Untitled


356 REINWARDTIA [VOL. 7

Phoebe nitida BL; cf. Kostermans, I.e. 1292 no 114. — Typus: Praetorius s.n.,
Palembang, ster. (L).

Phoebe lamponga Miq.; cf. Kostermans, I.e. 1284 no 84. — Typus: 4.222 H. B.
(U), Tarabangi, Lampongs, "Kajoe Helah (Hoeroe mehrang)".

Phoebe sumatrana Miq.; cf. Kostermans, I.e. 1302 no 165a. — Typus: Junghuhn
s.n., Sumatra, Hochankola, ster. (L, U) et Junghuhn s.n., herb, de Vriese, leaves with
large, hairy galls, Battak region, Sumatra (L, 2 sheets, U).

Ocotea excelsa Bl., p.p., no collector, no number, ster. (L).
Laurus longifolia Reinw. ex Blume; cf. Kostermans, I.e. 649 no 214d. — Typus:

two loose leaves (L).
Laurus gewmdflora Reinw. ex Blume (nomen) ; cf. Kostermans, I.e. 628 no 156b

(non 156a). — Typus: Reinwardt 831 (L), "Huru hiries", Java, fl. (in Leiden the
number 831 appears also on one type sheet of Ocotea declinata BL, marked "Huru leu-
ur", Salak, Dec. 1822).

Persea rimosa Zoll. ex Meissn. (non Machilus riinosa Bl.) ; cf. Kostermans, I.e.
1253 no 250, quoad Zollinger 13484 (L).

REINWARDTIA

Published by Herbarium Bogoriense, Bogor, Indonesia
Volume 7, Part 4, pp. 357—374 (1968)

THE HYPHOMYCETE GENUS DACTYLARIA SACC.

MIEN A. RIFAI *)

SUMMARY

An emended delimitation of the genus Dactylaria Sacc. is proposed and
the two accepted species, which are non-predaceous and dematiaceous,
are redescribed and illustrated. The affinity of many nematode-trapping
species currently classified in Dactylaria with the didymosporous genera
Arthrobotrys Corda, Candelabrella Rifai & R. C. Cooke and Genicularia,
Rifai & R. C. Cooke is discussed and the scopes of the latter genera are
enlarged, and consequently several new combinations are made.

INTRODUCTION

In an earlier attempt to find the appropriate genera for some species
of didymosporous nematode-trapping Hyphomycetes of the Trichothecium-
Arthrobotrys complex (Rifai & Cooke, 1966), several phragmosporous
genera have been seriously considered. Of these the genus Dactylaria
Sacc. has received a special attention, especially because Soprunov (1958)
transferred Arthrobotrys dactyloides Drechsler (1937) to it. This dispo-
sition was tentatively rejected by Rifai & Cooke (1966) because they
believed that the affinity of A. dactyloides with A. superba Corda, the
type species of the genus Arthrobotrys Corda, was sufficiently close to
warrant their inclusion in one genus. They further pointed out that the
limits of the genus Dactylaria was not yet fully understood. The result of
the subsequent study on the taxonomy of Dactylaria reported below does
not only prove that the retention of A. dactyloides in Arthrobotrys is fully
justified, but also shows that the majority of species now included in
Dactylaria cannot be retained in this genus any longer.

In 1877 Saccardo described and illustrated Acrothecium purpurellum
Sacc, a purplish phragmosporous hyphomycete which was found growing
saprophytically on a piece of decayed Quercus wood in Italy. Based on this
species the new genus Dactylaria was created by Saccardo (1880) three
years later, which he described as follows: " Saprophila. Hyphae
fertiles erectae, simplices, apice capitulum conidiorum gerentes; conidia
fusoidea v. clavulata, 2-pluriseptata " In compiling the fourth

*) Herbarium Bogoriense, Bogor (Java), Indonesia.

— 357 —



358 R E I N W A R D T I A [VOL. 7

volume of Sylloge Fungorum Saccardo (1886) transferred four more phrag-
mosporous species to this genus, which he placed in the family Mucedineae
(= the Moniliaceae). Since then all species with similar kind of conidia
which cannot be ranged in Arthrobotrys, Dactyllela Grove or Monacro-
sporium Oud. have been assigned to Dactylaria. In a long series of excellent
papers Drechsler (1937, 1940, 1944, 1950) described and illustrated in great
details the morphology and biology of several species of Dactylaria. Un-
fortunately, however, the morphological charaters and the biology of the
type species of this genus is still imperfectly known.

While investigating the ecology of microfungi colonizing forest leaf
litter in Great Britain, Hering (1965, 1965a) came across with an interest-
ing hyphomycete growing on decaying leaves of Quercus which had been
in the ground for over six months. Dr. M. B. Ellis of the Commonwealth
Mycological Institute (Kew) identified this fungus as Dactylaria purpurel-
la (Sacc.) Sacc.; he (pers. comm., 1967) has also compared this British
collection with the original specimen of the latter and found them to be
identical. A study of an isolate of this species kindly supplied to me by
Dr. T. F. Hering shows that it differs considerably from almost all species
currently classified as Dactylaria, so that it has been found necessary to
emend the limits of this genus.

Dactylaria purpurella, which is a non-predaceous species, markedly
differs from the nematode-trapping species of Dactylaria in its macrosco-
pic as well as its microscopic characters. In culture D. purpurella forms
compact, fluffy or thickly cottony colonies with a very slow growth r a t e ;
they only attain a 19 mm diameter after 12 days at 25°C (Hering, 1965a).
These colonies are greyish brown to dark brown because most of their
mycelia are composed of distinctly brown walled hyphae. When seen from
the reverse of the Petri-dish these colonies often appear almost opaque.
In contrast colonies of the nematophagous group of species have a much
faster growth rate and are capable of covering a 90 mm diameter Petri-dish
in about one or two weeks at 25 °C. Their colonies are translucent and
smooth surfaced because they do not normally form aerial hyphae and the
network of their creeping hyphae are thin and sparse. They are whitish
or pale pinkish and their individual hyphae are colourless; the colonies do
not also discolour their media. The difference in the cultural characters
of D, purpurella and the other species of Dactylaria is such t h a t those
who have seen the colonies of the two groups side by side would certainly
classify them in different genera. Although in recent years the significance
of the hyphal pigmentation as a major taxonomic evidence has occasionally
been doubted (Barron, 1962, 1964; Rifai, 1964), it is worth noting here

1968] RIFAI: The genus Dactylaria 359

t h a t should Saccardo (1877, 1877a, 1880, 1886) see the colonies of D.
purpurella in culture, undoubtedly he would have put the genus Dactylaria
in the Dematiaceae and not in the Moniliaceae.

The habit of conidiophores of D. purpurella shows also some dif-
ferences from t h a t of the nematode-trapping series. As has been well
known, in the latter group of species the conidiophores are usually long,
erect, regularly septate, generally straight or sometimes jointed, and nor-
mally they arise from the creeping or submerged hyphae. Depending upon
the species the apices of the conidiophores of the nematode-trapping
species of Dactylaria may become slightly swollen and bear numerous small
denticles, or they may become simply geniculate, or form a candelabrum-
like branching system to which their obovoid or ellipsoid conidia are at-
tached. On the other hand the conidiophores of D. purpurella are rather
short, frequently and irregularly septate; in colonies grown on artificial
media these conidiophores normally arise from the aerial hyphae and are
straight or curved, whereas their apices become irregularly dilated and
bear a few denticles which produce narrowly subcylindrical or subclavate
conidia.

Cooke & Satchuthananthavale (1966) considered that D. purpurella
and the nematode-trapping species of Dactylaria could be placed in one
genus, but the cultural and morphological characters enumerated above
do not seem to support their view. In fact among the many species which
in the last thirty years have been assigned to Dactylaria there is only one
species that can be accepted as a member of the genus Dactylaria, namely
the Indian species Dactylaria fulva Roy & Gujarati (1965), which is also
a non-predaceous fungus.

It is very likely t h a t Diplorhinotrichum juncicola MacGarvie (1965)
belongs also to Dactylaria as this genus is understood here, but I have not
seen this species. MacGarvie (1965), who described the cultural behaviour
of this species very briefly, stated that it had a slow growing colony which
at first appeared white, becoming light grey and finally dark grey towards
its centre and produced short, flask-shaped, subcapitate and pigmented coni-
diophores and 1-septate subcylindrical conidia. The shape of the sporo-
genous cell of Diplorhinotrichum juncicola seems to be basically different
from that of the two species of the genus Diplorhinotrichum von Hohnel
accepted by Hughes (1951), namely Diplorhinotrichum candidulum von
Hohnel and Diplorhinotrichum africanum Hughes. Papendorf (1967) sug-
gested t h a t Diplorhinotrichum might not be generically distinct from
Dactylaria but the cylindrical shape of its sporogenous cells appears to be
acceptable as the leading character which distinguish this genus from



360 R E I N W A R D T I A [VOL. 7

Dactylaria. Unfortunately these two hyaline species of Diplorhinotrichum
have not been grown in culture so t h a t it has not been possible to assess
the taxonomic value of their cultural charaters.

The generic name Dactylaria has been much used in mycological and
phytopathological literature for the nematophagous species that its con-
servation might be preferred to preserve current usage. Since it will be
shown later that the commonly accepted circumscription of Dactylaria
(Clements & Shear, 1931; Drechsler, 1937, 1940, 1944, 1950; Dollfus, 1946;
Bessey, 1950; Soprunov, 1958; Barnett, 1960; Cooke & Godfrey, 1964;
Cooke & Satchuthananthavale, 1966) is a heterogeneous one and because
its species can be referred to some existing genera, the conservation of
Dactylaria which will exclude its original type species is unnecessary.

THE TAXONOMY OF DACTYLARIA

DACTYLARIA Sacc. emend. Rifai

Dactylaria Sacc. in Michelia 2: 20. 1880; Syll. Fung. 4: 194. 1886; Lindau apud
Eng-ler & Prantl, Nat. Piflanzenfam. I, 1: 448. 1900; Clem. & Shear, Gen. Fung.: 207.
1931; Bessey, Morph. Tax. Fung.: 615. 1950; Barnett, 111. Gen. Imp. Fung.: 70. 1960.

In pure culture colonies grow very slowly, restricted, fluffy or thickly
cottony, compact, grey to dark brown; colour of reverse also brown or
dark brown to almost opaque. Mycelium composed of compact network of
septate, much branched, subhyaline to brown coloured and smooth walled
hyphae, with abundant but paler aerial hyphae. Conidiophores arise from
aerial or creeping hyphae, generally short, straight or flexuous, frequently
septate, smooth walled, occasionally irregularly branched, pale brown to
suhyaline below, paler towards the dilated subcapitate apex; the latter
bears a few small conical truncate conidial pegs which arise through
repeated subapical proliferations, a process which makes the conidiophores
very slightly elongate and dilated at the apex; sometimes a renewed growth
of conidiophore occurs after the formation of the first group of conidial
pegs and a second group will be formed at some distance above the first.
Conidia arise singly as blown out ends of the conidiophores, blastogenous,
hyaline to pale pink, 1—many-septate, smooth walled, subcylindrical or
narrowly subclavate, distally rounded but truncate at the base.

The slow growing, restricted, fluffy, compact, dematiaceous colonies,
t h e generally rather short conidiophores with subcapitate apices which
produce subcylindrical conidia and the non-predaceous habit serve to
distinguish this genus from other radulasporous nematode-trapping genera
of Hyphomycetes, to which Dactylaria is only remotely related.

HABITAT: mostly saprophytic in soil or on decayed vegerable matter.
TYPE SPECIES: Acrothecium purpurellum Sacc.

1968] RIFAI: The genus Dactylaria 36t

KEY TO SPECIES OF DACTYLARIA

la. [Parasitic on Juncus; conidiophores bulbous, conidia at maturity regularly 1-septate
; "Diplorhinotrichum" juneicola]

b. Saprophytic; conidiophores not bulbous, conidia at maturity 1—3-septate '. 2
2a. Colonies grey brown to dark brown; conidia 1—3-septate, 17.2—26.3(—30) X 2.7—

4 n. Italy, Great Britain Dactylaria.purpurella
b. Colonies at maturity grey to brownish grey; conidia 1—2-septate, 13.6—23.9

(—50) x 1.5—2.2(—3)n. India Dactylaria fulva

< DACTYLARIA PURPURELLA (Sacc.) Sacc.

Acrothecium purpurellum Sacc, Fung. Ital. Autograph. Del., Fasc. I-IV: t.8.
May 1877; Fung. Ital. Autograph. Del. Comm.: 75. June 1877. •— Dactylaria purpurella
(Sacc.) Sacc. in Midhelia 2: 20. 1880.

In pure culture on corn-meal agar colonies grow very slowly (only
attain a 60 mm diameter after 6 weeks at room temperature of 25 °C),
restricted, at first smooth surfaced but soon becoming compactly fluffy
or cottony from the formation of abundant interwoven aerial hyphae which
usually form raised thick cushions, indistinctly zonate, greyish brown to
blackish brown or rarely dark olive brown, often tinge with some shades of
pink; colour of reverse also dark blackish or reddish brown, often almost
opaque, whereas the media are discoloured by a red brown pigment; ac-
cording to Hering (1965a) on its natural substrates colonies of this species
form conspicuous white patches. Mycelium composed of a compact network
of profusely branched hyphae which are subhyaline, pale brown to dark
brown, 1.3—4µ. diameter, frequently septate and their cells sometimes
slightly inflated or barrel shaped, their walls rather thick, smooth or very
rarely minutely echinulate; the aerial hyphae mostly composed of lighter
coloured elements, but some of these may become tightly aggregated with
each other to form irregular capillitium-like hyphal cords which are dark
blackish brown and appear shining under the reflected light. Conidio-
phores arise singly as lateral branches of the aerial or sometimes also, of
the creeping hyphae, straight or gently curved, simple or irregularly
branched, pale brown below and becoming paler towards the subhyaline
apex, smooth walled, many septate, 2—4.8 µ. diameter by 20—100 µ long
(but on its natural substrate the length of the conidiophore may reach
500µ according to Hering, 1965a) ; the apices of the conidiophores ulti-
mately become slightly dilated and bear a few pronounced, flat-topped,
irregularly disposed conidial pegs or scars which arise by successive sub-
apical proliferations of the conidiophores; through a renewed growth
another group of conidial pegs may be formed at a higher level. Conidia
arise singly and successively as blown out ends of the conidiophores, hyaline
to subhyaline, smooth walled, subclavate to subcylindrical, rounded at the
apex, base obconical and truncate, mostly 1-septate but a few are 3-septate,
17.2—26.3(—30) X 2.7—4µ; Saccardo (1877a, 1886) and Hering (1965a)



362 R E I N W A R D T I A [VOL. 7

Fig. 1. Dactylaria purpurella: A, eonidiophores and conidia (from T.F. Hering BC/
1040, living culture) ; B, conidia (from T.F. Hering BCJ10AO, on leaf litter).

1968] RlPAI: The genus Dactylaria 363

gave the range of the spore size J.S being 20—25 x 4 \i. and 16—23 X 3 5
p. respectively (fig. 1).

HABITAT: on decayed leaves and wood of Quercus.
DISTRIBUTION: Italy (type locality) and the Great Britain.
ILLUSTRATIONS: Saccardo, Fung. Ital. Autograph. Del., Fasc. 1-1V:

t.8. 1877; Lindau apud Engler & Prantl, Nat. Pflanzenfam. I, 1: 449, fig.
232J. 1900; Hering in Trans. Br. mycol. Soc. 48: 665, fig. 5. 1965; Cooke &
Satchuthananthavale in Trans. Br. mycol. Soc. 49: 30, fig. 2E. 1966.

GREAT BRITAIN. On Querous leaf litter, Merlewood, Lancashire, 8 January
1963, T.F. Hering BC/lOiO (IMI 101627) ; a culture was isolated from the latter and
a dried culture on corn-meal agar is preserved in BO.

DACTYLARIA FULVA Roy & Gujarati :

Dactylaria fulva Roy & Gujarati in Lloydia 28: 53. 1965.

The following description is drawn from the dried type culture isolate
and supplemented with Roy & Gujarati's (1965) original account: In pure
culture on potato-dextrose-agar colonies grow very slowly, only reach a
9 mm diameter after one week at 25 °C, restricted, fluffy or compactly
cottony, at first dirty white, becoming grey to brownish grey at maturity;
colour of reverse also brownish at maturity. Mycelium made up of intricate
network of hyphae which are subhyaline to yellowish brown or brown, 1—3
µ diameter, smooth walled, septate, much branched, sometimes aggregated
with each other to form hyphal cords. Chlamydospores present in older
cultures, subglobose to elongated, mostly intercalary or arise laterally to
the hyphae, thick walled, dark reddish brown and measure 5.4—
36.4 x 5.4—21.8 µ. according to Roy & Gujarati. Conidiophores mostly
arise as lateral branches of the hyphae, straight or flexuous, irregular in
length, may be very short and only 4µ, long but mostly vary 'from 6 to 50
µ, long by 2—3µ. diameter, septate, subhyaline to pale yellowish brown
below; the apex of conidiophore hyaline, subhyaline or yellowish, distinctly
dilated to 2.7—4.5 µ. diameter, bearing numerous irregularly disposed,
flat-topped conidial pegs to which conidia formerly attached; a conidium-
producing apex may be formed at a higher level through a renewed growth
of the conidiophores. Conidia arise singly as blown out ends of the conidio-
phores, smooth walled, hyaline, subcylindrical to almost subclavate, only
slightly or hardly narrower at the base, at maturity generally regularly
2-septate, 13.6—23.9 X 1.5—2.2 µ.; Roy & Gujarati gave the range of the
spore size as being 13—50 x 1.5—3µ (fig. 2).

HABITAT : in rhizosphere of Dichantium annulatum.
DISTRIBUTION: India.
ILLUSTRATION: Roy & Gujarati in Lloydia 28: 54, fig. 1—11. 1965.
INDIA. Isolated from decayed roots of Dichantium annulatum, Botanical Garden,

Banaras Hindu University, Varanasi, U. P., 1963, S. Gujarati (IMI 104480, isotypus).



864 R E I N W A R D T I A [VOL. 7

A REASSESSMENT OF THE EXCLUDED SPECIES

In recent years it has been more or less generally accepted that the
taxonomic value of the spore septation in delimiting genera of some
groups of fungi has been given an undue emphasis. In many cases modern
authors have ignored this character and classified numerous species of
Hyphomycetes with various types of spore septation in one genus, which
in Saccardoan (1880, 1886) system of classification will require two, three
or four separate genera. Boedijn & Reitsma (1950) pointed out t h a t the
genera Cylindrocladium Morgan and Candelospora Rea & Hawley were
separated form each other solely by the formation of 3—many-septate
conidia and 1-septate conidia respectively; since all their other morpho-
logical and cultural characters are basically identical, they justifiably
merged the two genera. Hughes (1958) erected the genus Scheleobrachea
Hughes, which has been shown to represent a later synonym of the genus
Pithoniyces Berk. & Br. by Ellis (1960) ; in both Hughes' and Ellis' accounts

Fig. 2. Dactylaria fulva: conidiophores and conidia (from S. Gujarati).

1968] RIFAI: The genus Dactylaria 365

species with 1-celled, 1-—3-septate conidia and even those with muriform
conidia are accommodated in the same genus. Endophragmia Maire &
Duvernoy now does not only contain species with phragmosporous conidia
but also a few species which have 1-septate conidia (Ellis, 1959). Sutton
& Pirozinsky (1965) united Taeniophora Karst, and Alysisporium Pyronel
with Pragmotrichum Kunze & Schmidt because of their overall similarity
except in the possession of phragmosporous and muriform conidia re-
spectively ; similarly the genus Hansfordiella Hughes also embraces species
with phragmosporous and muriform conidia (Hughes, 1951). Madelin (1966)
correctly transferred Botrytis acridiorum Trabut to the traditionally didy-
mosporous genus Trichothecium Link ex Fr., although this species has
1-celled, 1—3-septate conidia; because of this the taxonomic relation of
Cylindrophora apiculata Tubaki and Trichothecium roseum (Pers.) Link
ex S. F. Gray should now be reconsidered (cf. Tubaki, 1954; Rifai & Cooke,
1966). In the Ascomycetes we find that the helotiaceous genus Hymeno-
scyphus S. F. Gray emend. Dennis (= Helotium auct., non Helotium Tode
ex Lehman, which is agaricaceous) has species with both 1-celled as well as
1—many-septate ascospores (Dennis, 1964) ; the same situation can also be
found in the hyaloscyphaceous genus Dasyscyphus S. F. Gray (Dennis,
1960). In the pyrenomycete genus Massarina Sacc. there are many species
with 1-septate ascospores besides those with phragmosporous ones (Bose,
1961; Muller & von Arx, 1962).

Although it must be admitted that a character widely accepted as
having a generic value in one group will not always be of equal importance
in a different context, it seems t h a t the unsuitability of the spore septation
to serve as a major taxonomic evidence in delimiting the genera of nema-
tode-trapping Hyphomycetes is manifest. The currently accepted conception
t h a t Arthrobotrys is a didymosporous genus with Dactylaria sensu lato
serving as its phragmosporous counterpart has not been consistently put
into practise: though definitely exceptional, conidia of Arthrobotrys super-
ba and A. dactyloides (Drechsler, 1937) may have more than one septum.
This anomaly, however, has always been played down largely because
there is no doubt whatsover that these species are good members of
Arthrobotrys. It follows t h a t it would not be possible to use the number
of spore septation as a key character to separate Arthrobotrys from Dacty-
laria sensu lato because of the existence of these intermediate forms.
The taxonomic value of the spore septations is further weakened by the
fact t h a t in nematode-trapping Hyphomycetes the types of media appear
to have an effect on the number of septa produced. When grown on a
nematode-infected agar Dactylaria clavispora R. C. Cooke produces conidia



366 BEINWAEDTIA [VOL. 7

which are " 1—2-septate, about twice as many having one septum
as two " but on corn-meal agar the conidia of this species become
1—3-septate, the majority of which have " 2 septa, about three
times as many with 1 or 3 septa " (Cooke, 1964). A similar
situation can also be found in Dactylaria eudermata Drechsler (1950).

As has been hinted earlier, the conidiophores of the nematode-trapping
species to be excluded from Dactylaria can be grouped into at least three
types, each of which corresponds to the conidiophore type found in the
didymosporous genera Arthrobotrys, Genicularia Rifai & R. C. Cooke and
Candelabrella Rifai & R. C. Cooke respectively. The only character which
these species have in common with each other is the phragmosporous nature
of their conidia, which is also the only character which so far has prevented
them from being referred to the three didymosporous genera. From the
above consideration it would be obvious, however, that for the nematode-
trapping species the number of spore septation should not be accorded a
generic value. Therefore it would be fully justifiable to distribute these
species among the genera Arthrobotrys, Candelabrella and Genicularia and
at the same time to enlarge the scopes of these originally didymosporous
genera to enable them to accommodate the phragmosporous species as
well. The other alternative would be the proliferation of newly described
phragmosporous genera for the reception of the numerous species current-
ly classified as members of Dactylaria, but this is wholly undesirable
because it will only make the ralationship of these species becomes more
obscure than as it is.

GENICULARIA Rifai & R. C. Cooke emend. Rifai
Genicularia, Rifai & R. C. Cooke in Trans. Br. mycol. Soc. 49: 153. 1966.

In pure culture colonies grow quite rapidly, effused, pale pink to
whitish. Mycelium composed of septate, hyaline, smooth walled, branched
hyphae which mostly form thin and sparse mat over the surface of the
media, with scanty aerial growth. Conidiophores arise from creeping or
submerged hyphae, hyaline, smooth walled, septate, erect or ascending,
at first straight, becoming geniculate or flexuous, elongating, sometimes
considerably, by repeated subapical renewal of growth; its apex is blown
out blastogenously to form a conidium and after the first conidium has
been formed a new growing point appears at one side of it and the second
conidium is formed at the new apex, displacing the first conidium to a
lateral position, this process being repeated several times. Conidia short
obpyriform, obpyriform to obvoid-turbinate or broadly fusoid-ellipsoidal,
1—many-septate, the basal cell usually obconical and truncate, hyaline
when viewed singly but appearing pale pinkish-white in mass, smooth
walled.

1968] RIFAI : The genus Dactylaria 367

HABITAT: mostly (? always) capturing and parasitizing nematodes
by means of various kinds of traps in soil or debris.

TYPE SPECIES: Trichothecium cystosporium Duddington.

The species included in this genus by Rifai & Cooke (1966) so far are
didymosporous. As has been indicated above in Geniculairia clavispora
(R. C. Cooke) Rifai, comb. nov. (basionym: Dactylaria clavispora R. C.
Cooke in Trans. Br. mycol. Soc. 47: 307, fig. 1. 1964) the number of spore
septation in each conidium varies from 1, 2 or 3, mostly one or two
depending on the type of media used. As far as the spore septation is
concerned, this species appears to represent an intermediate form between
the didymosporous members of Genicularia and Genicularia psychrophila
(Drechsl.) Rifai, comb. nov. [basionym: Dactylaria psychrophila Drechsl.
in Mycologia 36: 161. 1944. — Monacrosporium psychrophilum (Drechsl.)
R. C. Cooke & Dickinson in Trans. Br. mycol. Soc. 48: 622. 1965], because
the latter has 2—{mostly 3—)4-septate conidia. The type of conidiophore
of G. clavispora and G. psychrophila is similar to that of G. cystosporia
(Duddington) Rifai & R. C. Cooke.

Drechsler (1950) described the conidiophores of Dactylaria eudermata
as follows: " simple conidiophores bear terminally a single coni-
dium much like the simple conidiophores, for example, of Dactylella aphro-
brocha and D. bembicodes, which, indeed, in their dimensions they resemble
rather closely. The fungus likewise puts forth distally branched conidio-
phores that beside producing a spore at the tip of the main hypha bear
additional spores singly on its primary branches and also on its secondary
branches if such are present. As the branches are often rangy, sometimes
exceeding 50 JJ, in length, the conidia produced plurally are in many in-
stances attached at generous distances from one another " There
is no doubt that this species (fig. 3) likewise should be known as Geni-
cularia eudermata (Drechsl.) Rifai, comb. nov. [basionym: Dactylaria
eudermata Drechsl. in Mycologia 42: 40. 1950. — Monacrosporium euder-
matum (Drechsl.) Subram. in J. Indian bot. Soc. 42: 293. 1963]. As
Cooke & Dickinson (1965) already stated this species is closely related to
Genicularia bogoriensis Rifai, nom. nov. [basionym: Monacrosporium
cystosporum R. C. Cooke & Dickinson in Trans. Br. mycol. Soc. 48: 623.
1965; non Genicularia cystosporia (Duddington) Rifai & R. C. Cooke].
On nematode-free media these two species often produce conidia which
are similar in outline to those of G. perpasta R. C. Cooke apud Rifai & R. C.
Cooke.

Subramanian (1963) and Cooke & Dickinson (1965) placed G. psychro-
phila, G. eudermata and G. bogoriensis in the genus Monacrosporium.





370
REINWARDTIA

[VOL. 7

apparently one or two more species of Dactylaria will have to be included

in the present genus as well.

ARTHROBOTRYS Corda emend. Rifai

Pracht Fl • 43 1839; Sacc. Syll. Fung. 4: 181. 1886; Lindau
Nat.Planzenfam . I, 1: 445. 1900; Clem. . Shear, Gen. Fung.

Fig 4. CandelobreUa haptotyla: comdiophores and conidia
(redrawn from Drechsler, 1950).

1968] RlFAl: The genus Dactylaria 371

206. 1931; Bessey, Morph. Tax. Fung.: 615. 1950; Barnett, 111. Gen. Imp. Fung.: 66.
1960; Rifai & Cooke in Trans. Br. mycol. Soc. 49: 164. 1966.

Didymozoophaga Soprunov & Galiulina in Mikrobiologiya 20: 494. 1951 (sine
diagnose latina).

In pure culture colonies grow rapidly, translucent, watery white to
pale pinkish white, mostly smooth surfaced. Mycelium made up of smooth
walled, septate, branched and hyaline hyphae forming a thin sparse
mycelial mat on the media, typically with scanty aerial growth. Conidio-
phores arise from creeping or submerged hyphae, hyaline, smooth walled,
septate, subulate or subcylindrical, mostly erect and straight but sometimes
jointed, very rarely branched; at maturity conidiophore apex typically
capitate, namely terminated by a swelling which is mostly subglobose or
sometimes irregularly elongated, lobed or clavate and bearing numerous
minute and sterigma-like conidial pegs which previously produce t h e coni-
dia ; sometimes renewed growth of the conidiophore occurs after the
formation of the first conidial head and a second head will be formed at
some distance above the first; the process may be repeated and leads to
the production of several such swellings along the length of the conidio-
phores, which cause the latter to become jointed. Conidia distinctly blasto-
genous, arising singly as blown out ends of the conidiophore and the ends
of the newly formed growing points, narrowly obovoid to oblong-ellipsoidal,
sometimes slightly curved, 1—many-septate, smooth walled, hyaline.

HABITAT: in soil or debris, mostly capturing nematodes by means of
simple or elaborate traps.

TYPE SPECIES: Arthrobotrys superb a Corda.

Besides A. superba and A. dactyloides, there are other normally didy-
mosporous species of Arthrobotrys which occassionally have been observed
to produce conidia with more than 1 septum, such as A. oligospora Fres.
(Drechsler, 1937), A. anchoina Drechsler (1954) and A. longispora Soprunov
(1958; non A. longispora Preuss, 1851). In Arthrobotrys vermicola (R. C.
Cooke & Satchut.) Rifai, comb. nov. (basionym: Dactylaria vermicola R. C.
Cooke & Satchut. in Trans. Br. mycol. Soc. 49: 27. 1966) the conidia pro-
dused have 1 to 3 septa, but mostly have 2 septa.

It is of interest to note t h a t in referring the predominantly 4-septate
spored species Arthrobotrys polycephala (Drechsl.) Rifai, comb. nov.
(basionym: Dactylaria polycephala Drechsl. in Mycologia 29: 530. 1937)
to the genus Dactylaria a remark was made by Drechsler (1937) t h a t this
species " shows such obvious parallelism, both in reproductive
habit and in make up of predaceous apparatus, to the retiary species of
Arthrobotrys t h a t its assignment to another genus, however clearly neces-
sitated by the plural septation of its conidia, cannot be regarded with any



372
R E I N W A R D T I A [VOL. 7

gratification " Since the number of spore septation is no longer
maintained to be of important generic value there is nothing to prevent the
inclusion of this species in Arthrobotrys, especially because the morpho-
logy of its conidiophore and conidium-producing organ is essentially identi-
cal with that of A. superba.

I have not been able to study the developments of the first conidia
of a few more species currently classified as Dactylaria so that their
eventual classification cannot be proposed at the moment. Dactylaria
haptospora Drechsler (1940), however, does not seem to belong to any of
the genera discussed above because of its peculiar sporogenous cells as
well as the unusual morphology of its conidia; ultimately it may be neces-
sary to propose a new genus for it.

A C K N O W L E G D E M E N T S

I would like to thank Dr. T. F. Hering (Loughborough) for kindly
supplying me with a living culture of Dactylaria purpurella and to Dr.
M. B. Ellis (Kew) for generously placing the Commonwealth Mycological
Institute material of Dactylaria at my disposal.

R E F E R E N C E S

BARNETT, H. L. (1960). Illustrated genera of imperfect fungi. Minneapolis.
BAERON, G. L. (1962). Stachybotrys aurantia sp. nov. from soil. In Can. J. Bot. 40:

257—261.
BARRON, G. L. (1984). A note on the relationship between Stachybotrys and Hyalo-

stachybotrys. In Mycologia 56: 313—315.
BESSEY, E. A. (1950). Morphology and taxonomy of fungi. Philadelphia.
BOEDIJN, K. B. & REITSMA, J. (I960). Notes on the genus Cylindrocladium ( F u n g i :

Mucedinacaas). In Reinwardtia 1: 51—60.
BOSE, S. K. (1961). Studies on Massarina Sacc. and related genera. In Phytopath. Zi..

4 1 : 151—213.
CLEMENTS, F. E. & SHEAR, C. L. (1931). The genera of fungi. New York.
CoOKE, R. C. (1964). Dactylaria clavispora,, a new nematode-trapping hyphomycete. In

T r a n s . Br. mycol. Soc. 47: 307—309.
COOKE, R. C. & DICKINSON, C. H. (1965). Nematode-trapping species of Dactylella and

Monacrosporium,. In T r a n s . Br. mycol. Soc. 48: 621-—629.
CoOKE, R. C. & GODFREY, B. E. S. (1964). A key to the nematode-destroying fungi. In

T r a n s . Br. mycol. Soc. 47: 61—74.
COOKE, R. C. & SATCHUTHANANTHAVALE, V (1966). Some nematode-trapping species' of

Dactylaria. In T r a n s Br. mycol. Soc. 49: 27—31.
DENNIS, R. W. G. (1960). British cup-fungi and their allies. London.

1968] RIFAI : The genus Dactylaria 373

DEWNIS, R. W. G. (1964). Remarks on the genus Hyttienoscyphus S. F. Gray, with obser-
vations on sundry species referred by Saccardo and others to the genera
Helotium, Pezizella or Phialea. In Persoonia 3: 29—80.

DOLLFUS, R. P. (1964). Parasites (animaux et vegetaux) des Helminthes. In Encycl.
biol. 27: 1—481. .. : • '

DRECHSLER, C. (1937). Some hyphomycetes t h a t prey on free-living terricolous nema-
todes. In Mycologia 29: 447—552.

DRECHSLER, C. (1940). Three new hyphomycetes preying on free-living terricolous
nematodes. In Mycologia 32: 448—470.

DRECHSLER, C. (1944). Three hyphomycetes t h a t capture nematodes in adhesive network.
In Mycologia 36: 138—171.

DRECHSLER, C. (1950). Several species of Dactylella and Dactylaria t h a t capture free-
living nematodes. In Mycologia 42: 1—79.

DRECHSLER, C. (1954). Some hyphomycetes t h a t capture eelworms in southern states.
In Mycologia 46: 762—782.

E L L I S , M. B. (1959). Clasterosporium and some allied Dematiaceae—Phragmosporae.
II. In Mycol. Pap. 72: 1—75.

E L L I S , M. B. (1960). Dematiaceous Hyphomycetes. I. In Mycol. P a p . 76: 1—36.
HERING, T. F. (1965). Succession of fungi in the litter of Lake District Oakwood. In

Trans. Br. mycol. So. 48: 391—408.
HERING, T. F. (1965a). British Records: 89. Dactylaria purpurella (Sacc.) Sacc. In

T r a n s . Br. mycol. Soc. 48: 666—667.
HUGHES, S. J. (1951). Studies on micro-fungi. XIII. Beltrania, Cerutocladium, Diplorhi-

notrichum and Hansfordiella (gen. nov.). In Mycol. P a p . 47: 1—15.
HUGHES, S. J. (1958). Revisiones Hyphomycetum aliquot cum appendice de nominibus

rejiciendis. In Can. J. Bot. 36: 727—836.
MACGARVIE, Q. D. (1965). Diplorhinotrichum juncioola sp. nov., causing a disease of

Juncus effusu8. In Trans. Br. mycol. Soc. 4 8 : 269—271.
MADELIN, M. F. (1966). Triehothecium acridiorwm (Trabut) comb. nov. on red locusts.

In T r a n s . Br. mycol. Soc. 49: 275—288.
MULLER, E. & VON ARX, J. A. (1962). Die Gattungen der didymosporen Pyrenomyceten.

In Beitr. Kryptogamenfl. Schweiz I I , 2: 1—922.
PAPENDORF, M. C. (1967). Two new genera of soil fungi from South Africa. In T r a n s .

Br. mycol. Soc. 50: 69—75.
RIFAI, M. A. (1964). Stachybotrys bambusicola sp. nov. In Trans. Br. mycol. Soc. 47:

269—272.
RIFAI, M. A. & COOKE, R. C. (1966). Studies on some didymosporous genera of nematode-

t r a p p i n g Hyphomycetes. In Trans. Br. mycol. Soc. 49: 147—168.
ROY, R. Y. & GUJARATI, S. (1965). A new species of Dactylaria from soil. In Lloydia 28:

53—54.
SACCARDO, P. A. (1877). Fungi italici autographice deliniati. Fasc. I—IV. Patavii.
SACCARDO, P. A. (1877a). Fungi italici autographice deliniati. Commentarium. In Mi-

chelia 1: 73—100.
SACCARDO, P. A. (1880). Conspectus generum fungorum italiae inferiorum. In Michelia

2: 1—38.
SACCARDO, P. A. (1886). Sylloge Fungorum 4. Patavii.



S74 R E I N W A R D T I A [VOL. 7

SOPRUNOV, F. F. (1958). [Predaceous fungi — Hyphomycetes and their application in
the control of pathogenic nematodes. Ashkabad. — In Russian].

SUBRAMANIAN, C. V. (1963). Dactylella, Monacrosporium and Dactylina. In J. Indian
bot. Soc. 42: 291—300.

SUTTON, B. C. & PIROZYNSKI, K. A. (1965). Notes on microfungi. II. In Trans. Br. mycol.
Soc. 48: 349—366.

TUBAKI, K. (1954). Studies on the Japanese Hyphomycetes. I. Coprophilous group. In

Nagaoa 4: 1—20.

R E I N W A R D T I A

Published by Herbarium Bogoriense, Bogor, Indonesia
Volume 7, Part 4, pp. 375—381 (1968)

KOSTERMANSINDA RIFAI GENUS NOVUM HYPHOMYCETARUM

MIEN A. RIFAI *)

SUMMARY

The conidial development of Sclerographium magnum Boedijn is described
and illustrated; based on this species the new aleuriosporous genus Koster-
mansinda Rifai is proposed.

The morphology of the conidiophores and conidia of Sclerographium
aterrimum Berk., the type species of the stilbaceous genus Sclerographium
Berk., was described and illustrated in details by Hughes (1951). The
murif orm conidia of this species are radulaspores because they arise blasto-
genously and produced on numerous small denticles on the somewhat
dilated apices of the fasciculated conidiophores, which during the process
of the conidial development elongate slightly by subapical proliferations.
Therefore this genus has been correctly included in the section II of
Hughes' (1953a) experimental system of classification of Hyphomycetes
or in the Radulasporae of Tubaki (1963), Nilsson (1964) and Rifai & Cooke
(1966).

In 1960 Boedijn described Sclerographium, magnum Boedijn, based
on a collection of a fungus which he found growing on the decaying petiole
of a palm species in Bogor Botanic Garden, Java. From Boedijn's descrip-
tion and illustration, as well as from the results of my own observations
on more recent collections of this species which were made from decaying
petioles of several palm species cultivated in Bogor Botanic Garden, it is
evident that Sclerographium magnum has murogenous conidia produced
by simple conidiophores, so that it is an aleuriosporous species which con-
sequently should be referred to the Aleuriosporae or to the section III of
Hughes' system of classification. Although Boedijn (1960) was fully aware
that this species was not at all related to Sclerographium aterrimum, he
nevertheless preferred to place it in the genus Sclerographium because
of the superficial similarity between the two species, and also because of
the artificial nature of the classification of Deuteromycetes; in recent
years, however, substantial evidence are available to show that the system
of classification proposed by Hughes (1953a), which is based on the methods

*) Herbarium Bogoriense, Bogor (Java), Indonesia.

— 375 —


	Rein.Vol.7 Part 4,pp 291-420_Page_34
	Rein.Vol.7 Part 4,pp 291-420_Page_35
	Rein.Vol.7 Part 4,pp 291-420_Page_36
	Rein.Vol.7 Part 4,pp 291-420_Page_37
	Rein.Vol.7 Part 4,pp 291-420_Page_38
	Rein.Vol.7 Part 4,pp 291-420_Page_39
	Rein.Vol.7 Part 4,pp 291-420_Page_40
	Rein.Vol.7 Part 4,pp 291-420_Page_41
	Rein.Vol.7 Part 4,pp 291-420_Page_42
	Rein.Vol.7 Part 4,pp 291-420_Page_43