th e r developm ent of the plants, e.g., the form ation of resting spores, continued a f te r they were placed in sterile water.

T he filam entous fungi, e.g., Saprolegniaceae, which usually • occurred on tw igs,, but also on some fru its , rose hips, and Crataegus berries, did not appear fo r a few days a f te r the baits were brought in. T hey w ere then tran sfe rred to dishes of sterile w ater containing boiled halved hem p seeds, on w hich they grow readily. T he descriptions of the fo rm s included in this paper are from the grow th on hem p seeds.

In a couple of instances, pond w ater was brought in to the laboratory, and im m ediately baited w ith boiled hem p seeds. A fte r a few days, the filamentous form s, e.g., D ictyuchus and A chlya , appeared. Pond w ater was also baited w ith grapes and prunes, when P yth iom orpha was obtained. T h is genus also appeared on rose hips a f te r they had been in the laboratory fo r some days. F o r Blastocladia, Saprom yces, R h ip id ium , and Gonapodya, the satisfactory baits are apples, rose hips, Crataegus berries, and Japonica fru its. O ranges w ere used as baits on two occasions but only P yth iom orpha was obtained.

O w ing to the difficulty of keeping perm anent cu ltures o f the B lastocladiales and Leptom itales, p lants o f these mem bers were m ounted on slides in g lycerine jelly fo r reference purposes.

208 K ath leen M . C rooks:

C u ltu r a l M e th o d s .

( i ) S A P R O L E G N IA L E S .

.W hen the filam entous threads appeared on the bait, they w ere lifted off, and tran sfe rred to dishes o f sterile w ater containing boiled halved hem p seeds. Sometim es m ixed cultures were obtained, but as Saprolegnia, . A ch lya , or D ictyuchus readily fo rm ed sporangia, one of the la tter was cut off, tran sfe rred to ano ther dish, an d thus pu re cu ltures w ere obtained. Sometimes the sex organs developed readily, e.g., in Sapro legnia fe ra x , but in o ther'cases, difficulty was experienced in obtaining them , as in A ch lya conspicua. T he ir developm ent was o ften very localized — appearing in only one place in a dish contain ing several hemp seeds.

A species o f A chlya , also one of D ic tyuchus (la te r regarded asD. sterile, or one grain of a heterothallic D ic tyu c h u s), did not develop sex organs on hemp seed, and attem pts w ere m ade to obtain them by grow ing the fungi on hemp seeds in different cu lture media, also on solid media.

T he follow ing m edia w ere u s e d :—

(a ) A gar fo r Saprolegniaceae (recom m ended by W . H ohnk1932)— 1 litre distilled w ater, 8 -10 gm.- agar, 5 gm. carragen (o r if no t available 5 gm. ag a r) , 0.5 gm. gm. dextrose, 0.05 gm. citric acid.

Stock solution— 0.005 gm. K H 2P 0 4, 0.000025 gm. N,H4N 0 3, 0.000025 gm. M g .S 0 4, 0.000025 gm. ( N H J 2S 0 4.

( b) Levulose and peptone agar— Levulose 1.2 per cent.,peptone 0.1 per cent.

( c) G lucose and peptone agar—-Glucose 1.2 per cent., peptone, 0.1 per cent., agar 1.5 per cent.

( d ) Solutions o f m altose and peptone in w ater— 1. M altoseand peptone .0125 per cent. each. 2. 0.3 per cent, m altose and 0.1 per cent peptone. 3. 0.05 per cent, m altose and 0.025 per cent, peptone.

( e) Solution of W itte ’s peptone— 0.1 per cent., leucin 0.1per cent., m altose 0.5 per cent.

( / ) Solution of saccharose— 0.025 per cent, in w hich cubes of egg album en w ere placed instead of the hem p seeds.

In all o f these good g row th was obtained, bu t no sexual organs. K anouse (1932) had considerable success w ith some of these peptone cu ltures in com bination w ith m altose or glucose, and also w ith the addition of leucin, in inducing the form ation of oogonia and an therid ia in Sapro legnia parasitica— a form w hich until then was regarded as s te r i le ; but w ith these m edia, the w rite r could not obtain sex organs in either th & A ch lya sp. or D ictyuchus

S tu d ies on A ustra lian A qua tic Phycom ycctes. 209

sp. A m ethod described by Couch (1932) w hich was very successful w ith Lepto legnia candata was also tried , together w ith certain m odifications of it, by tran sfe rr in g the grow th from the maltose and peptone solutions described above, to sterile w ater. In only one case w ere oogonia obtained ; the A ch lya grow th was transferred from a solution containing 0.025 per cent, m altose and 0.025 per cent, peptone to sterile w ater, and af te r th ree to fou r days, num erous oogonia appeared, but no antherid ia. T he oogonia' degenerated w ithout the developm ent of eggs.

E xperim ents w ere then tried using heteroaux in , a p lant grow th horm one, and an A -Z solution, a solution contain ing traces of th e ra re r e lem ents; in this case 35 were present. The follow ing concentrations w ere used :—

(a ) j q o q̂ q q - heteroaux in in maltose and peptone, 0.025 per

cent each.

( b) gQ qqq heteroaux in in 0.025 per cent, m altose and

0.025 per cent, peptone.

' (c ) rn Jinn- heteroaux in in 0.025 per cent, saccharose.v ' o0,000 r

(d ) | q q q q q heteroaux in in w ater.

( e ) A - Z solution, 2 -6 drops in 0.025 per cent, m altose and0.025 per cent, peptone.

( / ) A -Z solution, 2 -6 drops in 0.025 per cent, saccharosesolution.

In all cases, a good vegetative grow th was obtained but no sexual organs.. W hen the mycelium was tran sfe rred from these solutions to sterile w ater, oogonia appeared in all cases a f te r a

few days, but w ere most num erous in the solutions of ,-q q q q

heteroauxin. Tw o an therid ia appeared, and, in all cases the oogonia again degenerated a f te r a day or tw o, suggesting either that the sexuality-inducing substance was not present in sufficient am ounts to carry the developm ent fa rther, o r tha t the p lant in question is the fem ale stra in of a heterothallic species.

A series of experim ents was also carried out, follow ing those of Leonian (1936), an account o f which appeared ju st before the completion of this paper. U sing pea-in fusion as the sexuality -' prom oting substance, Leonian obtained considerable success w ith P hytoph tho ra cactorum . H ow ever, in the case of A ch lya sp., the pea-in fusion did not induce the form ation of any sexual organs, not even the oogonia obtained in the previous solutions.


I t is evident th a t physiologically each species is extrem ely specialized, as has been show n in the past by the experim ents of d ifferent w orkers, and substances w hich m ay induce the fo rm ation of oogonia in one species will have no effect in even another species of the genus.

( i i ) B L A S T O C L A D IA L E S A N D L E P T O M IT A C E A E .

Prev ious w orkers have show n the ex trem e difficulty o f obtaining these form s in culture. Coker (1923) has obtained a slow grow th of Saprom yees in vegetable juices obtained by boiling corn gra ins or peas, and V on M inden (1916) described a m ethod of grow ing A ra iospora and R h ip id iu m europaeum in culture, bu t all attem pts w ith the B lastocladiales have given negative results. Since these form s grow readily on rose hips, a solid m edium was tried in w hich an ex trac t from rose hips was solidified w ith agar. A n in fusion from rose hip’s was also used, but no grow th was obtained.

' Slopes of carro t partly covered w ith w ater w ere inoculated w ith p lants of Blastocladia, Saprom yees, and Gonapodya, but gave negative results. A bacterial scum quickly develops, but th is is not the only facto r proh ib iting grow th. Saprom yees was found to grow slowly in prune ju ice, bu t no new plants appeared, show ing th a t neither the zoospores nor resting spores germ inated. I t is suggested th a t ow ing to the extrem e sensitiv ity of the protoplasm in these form s, the mere act o f d isturb ing them by transference from the original -substratum , renders them incapable of germ ination.

A nother m ethod was tried , so as to have cu ltu res of these form s fo r continuous study. I t was found that w hen baits w ere b rought in to the laboratory, pustu les w ere form ed in addition to those already present. F resh apples w ere added to dishes contain ing infected apples, in the nope that they would be attacked, but the result was negative. E ven w hen pustu les were tran sfe rred directly to the fru its no' fu r th e r in fection occurred, although the fungi apparently continued to grow. I t would appear that, in the lakes or stream s in which the traps are placed, bacterial action is responsible fo r the soften ing of the fru it surface- before fungal attack. N atu rally baits brought into the laboratory were w ashed frequently to keep dow n bacterial contam ination, and it was thought that th is m ay have prevented the apples from being .attacked. S terile tap w ater was used fo r w ashing the baits at first, but la ter on ju s t o rd inary tap w ater. A nother experim ent was tried in w hich fresh apples w ere placed ‘in dishes w ith the bait, and the w ater was not changed. A f te r two m onths, the apple skin was softened, bu t no in fection had occurred. I t is there fo re concluded th a t in lakes, etc., decom posing bacteria and organism s, not found in tap w ater, p repare the f ru it surface fo r fungal attack.

The fact th a t these fungi o ften possess rhizoidal system s should not be wholly responsible fo r the difficulty in obtaining them in culture, bu t evidently, parts o f p lants, e.g., filaments broken off, do not produce new plants as in Saprolegniaceae, Pythiaceae, etc., and the fo rm ation of p lants m ust be entirely dependent on the germ ination of the spores— the protop lasm of which is extrem ely sensitive to any d isturb ing forces and to environm ent.

D e s c r ip t io n o f t h e F u n g i in C u ltu r e .

S A P R O L E G N IA L E S . S a p r o le g n ia c e a e .

S z p r o l e g n i z f e r z x (G ru ith .) T huret.

A n n . S c i. N a t . B o t., S e r ie s 3, 14: 214, p l. 22, 1850.

Saprophytic on Crataegus fru its a t U niversity , collected 5th A pril, 1935; grow th ra the r coarse on hemp see d ; sporangia, term inal, cylindrical, p ro liferating w ith an average diam eter of 54 .5 /a, and a great varia tion in length, o ften 'co n str ic ted by the m ouths of earlier sporangia; gemmae py rifo rm to oval, and occasionally bearing oogonia, frequently in chains (F ig . 1, B ) ; oogonia spherical to oval, occasionally cylindrical w hen developed in an em pty sporangium , borne in a term inal, lateral, or in te rcalary position on the hyphae; wall about 1.9/* th ick, w ith conspicuous pits w hich have a d iam eter range of 6-10/a, average d iam eter 7 /a ; frequently , a narrow hypha is observed to pass into the cavity of the ogonium, from the apex of the oogonial s ta lk ; spores-—1-30 per oogonium ; centric, 18-34/a in d iam eter, usually 25/a; antherid ia found on about 60 per cent, o f the oogonia; mainly androgynous but of diclinous orig in when the oogonia are borne term inally. W hen the oogonium arises on a short lateral branch, the antherid ium arises from the stalk o f the oogonium ; an therid ia usually slender and branched, but only one or few found on any one oogonium (F ig . 1, D -G ).

S tu d ies on A ustra lian A qua tic P hycom ycetes. 211

F ig . i .— A-G , S a p ro le g n ia f e r a x (G ru i th ) T h u re t. H *L, D ic t y u c h u s m o n o s p o ru s Leitgeb. A , S porang ia , B , Gem m ae, C, Gem m a con ta in ing eggs, D , E , F,O ogonia and A n th e rid ia . A ll X 3 3 - G, Oogonium con ta in ing m a tu re oospores. X 2 0 7 . H , I , Spo rang ia o f no rm al type. X 3 3 . J* R esting sporang ium about

to fa ll off. X 3 3 . • K , P o rtio n o f sporang ium show ing netw ork . X 2 0 7 . L , O ogonia w ith enc irc ling a n th e rid ia . X 2 0 7 .

212, .K a th leen M. . C rooks:

T here has been m uch discussion as to w hether 5'. fe r a x and 6". m ix ta should be regarded, as two distinct, species. Formerly S', fe ra x was distinguished from S', m ixta , in possessing an therid ia on usually less than 10 per cent, of the oogonia, but it has been found that the same cu ltu re m ay at d ifferent times vary in the percentage of oogonia-bearing antheridia.’ A ccording to Forbes (1 5 ) , the species form erly know n as S', m ix ta is only a grow th fo rm of S'. fe ra x , and the same applies to S', m o n o i c aso that S’, fe ra x is regarded as one good species which shows different grow th form s under vary ing environm ental conditions. H ence, the form in question, since it possesses antherid ia on approx im ately 60 per cent, of the oogonia, m ost closely approxim ates to S’, fe ra x “ fo rm a m ix ta .1’ .

A c h l y z c o n s p i c u z .

Coker in “ T h e Sap r o legn ia ceae,” 1923, p. 131.

Saprophytic on Crataegus fru its and tw igs in lake a t U niversity , 5th A pril, 1935 : th is fungus has a coarse w hite grow th on hemp seed, the hyphae reaching a d iam eter of 80ft,; sporangia abundant, arising by lateral branching, 9 0 - 4 0 0 /a in length and 1S—50/x in bread th (F ig . 2 C ) ; gem m ae frequently form ed especially in -old cu ltu res ; they resem ble sporangia in shape but are larger, frequently have prongs, giv ing a fo rked appearance; oogonia not always developed, but when present are borne laterally on the m ain hyphae; spherical in shape, 55-80/a d iam eter; the walls are not very thick but pitted, the d iam eter of the p its vary ing from5.5-7.5j». U sually 4 -1 0 oospores are developed in a single oogonium but over 20 have been observed in a few cases, ranging in d iam eter from 2 1 - 3 0 /a ; eggs rarely m ature, bu t nearly always degenerate im mediately and become filled w ith oil drops. W hen they do m ature, the eggs are sub-centric, sm aller droplets su rround ing the subcentric larger g lobu le ; A n theridia always present, usually androgynous, but occasionally diclinous, arising near the oogonial stalk from the m ain hypha, o r frequently from the oogonial stalks them selves; an therid ia are simple o r branched, one o r som etimes tw o to each oogonium , usually touching the oogonium by foot-like pro jections (F ig . 2, D. E ) .

A (c h l y z z m e r i c z n z var.—m e g z s p e r m z var. nov.

Saprophytic on rose hips, in lake in F itz roy G ardens, M elbourne, 20th July, 1 9 3 5; hyphae variable in d iam eter, bu t mainly coarse reaching a d iam eter of 100/a; sporangia tapering tow ards the tip, usually up to 3 5 0 /a in length, and about 4 0 /a in the w idest p a r t; gem m ae few, in chains o f 2 o r 3 (F ig . 3, E ) ; oogonia are num erous, borne in a racem ose fash ion along the h y ph ae ; oogonial stalks usually about the sam e length as the d iam eter of the oogonium or less than rarely \ \ tim es the d iam eter of the oogonia; the la tter are occasionally borne term inally on the

S tu d ies on A ustra lian A quatic Phycom ycetes. 213

h y p h ae ; oogonia sp h er ica l,' m ainly 40-100/;. in d iam eter, walls th in and p itted, th e -p its usually fa ir ly conspicuous; oospores 2 4 -3 1 /i in d iam eter, average 27//, 2 -2 0 per oogonium, usually 6 -8 ,.eccentric; an therid ia branched, androgynous, but occasionally diclinous, one to fou r on each oogonium, one an therid ium often supplying tw o oogonia: an therid ia usually arise from the m ain hyphae, occasionally from the oogonial stalk, and are slender, long and branching (F ig . 3, F. G ) : In the cu ltu res o f th is fungus, it was found th a t the eggs frequently disorganized before reaching m aturity , leaving the oogonium filled w ith oily masses.

T h is species appears to resem ble A . am ericana but d iffers from the description of the type species in the larger size of the oospores, which rang from 24 -3 1 p , average diam eter 27^, as against 18.5-25/*, average 22/x of the latter. Several w orkers found it impossible to separate A ch lya de B aryana and A ch lya americana as two distinct species; but Forbes (1 9 3 5 ) distinguishes them as fo llo w s:— In A . de Baryana, the oogonial walls are u n p itted and some diclinous an therid ia occur, while in A . americana, the oogonial walls are pitted, and the an therid ia are consistently androgynous. C oker describes a fo rm re ferred to A . americana, in which the oogonial walls are p i t te d ,, but the antherid ia m ay occasionally be diclinous, so that our species, having p itted oogonial walls, occasional diclinous antherid ia, the la tte r sometimes arising from the oogonial stalk, a featu re never found inA . de Baryana, seems to be a varie ty of A . americana.

O w ing to. the large size of the oospores, this fo rm is re ferred to A . americana var. m egasperm a var."nov.

A nother variety of A . americana, namely, A . am ericana var. cambrica, possesses characters not found in th is species, nam ely oogonia w ith th ick yellow ish walls, and oospores w ith a very thick tw o-layered wall, and 23 -2 6 / i in diam eter.

F i g 2 .— A, B , A c h ly a s p . ? C -E, A c h ly a c o n s p ic u a Coker. A . Oogonia. X 33-. fr , Oogonia w ith androgynous an th erid ia . X 33- E , O ogonium w ith an th erid iu m . X 12 5.

214 K ath leen M . CrOoks:

A c H L y a o b l o n g a t a .

de B a ry . B o t. Z e it . : 646, p i. it), f ig s. 7 -9 , 1888.

Saprophytic on silver-fish; collected 17th June, 1936; mycelial grow th fa ir ly coarse, hyphae up to 9Q(i d iam ete r; sporangia typical of the genus, bu t occasionally dictiosporangia occur; zoospores 9-12/x d iam eter; gem m ae frequently form ed and are oval o r elongated; oogonia very large, oval, pyrifo rm or occasionally spherical, 85-300jU, x 70-160/x, borne on lateral branches, term inally on the m ain hyphae or rarely in tercalary ; wall of oogonia th in and u n p it te d ; eggs 12-30 per oogonium, fa ir ly small, 23-30/x in d iam eter, m ostly about 26/x, sub-centric, w ith a sheath of oil droplets all or .most of the way around the protop lasm (F ig . 3, C, D ) ; eggs at first dark, becoming lighter in appearance, but rarley m a tu r in g ; an therid ia long, slender, diclinous, usually several w ound around each oogonium, tuberous at the point of contact w ith the oogonium (Fig.. 3, A , B ).

F i g . 3 .— A -D , Achlya oblongata de B ary . E -H , Achlya americonu . v a r. megaspertna nov. v a r. A , O ogonium w ith long, d ic linous a n therid ia . X 3 3 . B , In te rc a la ry oogonium w ith a n th e rid ia .X 3 3 . Cj M a tu re oospore w ith oil g lobules all a ro u n d p ro to

p lasm globule. X 2 0 7 . D , M atu re oospore w ith oil g lobulesonly p a rtly su rro u n d in g protop lasm globule. X 2 0 7 . E , G em m ae. X 3 3 - F , G, O ogonia w ith a n th e rid ia . X 33-H , O ospore of eccen tric type. X 2 0 7 .

Saprophytic on Japonica fru its, lake at M aryborough, collected 13th February , 1936; grow th on hem p seed fa ir ly coarse, hyphae, 15—100/x in d iam eter; sporangia developed fa ir ly abundantly , p ro ducing v ery sm all zoospores; gem m ae num erous, cylindrical, o ften very long, term inal ones o ften tapering to a point, frequently developed in chains along the length of a hypha (F ig . 2, B ) ; oogonia sm ooth, spherical, 40-75fx in d iam eter, w ith a th in, unp itted wall, borne laterally on the hyphae; a t first denser in appearance, bu t rap id ly degenerate and lose the ir contents (F ig .2, A ) ; eggs never observed ; occasionally slender branches w ere noticed close to oogonia, bu t as they were never observed to

A c h l y z s p . ?

a p p r o a c h t h e O O g o n ia , th ey canno t b e r e g a r d e d a s a n t h e r i d i a , a n d

iti m ost c a s e s , t h e r e W a s no s i g n o f a n t h e r i d i a l b r a n c h e s n e a r t h e

o o g o n i a .

I t is considered th a t th is m ay be a fem ale stra in of a heterothaliic specie's o f A ch lya . Oogonia have been observed in large num bers, when the fungus was grow n in various m edia (an account o f these experim ents is given earlier in th is p ap er), but they always disorganized w ithout any sign of oospore form ation. H eterothallisfn in the Saprolegniaceae has been reported previously— in D ictyuchus by Couch (1926) and for A ch lya by Coker (1923). Coker states th a t his heterothaliic species—A ch lya b isexualis n.sp.— is probably the same as one previously described by him ih the Saprolegniaceae, as A ch lya sp., form w ithout oogonia. O u r plant differs from A . b isexualis in the shape of the gem m ae which are always cylindrical, in contrast to the spherical or pear-shaped gemmae of the latter.

D i c t y u c h u s s p .

Saprophytic on rose hips, pond in Botanical G ardens, M elbourne ; on P ru n u s tw igs in lake in G ardens, a t M aryborough, V ic to r ia ; the m ain hyphae are 20-60/x in d iam eter, tapering tow ards the end, w here they have a d iam eter range of 18-35/n; sporangia form ed at the tips of the hyphae, and lateral ones are form ed by cymose branch ing ; in o lder cu ltures, the hyphae them selves become segm ented into sporangia, o ften irregu la r and b e n t : frequently the sporangia break aw ay from the hyphae bearing them , and a.re know n as resting sporangia. Sporangia developed along the length of a hypha, if the la tter is a fine one, m ay be thread-like, w ith only a single row of sp o res; zoospores ll-14 ju in d iam eter, and the spores encyst w ith in the sporangium for a period a f te r which they em erge by indiv idual openings in the sporangium wall, leaving a netw ork of walls in the la t te r ; sexual reproduction not observed.

T h is isolation agrees in all particu lars w ith D ic tyuchus sterile. H ow ever, Couch has show n th a t heterothallism is common in the genus D ictyuchus, so it is likely also that this m ay be One stra in of a heterothaliic form .

D i c t y u c h u s m o n o s p o r u s L e i t g e b .

Jahrb . f . W is s . B o t., 7 : 3S7, p l. 22, f igs. 1-12, p l. 23 , f ig s . 1-8 , 1869.

Saprophytic on rose hips in lake in F itz roy G ardens, -20th July, 1935; hyphae fa ir ly coarse, reaching d iam eter of 60/*; sporangia form ed abundantly , in sym podia and also along the length of the hyphae; the m ethod o f ejection of the spores is typical o f the genus; zoospores 9 -1 5/x in d iam eter; oogonia term inal on the branches,- spherical, 25-35/z in d iam eter, average 31.5/x, w ith unp itted w alls; oospores borne singly in the oogonia, Centric ,

S tu d ie s on A u s tra l ia n A q u a t ic P h y c o m y c e te s . 215

216 K a th le e n M . C ro o k s :

sm ooth, 22 -31 /i diam eter, average 25.7/i.; 'an th e rid ia always developed, usually several in contact w ith each oogonium, diclinous and encircling the oogonia (F ig . 1, L ) .

T h is fo rm agrees in general characters w ith the type species, except th a t in our species, the oogonia and oospores are slightly larger. L und (1934) has described a species agreeing w ith the one in question except th a t L u n d ’s fo rm has eccentric eggs. L indsted t (1872) stated th a t in D. m onos porus, the oogonia are 25/i th ick, and the sporangia borne only in sympodia, while in D . M agnusii, the oogonia are 30-35,j, thick, and the sporangia borne only in rows. In th is species other sporangia showed both m ethods of fo rm ation, and while the oogonial size is large, sim ilar to th a t o f D . M agnusii, it is thought th a t th is form approx im ates m ore closely to D. m onosporus, particu larly in the character of the antherid ia, which are alw ays found to w rap around the oogonia. L und (1934) states th a t D . m onosporus developed no sexual organs w hen grow n on hem p seed, but the au tho r has obtained them in hem p seed, cultures, although they w ere not observed in the orig inal substra ta, namely, rose hips and tw igs.

L E P T O M IT A C E A E .

S z p r o m y c e s R e i n s c h i i (S ch ro e ter) F ritsch .

O ste r r . B o t . Z e itsch r . 43, p. 420, 1893.

Saprophytic on rose hips and apples in lake at U n ivers ity ; collected 5th A pril, 1935; saprophytic on apples in B arham R iver, near Apollo Bay, collected 1st June, 1936. T h is fo rm was found in abundance, and was frequetn ly associated w ith G onapodya and Blastocladia. W hile Blastocladia produces a short com pact tu f t, on tlie fru its, tu f ts contain ing Saprom yces alone w ere no t so dense, and pro jected fu r th e r from the fru its.

P lan t attached to substratum by well-developed rhizoids. the rhizoidal system being o ften very ex tensive; basal cell including the rhizoids, varies fro m 3 0 0 -1 2 0 0 / i in length and 15 -3 0 / i in breadth, and produces d istally finer branches which are cylindrical, segm ented and 8 - 1 5/t in d iam eter (F ig . 4 , A ) ; reproductive organs arise from these branches o r fro m the finer filaments w hich arise from th em ; constrictions between the segm ents usually closed w ith cellulin p lu g s ; asexual reproduction by zoosporangia borne singly o r usually, in groups of 2 -5 a t the end o f the filam ents or a t the constrictions. Zoosporangia 4 0 - 1 7 0 / i x 1 5 -3 0 / i elongate and usually w ider about the m iddle than at either e n d ; zoospores escape by a term inal pore o r occasionally germ inate inside the sporangium . T he m ethod of escape m en tioned by K anouse ( 1 9 2 7 ), in w hich t h e ’whole zoospore mass em erged, surrounded by a m em brane w hich is im m ediately rup tu red , w as.-a lso observed; sexual reproduction, no t always

present in the m ateria l exam ined, bu t if present a t all was abundan t; oogonia borne term inally o r laterally, and o ften associated w ith sporangia iii the same w horl; oogonia 41-63/i x 29-40//, p y rifo rm to sub-spherical, w ith a well-developed pedicel. W hen m ature the oogonial walls are covered w ith a yellow ish-brow n incrustation w hich is usually th icker a t the apex, and a single spherical oospore is developed with in each oogonium ; oospores vary in size from 26-34/t, wall fa ir ly th ick, b row n externally but colourless w hen viewed in optical section, occasionally slightly irregu lar, bu t the series of irregu lar ridges noted by Sparrow (1936), w ere not v isib le; an theridia always present and usually of diclinous o rig in ; swollen and clavate iii shape, 30-40/a x 9-1 3 / x , borne on long, w inding, tw isted branches, 4-6/x in d iam eter, and send a prom inent fertilization tube into the oogonium (F ig . 4, B ). O ne exam ple was found of an androgynous antherid ium said to be characteristic of. S , androgynous. O n the sam e p lant there were num erous oogonia fertilized by an therid ia from different plants, and th ere fo re of diclinous orig in (F ig . 4, C ). T h is is in teresting, in view of the fact th a t Sparrow (1932) m entions th a t experim ents w ere being carried out by P . H . Jo rd an which seetited to indicate th a t th is

S tu d ie s on A u s tra l ia n A q u a t ic P h y c o m y c e te s . 217

F IG. 4 .̂ -Sixpromyces Reinsckii (Schrfleter) Fritsch.A , P la n t s h o w i n g w e l l d e v e l o p e d f h i z d i d a l s y s t e m a n d b e a r i n g r e p r o d u c t i v e o f g & n s . X 3 3 - B » Oogo- n i u f t i w i t h d i c l i n o u s a f i t h d f i d i i i n i . X 2 0 7 . C, T w o o o g o n i a f r o m t h e S a m e p l a n t , one W ith diclinous a n t h e r i d i u m , t h e o t h e r w i t h androgynous a n t h e r i

d i u m . X 2 0 7 .

species is heterothallic. I t has also been noted tha t a well- developed rhizoidal system is present, as previous w orkers have suggested that they are not developed. Sparrow (1932) stated th a t the narrow basal cell is “ presum ably anchored by ho ld -fasts,” bu t in all the p lants exam ined in th is collection except those that w ere obviously broken above the point o f attachm ent of the substratum , rhizoids w ere present (see P la te X ., 7, 8 ) .

6 5 8 . - 5


R h i p i d i u m a m e e i c a n u m T hax ter.

B o t. Gaz.. 2 1 : 320, p l. 22, f ig s . 1-15, 1896.

Saprophytic on rose hips, in Lake at U niversity , 5th A pril, 1 9 3 5; on apples, Japonica fru its , Cotoneaster berries, in lake in 'M aryborough G ard en s; p lan t attached to substratum by m eans of rh izoids, sometimes copious, a t o ther tim es scanty ; basal -cell pe lta te,' 180-300/u. from rh izoids to apex, and in surface view, the apex has an irregu larly stellate appearance (P la te X . 6 ) ; stalk varies in d iam eter from 23-50//,, and at the top m ay divide dichotom ously to form branches or lobes w hich spread in a rad iate fashion, or m ay expand into a flattened head which is lobed or branched around its circum ference (F ig . 5, A, B ) ; d iam eter of the apex varies fro m 9 0 -2 5 0//., and has a fa ir ly th ick wall 3 -7 //, but the d iam eter o f the indiv idual lobes range from 10 //-90 ,/. F rom th is basal cell arise filam entous branches 1 5 0 -7 0 0 ,/ x .3 - 1 0 /a , and they are swollen just above their, attachem ent to the thallus. These filaments are usually segm ented, resem bling the filaments of Saprom yces R einsch ii. Zoosporangia b o rn e . term inally on the filam ents, usually 2 o r 3 together, less o ften singly, ovoid, 46-100// x 16-40/* (F ig . 5, C, D ) ; zoospores observed to emerge in a cylindrical m ass surrounded by a th in m em brane which rup tu res and allows th e zoospores to escape; sexual reproduction not observed, although a large num ber of p lants w ere exam ined.

F i g . s .— R h ip id iu m am e rx c a n u m T h a x te r. A , B , P e lta te th a ll i show ing i r re g u la r b ranch ing i X 33> G* S po rang ia occurr in g sing ly , o r’ in tw os ' o r th rees. X 3 3 . ' D , S porang ia . X 12 5. E , P o r tion o f th a llu s show ing th ic k w alls, and the basal p a r ts o f f ilam ents. X 20 7.. V

T he natu re of the thallus, although constant in form , in the p lants exam ined here, is very varied as show n -by the descriptions, of o ther w orkers. Forbes (1935) describes, p lants in w hich the basal p art is cylindrical and branched, resem bling the thalli of som e Biastocladias. S parrow has described R h ip id iu m am ericanum show ing exam ples of both the" peltate -th a ll i and: ther branched ‘ tree-like p lants. Since th e sp o ran g ia l ' measurem ents are sim ilar to those;'of p lants described in o ther collections it is


likely that R h ip id iu m am ericanum is a species like Blastocladia Pringsheim ii in which the thallus is extrem ely variab le in size and shape.

Saprophytic on tw igs, in lake a t U n iversity g ro u n d s; collected 5th A pril, 1935; mycelium delicate, hyphae of un iform d iam eter, vary ing from 2 - 5 /a ; a few bud-like pro jections from the hyphae are sometimes found, bu t are not characteristic as in P yth iom orpha gonapod io ides; sporangia developed in w ater cu ltures, spherical to py rifo rm in shape w ith a very long papilla", a t m aturity , they m easure 2 6 - 4 6 /a x 2 0 - 3 2 /a (F ig . 6, F ) ; p ro liferations occur, the secondary sporangia being form ed inside the old one, or the hypha may grow th rough it and form the new sporangium at some distance fu r th e r on ; sporangia frequently function as conidia producing gerrri-tubes, but they also w ere observed: to produce zoospores in the true P y th iu m fashion— th at is, the protop lasm is

F ig . 6 .— A-E,. Pythiomorpha gonapodioides P e te rsen . F -H , Pythium proliferum de B ary . A , B , Y oung sporang ia. X 2 0 7 . C, P ro l i fe ra tin g sporang ia. X 2 0 7. D , E m p ty spo rang ia w ith the hypha g row ing th rough . X 33. E , O ogonium w ith th ick th ree-layered wall. X 2 0 7 . F , S porang ium w ith pap illa . • G, P ro l i fe ra tin g sporang ia. A , E m p ty sporang ium th ro u gh which hypha has grow n. I , Zoospore. J , Spo rang ium w ith vesic le. A ll X 2 0 7 .

extended into a vesicle w here the zoospores are d ifferentiated (F ig . 6, J ) ; in one case observed, 30 m inutes elapsed from the tim e the vesic le 'w as fo rm ed until, the zoospores w ere set f ree ; in the vesicle they are spherical, and, appear to escape at a th in point in the. wall, they w ere constricted as they passed th rough, and assum ed a. ren ifo rm . shape in the sw im m ing stage,, but settling down, , they became spherica l. ag a in ; no sexual organs observed. " .,

P y t h i u m p r o l i f e r u m d e Bary.

P r in g sh . Jah rb . W iss . B o t . 2, p. 182, 186&.


P y t e u o m o r p h z g o n z p o d i o i d e s H . E . Petersen.

B o t . T id ssk r . 29, p. 3,91, f igs. V I . and V I I . , 1909; and in A n n .M yc. 8, p. 528, f igs. V I . and V I I . , 1910.

Saprophytic on rose h ips in pond in F itz roy G ardens, collected 20th July, 1 9 3 5; on grapes, prunes, and rose hips in pond in Botanical G ardens, collected 23rd M ay, 1 9 3 5; on oranges in M cC allum ’s Creek, near M aryborough, collected 12th February , 1936. In the case of the grapes, the pond w ater was brought into the laboratory and baited w ith the fru it. H yphae irregu lar in appearance, ow ing to the fact th a t a sirlgle hypha m ay vary in d iam eter from, 2- 8 / a ; bud-like pro jections are frequently developed on the m ycelium ; sporangiophores of un iform diam eter, slightly narrow er than the o rd inary hyphae; sporangia pyrifo rm , papillate and p ro liferate frequently , the new sporangium form ed w ith in an old one o r a t som e d istance beyond it, and frequently a row of sporangia is developed (F ig . 6, D ) ; range in size from 40-70/x x 20-40/a. Sporangia set free zoospores w ithout the fo rm ation of a vesicle; zoospores— spherical, 9 - 1 3 /a diam eter, usually m ore than 20 per spo rang ium ; sporangia w ere present on the m ycelium on the f ru it arid hips w hen brought in; oogonia spherical, 2 3 - 2 8 /a diam eter, and when m ature have a th ick three- layered wall (F ig . 6, E ) ; oospores yellow ish in colour and 20-22/t diam eter. O nly one an therid ium was noted, and th is was clavate in shape and of diclinous origin.

•This fo rm agrees in all particu lars w ith P yth iom orpha gonapodio ides except in the size of the sporangia which are slightly larger in ou r form .

BLASTOCL ADI ALES.

B la s to c la d ia c e a e .

G o n z p o d y z s i l i q u A e f o r m i s (R einsch) T hax ter.

B o t. G az., 20': 480, p l. 31, f ig s . 6 -1 0 , 1895.

Saprophytic on rose hips and tw igs in pond at U niversity , and on rose hips again in pond at the F itz roy G ardens; pustu les on the f ru it differed from those of B lastocladia in form ing loose spreading n tats of hyphae, in con trast to the short dense tu fts of the B la sto d a d ia s; vegetative p lant attached by scanty rhizoids, consists of hyphae m ore or less sparingly branched in the low er parts; but m ore copiously branched in the upper parts, g iv ing a successively um bellate appearance; whole p lant segmented, each constriction correspondir^g to a *■ pseudo-septum form ed by a deposit of ce llu lin ; som etimes segm entation is alm ost entirely absent (F ig . 7, D ) , although the cellulin plugs m ay be p resen t; iti the basal parts, the segm ents are long and narrow bu t in the upper part sh o rt'an d rounded ; sporangia, 60-220 x 1 5-35/a, podshaped, frequently p ro liferating (F ig . 7, C, D ) , borne term inally on the hyphae or in umbels som etim es separated by a definite

S tu d ies on A ustra lian A quatic P hycom ycetes.

co nstr ic tion ; zoospores num erous, hyaline w ith conspicuous nuclei and w eak granu lar fo rw ard end, one backw ard m oving c il iu m ; sexual reproduction no t observed.

G o n z p o d y z p o l y m o r p h z T hax ter.

B o t. Gaz., 2 0 : 481, p l. 31, f igs. 11-16, 189S.

Saprophytic on Japonica fru its in lake a t M aryborough G ardens; vegetative p lant consists of m ore slender hyphae m w hich segm entation is no t pronounced, which give rise to tu fted um bellate branches composed of oval.-o r irregu lar segm ents; sporangia borne term inally on the segm ents, one to th ree arising from each segm ent, ra ther oval in shape, and truncate at the apex a f te r dehiscence, size 8 5 -1 10/i x 35-45//,; sexual organs not observed. T he habit of th is p lant strongly resembles th a t o f

F i g . 7 — A , G o n a p o d y a p o ly m o rp h a T h a x te r F ilam en ts w ith p ro li fe ra tin g sporang ia. X 2 0 7 . B -E , G o n a p o d y a s iU q u a e fo r tm s R em sch. B , Yoiing sporang ia. X 3 3 . C, D , S po rang ia show ing ear ly stages 111 p ro life ra tion .X 3 3 . E , Z oosporang ium w ith zoospores. X 15 0 -

G. po lym orpha, bu t the size of the sporangia is definitely larger. H ow ever, the appearance is so sim ilar to G. po lym orpha, th a t it m ust be regarded as th a t species, in w hich the sporangia a ie larger, possibly due to d ifferent environm ental conditions.

B l z s t o c l z d i z (R einsch) Em end.

P r in g sh . Jah rb . W iss . B o t. 2 , p. 182, 1860.

P lan t consists of large basal cell of very variable form , short and swollen, or long, cylindrical, branched o r unbranched, the apices o f the tru n k o r lobes o ften expanded; plant attached to substratum by rh izo id s; filamentous hairs', simple o r branched, o ften arise from the basal cell on its branches, and these hairs have a bulbous base; zoosporangia sessile on the basal cell, variable in size and shape; zoospores num erous, usually uniciliate, em erging th rough a term inal p o re ; resting spores sessile like the sporangia, usually oval or spherical in shape; an therid ia have been observed in one species by K anouse (1927), who concludes th a t the “ resting-spores ” should really be regarded as oogonia.

K ath leen M y CroO’ks:

B

B . globosa

C

U p to the present, ten species of B lastoclad ia have been described, and a' new species is given here, nam ely Blastocladia aspergillo ides n.sp. ‘

K e y to, the Species.

A. Basal cell swollen o r cylindrical w ithswollen heads, branched o r u n branched . . . . 1

Basal cell cylindrical, usually m uch branched, the apex of the thallus

■ or its branches not swollen

B. Basal cell globose, slightly narrow edtow ards base, sporangia 55-130 x 16-40/t . .

Basal cell cylindrical in the lower part, w ith swollen h e a d s ; thallus branched or unbranched

C. P lan ts 140-260/i, basal cell u n branched, sporangia long, narrow , cylindrical, 85-150/i x &^-15/i

P lants usually larger, branched or u n branched, sporangia ovate o r broadly cylindrical

D. Sporangia p ro liferating Sporangia not p ro liferating

E. P lants tall, 600-950/i x 20^1-5/i,branched, hya line ; sporangia borne in a racem ose or corymbose m anner, 120-220/i x 22-34/i

P lan ts sm aller, sporangia on the apex o f the thallus o r its branches

F . Sporangia long, narrow , cylindrical,60-210/i x 8-25fx

■ Sporangia broadly cylindrical or ovalG. P lan t very hyaline, sporangia oval,

27-40/i x 7-13/t, not p ro liferating P lan t dark in colour, sporangia

broadly cylindrical, not narrow ed a t "base, p ro liferating , 80-140/i x 18-35/i . . . . . B.

B . aspergillo ides n.sp.

D

B . p ro life r a B . P ringsheim ii

B. gracilis

B . augustaG

B . raniosa

B l z s t o c l z d i z P r i n g s h e i m i i Reinsch.

Jah rb . W iss . B o t. 11 : 298, 1878. -, i E m e n d T h a x te r, B o t . Gaz. 21-: 51, 1896. ..

Saprophytic on rose hips, lake at U n iversity , 5th April, 1935:, -Botanic G ardens, 23rd A pril, 1935 ; F itzroy. G ardens, 20th Ju ly , 1935; on banana, Y a rra R iver, R udder Grange, 10th Novem ber,

1935; on- apples. Japonica fru its , w in ter-cherry ..fru its, lake: at M aryborough G ardens, 1.3th-February, 1936;. Cotoneaster berries, Japonica fru its, M cCallum ’s .Creek: , M aryborough, 12th F ebruary , 1936; on apples, B arham River,- Apollo Bay, 1st. June, 1936.

This fo rm is an extrem ely common saprophyte on any decaying fru its , bu t was never found on tw igs of any k in d ; the pustules of Blastocladia P ringsheim ii w ere dense, tu f ted , up to 1 mm. in d iam eter, and yellow ish w hite in colour. T here w ere usually a • large num ber o f indiv iduals in a pustule, and although occasionally a pustu le is composed entirely of p lants o f the one species, usually B . P r ingsheim ii was associated w ith o ther form s, e.g., R h ip id ium , o r o ther B lastocladia sp., bu t no t w ith the filam entous fungi, e.g., Saprolegnia, A ch lya .

P lan t consists of a large basal cell attached to the substratum by rhizoids, and very variable in size and sh ap e ; figure 8 shows the variations in the thallus of B. P ringsheim ii from m aterial obtained from d ifferent sources; basal cell m ay be fa ir ly short, unbranched and swollen tow ards the apex, or long cylindrical, branched or unbranched, the distal part o f the trunks or o f the lobes being ra th e r sw ollen; on the expanded region the sessile sporangia are borne, together w ith filam entous hairs, simple or branched and w ith a bulbous base; sporangia m ore o r less long,

S tu d ies on A ustra lian A qua tic P hycom ycetes. 223

F i g . 8 .— B la s to c la d ia P r in g s h e im i i 'R e in sch . , .P la n ts show ing v a ry in g size and shape o f basa l cell and sporang ia. X 2 5 .

cylindrical, rang ing in size from 70-280/x x 14—70/i over a large num ber of p lan ts; zoospores num erous, 12-15/i in d iam eter, escaping directly th rough the term inal po re ; the form ation o f -a vesicle^ described by V on M inden '( 1916) and S parrow (1932) Was .not observed; zoospore discharge seems to be largely influenced by external conditions, as in m ost o f the sporangia ■ observed, the zoospores d id..not escape but disintegrated- w ith in


the sporang ium ; resting spores frequently found, and always associated w ith the sporangia, oval o r spherical, thick-walled, punctate and 40-99 /* x 3 0 - 5 0 /a .

As suggested by S parrow ( 1 9 3 6 ), B . P r ingsheim ii appears to be a very variable form , both in the na tu re of the thallus, and in the size o f the sporangia. In practically each locality, whence th is fo rm was obtained, the habit varied, as also did the range in sporangial size, g iv ing from; all the p lants examined," a w ide range of 70-280 /* x 4 0 - 7 0 /a , although in indiv idual p lants the range was not o f th is o rder. F rom the m easurem ents o f the length and breadth of the sporangia in 265 indiv iduals, frequency

A.

mTO ISO So nJT-q

(.E»C7H OF SPORANGIA N Jl

etASTOd-AfllA PflNCjHEIMIICF 5PO«ANClA

F ig . g .— B la s to c la d ia P r in g s h e im i i . F req u en cy d iag ram s to i l lu s tra te va ria tio n in length (A ) a n d b read th (B ) o f sporang ia.

colum n diagram s have been prepared and from these the m ax im um frequency in length and breadth can be noted. In the length diagram!, the lengths are arranged in classes w ith an interval o f 10, e.g., 70-79.9, 80-89.9, etc. T hus there are 25 indiv iduals (sporang ia) w ith a length between 150 and 159.9, which class shows the greatest frequency, 21 indiv iduals w ith length between 160 and 169.9, etc. In the d iagram illustrating breadth (F ig . 9, B ) a m easurem ent w hich is not as variable as the length, and hence shows a g reater frequency in the classes (since the same num ber of indiv iduals w ere m easured in both cases) the breadths are arranged in classes w ith an in terval of 5, e.g., 10^14.9, 15-19.9, etc. Th is m ethod was used here instead of classes w ith an interval of 10, only fo r g reater convenience in p reparing the d iagram , ow ing to the g reater frequencies, show n in the classes, e.g., betw een 30-34.9, there is a frequency of 44, betw een 35-39.9, a frequency of 44 also, hence between 30-39.9 a frequency of 80 (w hich w ould be difficult to .express w ith the sCale in u se), w hich is the m ax im um 'frequency . A lthough these

form s appear w idely d ifferent in size, habit, etc., there does not a t present seem to be sufficient justification to separate them as o ther species, particu larly on the basis o f m easurem ents. Size is a very variable character, particu larly when studied over a large num ber o f indiv iduals, and m ay be influenced by ex ternal conditions, e.g., am ount o f food m ateria l present, etc.

B l z s t o c l z d i z p r o l i f e r z von M inden.

K ryp t ogam enf l. M a rk B randenbu rg , 5 : 606, 1912.

Saprophytic on rose hips, lake a t U niversity , collected 5th A pril, 1935; Saprophytic on apples and japon ic a fru its, lake at M aryborough, collected 13th F ebruary , 1936; this species was found in pustules and was sometimes associated w ith B lasto cladia P ringsheim ii or w ith Gonapodya. In the first collection obtained of th is form , on rose hips, the plants w ere a dark bluish- black colour and th is was also characteristic of the B . P r ingsheim ii p lan ts associated w ith it. In a later collection from a different locality, the p lants of B . p ro lifera w ere not strik ing in colour at all, only tinged w ith brow n, and B . P r ingsheim ii when found ' in o ther parts, and on rose hips again, did not have th is bluish-black colour, although p lants o f the latter are o ften dark brow n in colour. T he cause of th is unusual colour is not known.

P lan t attached to substratum by rh izo ids; basal cell cylindrical in the low er part, about 150-300//, x 5 0 - 7 0 /a but irregu larly and copiously branched in the upper parts, the branches o ften only 10/a in d iam eter; sterile hairs w ere seldom p resen t; sporangia sessile on the branches, cylindrical, o ften curved, rounded at the apex, but truncate a t the base, 8 0 - 1 4 0 /a x 1 8 - 3 5 /a, pro liferating by successive sporangia form ed inside the old o n es; resting spores


F i g . i o .— A , B la s to c la d ia p r o l i f e r a von M inden . A , Y oungp lan t show ing i r re g u la r b ranch ing and sessile {sporangia.X io o . B , P o r tio n o f th a iiu s show ing p ro li fe ra tin g sporang ia. X 2 70 . C» B la s to c la d ia g lo b o s a K anouse.P la n ts bear in g sporang ia and s ter ile ha irs . X 10 0 .

not observed ; the "plants, generally, are sm aller than those ofB . P ringsheim ii, and the lpbes o r branches are no t swollen at the apex as in B . Pringsheintdi except in one or two cases.


S parrow (1936) figures p lants of B . pro lifera in which the heads are definitely rounded. In the p lants exam ined here the sporangia w ere o ften so densly crow ded around the top of the p lant, th a t the na tu re of the sporangia could not be seen, but it gave the appearance o f a head sim ilar to th a t o f B . P ringsheim ii. W hen the p lants w ere dissected the crow ded, irregu lar branching w ere observed. P ro life ra ting sporangia w ere abundant in the older plants.

B l z s t o c l z d i z g r z c i l i s K anouse.A m er . Jou r . B o t., 14, p. 300, p l. 33, f ig s . 14-16, 1927.

B l z s t o c l z d i z r z m o s z var. l u x u r i z n s .K anouse.M ich ig a n A ca d . S c i. A r ts and L e t te r s : 105, p l. 1, fig . 1.

Saprophytic on apples, rose hips, and banana, Y a rra R iver a t R udder G range, collected 10th N ovem ber, 1935. T h is species occurred in the same pustu les as Blastocladia n.sp., Blastocladia gracilis, and Gon’apodya siliquaefo rm is. P lan t attached to- substra tum by w eakly-developed rh izoids, w hole p lan t 600-950/i in ie n g th ; basal cell cylindrical, slender, about 20—4-5/̂ in d iam eter, branched in a racemose to sub-dichotom ous m anner, hyaline, sm ooth, wall th in— 2-4/i. S terile hairs w ith bulbous base noted on some p lants (P ia te X., 3 ) ; zoosporangia borne in a racem ose or corymbose fashion, long cylindrical, 120-220/x x 23-34/i (F ig . 11, A ) ; resting spores usually term inal on the branches, oval to spherical w ith a truncate base, 45-60/i x 25-40/i (F ig . 11, B ) . ' Th is p lan t is very sim ilar to the species described by K anouse, except in the slightly larger size of the sporangia and the presence o f sterile hairs on some plants, also the p lants are som etim es m ore branched than K anouse suggests— a, featu re also observed by L und (1934).

B l z s t o c l z d i z r z m o s z T hax te r.B o t . G az. 21, 50, p l. 3, f ig s. U - 1 6 , 1896.

Saprophytic on banana fru its , Y a rra R iver at R udder Grange, collected 10th Novem ber, 1935; th is fungus occurred in pustules associated w ith Blastocladia . gracilis; p lant attached to substratum by poorly developed rhizoids, w hole p lan t 500-700/m in leng th ; basal cell cylindrical, 15-33/i in d iam eter, copiously and irregu larly branched in the upper parts, the finer branches which bear the reproductive organs term inally , hav ing a diam eter of. only 5 -1 4 /t; wall th in , sm ooth, hyaline, the whole p lan t having a very delicate appearance; zoosporangia oval, slightly narrow ed tow ards the base, 27-40/x x ‘7 -13 /i; resting spores H u n tly rounded a t the apex, narrow ing tow ards the base, 18-25/i x 11-13/i (F ig .

T he p lants seem to be ta ller than those described by T hax te r, bu t from its- delicate appearance, and-,small size of the sporangia and resting spores;-it is unm istakably B. ramosa. ■ - T he size' o f the

sporangia and resting spores agrees fa ir ly well w ith those- of T h ax te r, 1 8 9 6 z ) ' and of Sparrow ( 1 9 3 2 ) , but differs greatly--from those of ano ther collection described by Sparrow recently, in which the reproductive organs are much larger.

S tu d ies on A iistra lian A qua tic Phycom ycetes. 227

F i g . i i .— A , B , B la s to c la d ia g r a c i l is . • C, B la s to c la d ia a s p e rg i l lo id e s n . -sp. - D-G, B la s t o c la d ia a u g u s ta L und . H , X, B la s to c la d ia r a m o s a T h a x te r. A , P la n ts w ith sporang ia (a ) X 3 0 . B , P la n ts w ith res tin g spores (6 ) X 3 0 . C, P la n ts w ith sporang ia X 3 0 . T), E , F , P la n ts bear in g sporang ia (a ) and re s ting spores (b)X 3 0 . G, R estin g spore. X 2 0 5 . H , P o rtio n o f p lan t w ith sporang ia. X 9 0 .I , P a r t of p lan t show ing spo rang ia (a ) and re s tin g spores (b ) X 2 0 5.

B l z s t o c l z d i z z u g u s t z A. Lund.M em o ires de l ’A ca d . R o y . d es S c . arid L e t t , d e D an em ark ,.

C op en h agu e, p . 44, f ig . 21, 1934.

Saprophytic on rose hips, lake at U niversity , collected- 5th A pril, 1935; on rose hips in lake at Botanic G ardens, collected 23rd M ay, 1935; the small w hitish pustu les in w hich th is form was found consisted either entire ly of this, species or occasionally contained a few plants of B . P ringsheim ii.

' P lan t attached to substratum by th in branching rhizoids, o ften copiously developed; whole p lant very delicate and hyaline, but in a few plants observed, the contents w ere qu ite bro w n ish ; plants 160-500//, h igh ; basal cell slender, cylindrical, 10-35/t in d iam eter and Usually branched in the upper parts— occasionally un branched plants are obtained. T he tips of the branches on which the reproductive organs are borne are riot swollen and globose as in B . P ringsheim ii, a lthough sometimes they gradually increase in d iam eter tow ards the tip.- Sterile' ha irs usually present (P la te X ., 5, F igs. 11 D -F ) ; sporangia long cylindrical, narrow , 60-210/x •x 8-25/a ; resting spores oval, light coloured, usually narrow ed tow ards the t i p ; apex of spores rounded or sometimes -beaked (F ig . 11, G ) ; resting spores thin-w alled, and sometimes a suggestion of a p itted inner wall is o b ta in ed ;: size 25—4-5/* x 16^26//,.

In their light colour and thiri outer wall, these spores resemble those '-of B . ramosa, which, according to T h ax te r ( 1 8 9 6 z ) , may properly be spoken of as-conidia. ' A lthough mbsfc o£ the. p lants


showed only the sporangial stage, a few w ere obtained in which resting spores w ere present, and these have not been observed previously. T hey appear to resemble those described fo r B . rostrata (M inden 1912) but the latter, although not exam ined by the au thor, appears to be a larger p lant, d iffering in habit, w ith fusifo rm sporangia, in contrast to the narrow , cylindrical ones of th is species.

B l a s t o c l a d i a g l o b o s a Kanouse.A m e r . Jou r , o f B o ta n y , v o l. X I V . , p. 298, p l. X X X I I , f ig s . 1-4, 1927.

S a p rop h y tic on berr ies o f So lanum pseudo-capsicum , lak e a t M aryb o rou gh , co llec ted 13th F eb ru a ry , 19 3 6 ; p lan ts occu r red in v e ry sm a ll, d en se, w h it ish p u stu les , som etim es assoc ia ted w ith G onapodya siliquaefo rm is but m ore o f te n a lo n e ; p lan t a ttach ed to su b stra tu m b y a fe w rh izo id s, n o t as e x te n s iv e ly d eve lop ed as in B . P r ingsheim ii (F ig . 10, C ) ; basa l ce ll g lo b o se to su b -g lo b o se u su a lly n a r ro w ed to w a rd s base, som etim es lobed , th e lo b es b e in g v e ry sw o lle n ; p lan ts 1 2 0 - 2 0 0 / z h ig h an d up to 2 0 0 / z in d iam eter , th e n arrow part f r o m w h ich th e rh izo id s d ev e lo p h a vin g a d iam eter of 30-50 /a ; w a ll fa ir ly th ick and lam in a te, p ro top lasm dark b row n ish . S te r ile h a irs u su a lly p resen t, s im p le or b r a n c h e d ; sp o ran g ia sess ile , b road ly cy lin d r ica l, '55-130/a x 16-40/a (P la te X ., 2 ) ; rest in g sp o res ov a l to sp h er ica l, 2 5 -3 5 x 27 -32/a and are dark w ith a th ick p it ted w a ll.

The antherid ia l filam ents described by K anouse were not observed. T he hab it o f th is p lan t strongly suggests B . globosa, but the size of the sporangia, and in a g reater degree, th a t of the resting spores, is sm aller than those described by K anouse, and also by Sparrow (1936).

B l z s t o c l z d i z z s p e r g i l l o i d e s n . s p .

Saprophytic on rose hips, lake at U n iversity , collected 5th A pril, 1935; on apples at Y a rra R iver, R udder G range, collected 10th Novem ber, 1935; p lants occurred in small, w hitish pustules, e ither alone o r associated w ith B . gracilis.

P lan t attached to substratum by m eans of a few fa irly stout rhizoids— up to 5 /a in d iam eter; p lant consists of a single-cell, cylindrical in the low er part 2 5 - 4 5 /a in d iam eter, bu t expanded a t the apex to fo rm a swollen head—6 0 - 8 5 /a in d iam eter . ( P late X ., 1, 2 , Fig. 11 C ) ; branched fo rm s w ere not observed; p lants 1 4 0 - 2 0 0 /a high, dark in colour due to dark brow n protoplasm , wall o f basal cell fa irly th ick , 3 .5 - 5 /a ; sterile ha irs usually present, only 1 - 2 / a in d iam ete r; sporangia long, narrow , cylindrical, ar is ing from the swollen head, 8 5 -1 5 0 x 8 - 1 5 /a . Zoospores em erge th rough a term inal opening; resting .spores no t observed,

T he species is characterized by its unbranched basal cell w ith a swollen head bearing long th in sporangia, the whole plant hav ing a dark brow n colour. T he sporang ia in fo rm and size

a re sim ilar to those of B . augusta L und., although slightly narrow er, but the la tte r species has a branched basal cell usually, the branches not being swollen at the apex, and the w hole p lant is delicate and hyaline in appearance. Both species w ere found on the same fru it but not in the same pustule. •

Latin description :—

B l z s t o c l z d i z z s p e r g i l l o i d e s ( n . s p . ) .

P lan ta paucis filamen-tis rhizoideis crassis substrato a ff ixa; cellula basali simplici, parte iiiferiore cylindrica d iam etro 25^45//., capite inflato, d iam etro 60-85/*; a ltitud ine 140-260//., colore atro fusco, m uris cellulae 3.5-5/* crassis. A dsunt plerum que capilli steriles d iam etro 1-2/x. Sporangia longa angusta cylindrica 85-150/1 x 8 -15jx. O osporae ignotae.

S um m a r y .

1. A quatic fungi have been collected from d ifferent localities. T he m ethods of collection- w ere those recom m ended by o ther w orkers— placing traps contain ing baits in the pondS or stream s, although additional baits have been used.

A fte r fo u r to six weeks, the baits were brought into the laboratory , exam ined, and when possible, pure cu ltures o f the fungi w ere obtained.

2. T w enty species w ere exam ined, only one o f which, namely, Saprolegnia fe ra x , has previously been described from A ustralia.

3. A n account is given of the system atic and physiological characteristics of these species, including experiments regard ing the fo rm ation of the sexual organs in various media.

4. The species recorded a r e :—Sapro legnia fe ra x (G ru ith ) T huret,,A ch lya conspicua Coker,A ch lya americana var. m egasperm a var. nov.,A ch lya oblongata de Bary,

A ch lya sp. ?,D ictyuchus sp.,D ictyuchus m onosporus Leitgeb.,P y th iu m p to l ife ru m de B a ry , 'P yth iom orpha gonapodioides H . E . Petersen,.Saprom yces R e in sh ii■ (S ch ro e ter) F ritsch ,,

R h ip id iu m am ericanum T hax ter,G onapodya siliquaeform is (R einsch .) T hax ter,

G onapodya po lym orpha T hax ter,Blastocladia P ringsheim ii Reinsch,

Stu d ies on A ustra lian A qua tic Phycom ycetes. _ 229


B lastocladia pro life ra V on M inden,Blastocladia gracilis K anouse,Blastocladia ram osa T hax te r,Blastocladia augusta A. Lund,Blastocladia globosa K anouse,Blastocladia aspergillo ides n.sp.

A c k n o w le d g m e n t s .

T hanks are due to P ro fesso r E w art of the B otany School, for m aking available the facilities o f the D epartm ent, and to Dr. M cLennan fo r her help and guidance throughout. I am indebted to Rev. F a th er K elly o f N ew m an College, fo r the L atin diagnosis.

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Stu d ies on A ustra lian A qua tic P hycom ycetes. 231

* T h e se p ap ers w e re n o t ob ta in ab le fo r r e fe ren ce .

232 K ath leen M, Crooks:

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* R f . i n s c h , P . F . 1878. B eob ach tu n ger i fiber e in ig e n eu e S a p ro leg n ia cea e , iiber d ie P a ra s ite n in D e sm id ie n z e l len tind ub er d ie S a c h d k u g e ln in A ch ly a sch la u ch en . J a h r b . W i s s . B o t . , ii., p. 283.

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.............. 1936. A co n tr ib u t io n to ou r k n o w le d g e o f th e aq u a tic P h y c o -m y c e te s o f G rea t B r ita in . J o u r . L in n . S o c . L o n d o n , 1, N o . 334, n. 417.

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R e in sch . B o t . G a s ., x ix ,, p. 45.

-------------. 1895, a. C on tr ib . C ryp t. L ab . H a rv a rd U n iv . 27. N e w orp ecu lia r a q u a tic fu n g i. 1. M o n o b le p h a r is . B o t . G a s ,, x x , , p, 433.

-------------. 1895, b . C on tr ib . C ryp t. L ab. H a rv a rd U n iv . 28. N e w orp ecu lia r aq u a tic fu n g i. 2. G o n o p o d y a F is h e r an d M y r io b le p h a r i s n ov . g e n . . B o t . G a s ., x x . , p. 477.

------------- . 1896, a. C on tr ib . C ryp t. L ab . H a rv a r d U n iv . 30. N e w orp ecu lia r aq u a tic fu n g i. 3. B la s to c la d ia . B o t , G a s ,, x x i . , p. 45.

------------ 1896, b. C on tr ib . C fy p t . L ab . H a rv a r d U n iV i 35. N e w o rp ecu lia r aq u a tic fu n g i. 4.. R h ip id iu m S a p r o m y c e s and A r a io s p o r a

• n o v . gen . B o t . G a s ., x x i . , p. 317.

E x p la n a t io n o f P la t e s .

F i g . 1 .— B la s to c la d ia a s p e r g i l lo id e s n . sp . T w o p lan ts w ith sp o ra n g ia . X 150,

F ig . 2 .— B la s to c la d ia g lo b o s a K a n o u se . P la n t sh o w in g sw o lle n b a sa l-ce ll

w ith rh izo id a l sy s tem an d sp o ran g ia . X 450.

F i g . 3.— B la s to c la d ia g r a c i l i s K a n o u se . T h a l lu s w ith sp o ra n g ia . X 100.

F ig . 4 .— B la s to c la d ia a s p e r g i l lo id e s n . sp. A p e x o f th a llu s sh o w in g sw o lle n h ead w ith Sporang ia . X 620.

F ig . 5.-—B la s to c la d ia a n g u s ta L u fid . U p p e r p a r t o f th a l lu s S h o w in g b ra n ch es — n o t sw o lle n a t th e a p ex . X 620.

F i g . 6 .— R h ip id iu m d m e r ic a n u m T h a x te r . T h a l lu s b ea r in g f i la m en ts . X 450.

F i g s . 7, 8 .— S a p r o m y c e s R e in s c k i i (S c h r o e te r ) . F r i tsc h . P la n ts w ith sp o ra n g ia . X 100.

* T h e se pap ers w e re n o t ob ta in ab le fo r re fe ren ce .

P r o c . R o y . S o c . V i c t o r i z , 49 (2 ), 1937. P l z t e X.

Aquatic Phycomycetes.

[Page 233.]

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Transcript of [R ead 12th N ovem ber, 1936; issued separately 19th -July ... · The present writer gives an...