De HOOG 1972 TThe Genera Beauveria, Isaria, Tritirachium and Acrodontium

THE GENERA BEAUVERIA, ISARIA, TRITIRACHIUM AND ACR...
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Studies in Mycology, No. 1
http://www.cbs.knaw.nl/publications/1001/content_files/content.htm
15 September 1972
The genera Beauveria, Isaria, Tritirachium and Acrodontium gen. nov

G. S. DE HOOG
Centraalbureau voor Schimmelcultures, Baarn.
Summary
The genus Beauveria Vuill. is restricted to three species, of which descriptions and figures are provided: B. bassiana,
B. brongniartii and B. alba. The genus Isaria Fr. is described with two species, I. felina and I. orthopterorum. The
species formerly placed in Tritirachium Limber are arranged in two genera: Tritirachium, based on T. dependens,
characterized by the zig-zag shape of the fertile portions of the conidiogenous cells and the absence of distinct denticles,
and Acrodontium gen. nov., based on Chloridium crateriforme van Beyma, characterized by the straight, denticulate
fertile portions of the conidiogenous cells. Descriptions and figures are presented of two species of Tritirachium and
seven species of Acrodontium. In the latter genus two sections are distinguished: in section Acrodontium the conidia are
formed on solitary conidiogenous cells, in section Grisea, based on Tritirachium heimii (Saccas) Langer. var. griseum
Fassatiov, compound erect or suberect conidiophores are present which are mostly stiff and pigmented. A list of
doubtful and excluded species is given.
Introduction
Taxonomy. Some of the representatives of the genus Beauveria are common insect parasites. They have
been studied frequently, but still some problems of nomenclature exist.
The species now placed in Beauveria were initially known in Europe as Botrytis, while most authors in
America referred to them as Sporotrichum. The genera mentioned were considered to be closely related to
each other. Vuillemin (1912) separated the genus Beauveria from Botrytis on the basis of the geniculate fertile
portions of the conidiogenous cells. Limber (1940) defined the genus Tritirachium mainly on the same character.
Saccas (1948) supposed that Limber might not have known of the existence of Beauveria and considered both
genera as identical. MacLeod (1954) while examining the genus Beauveria found that other good criteria for
distinguishing this genus from Tritirachium were available. After testing the value of taxonomic criteria used
previously, he concluded that in Beauveria only two species could be distinguished.
For the present study fresh isolates, herbarium specimens and numerous [p. 2] strains maintained in the
CBS collection were compared. It appeared to be necessary to use other taxonomic criteria for delimiting the
genera Beauveria, Tritirachium and other genera discussed by Hughes (1953) in section II of his ontogenetic
scheme. Most attention was paid to the conidial production and the ramification of the conidiophores.
Conidial production. In all genera considered here, the conidia arise by the same mechanism. The first
conidium is a terminal swelling formed on the narrowed neck of the conidiogenous cell. The next one is formed
by lateral proliferation and is pushed upwards by sympodial growth (Kendrick, 1972). In some cases elongation
of the tip of the conidiogenous cell ceases after the first conidium is formed, and proliferation takes place at the
base of the first neck. The subsequent branchlets do not exceed the previous ones (Fig. 1a), all conidia are
formed side by side in a limited number, as in Isaria sensu von Arx (1970). In the other genera described in this
paper the conidiogenous cell elongates with the formation of every subsequent conidium. In the species of
Beauveria the second conidium is initiated half-way up the first neck, the next one half-way up the previous one
in another direction, etc. In that way a geniculate rachis with denticles of a width equal to the rachis is formed
(Fig. 1b). In Tritirachium sensu str. proliferation occurs immediately below the previous conidium. A geniculate,
cicatrized (Ellis, 1971) rachis is formed, on which the scars can be traced with difficulty (Fig. 1c). In
Acrodontium gen. nov. subsequent proliferations push the previous ones aside, the rachis becomes straight and
denticulate (Fig. 1d), giving rise to apiculate conidia. In the section Grisea of this genus the denticles are blunt
and the conidiogenous cells are constantly flask-shaped. In the section Acrodontium the denticles are mostly
smaller, sometimes directed downwards by subsequent proliferation, and the conidiogenous cells are flaskshaped or subulate, more variable; the rachis is often not clearly differentiated from the basal part. In both
sections the newly developing conidium does not grow in lateral direction, as in Beauveria, but strictly upwards.
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Fig. 1. Proliferating tips of conidiogenous cells. a. conidiogenous cell without elongation; b. geniculate, denticulate rachis;
c. geniculate, cicatrized rachis; d. straight, denticulate rachids.
Ramification. Vuillemin (1912) classified Beauveria in the family [p. 3] Verticilliaceae, because of the
resemblance of the clustered branches of B. bassiana (Bals.) Vuill. to the conidial apparatus of Spicaria aphodii
Vuill. [= Paecilomyces fumoso-roseus (Wize) Brown & G. Smith], though the latter had a more regular
branching. Also similar are the species of Tolypocladium W. Gams. However, the species of these genera are
defined as having phialides, whereas in Beauveria and Tritirachium blastic conidia are formed on sympodially
elongating conidiogenous cells.
The conidiogenous cells may be supported by differentiated side branches of aerial hyphae (prophialides
according to Petch, 1931, conidiophores and vesicles according to MacLeod, 1954). These hyphae may be of
three different types. The first type is creeping and little differentiated from the vegetative hyphae, the second
type is ascendent and somewhat differentiated but imperceptably merging into the vegetative mycelium, the
third type is suberect to erect and distinctly differentiated by having thicker and pigmented walls. Fertile hyphae
with ramifications including conidiogenous cells are called conidiophores (Pirozynski in Kendrick, 1972) when
they are markedly differentiated.
In some species of Beauveria the slightly differentiated fertile hyphae support swollen side branches which
often form a densely clustered conidial apparatus. The conidiogenous cells may also arise from stalk cells or
directly from the hyphae. In B. alba (Limber) Saccas the conidiogenous cells are formed in small whorls along
ascendent hyphae. This pattern resembles that of the species in Tritirachium sensu str. In Acrodontium the
conidiogenous cells form part of differentiated, compound conidiophores, or arise directly from the vegetative
mycelium. The sect. Grisea contains species with both differentiated conidiophores and simple conidiogenous
cells. In sect. Acrodontium no differentiated conidiophores are present.
Key to the genera and sections discussed in the present paper

1a.
1b.
2a.
2b.
3a.
3b.
4a.
4b.
Conidia globose, ellipsoidal or subcylindrical, mostly with a rounded base,

hyaline or nearly so
Conidia subglobose to fusiform, distinctly apiculate at the base, somewhat
pigmented or hyaline
Conidiogenous cells denticulate, with or without elongation, arising in clusters
from subtending cells or solitarily from undifferentiated hyphae; aerial
mycelium usually hyaline
Conidiogenous cells cicatrized, with elongation, arising in whorls from
ascendent hyphae; aerial mycelium never purely hyaline
Conidiogenous cells with elongation; synnemata mostly absent
Conidiogenous cells without elongation; synnemata mostly present
[p. 4]
Conidiogenous cells arising orthotropically from undifferentiated hyphae only;
conidia hyaline
Conidiogenous cells arising both plagiotropically from more or less
differentiated hyphae and orthotropically from undifferentiated hyphae;
conidia somewhat pigmented Acrodontium section
2
4
3
Tritirachium
Beauveria
Isaria
Acrodontium section
Acrodontium
Grisea
Beauveria Vuill.
Beauveria Vuill. - Bull. Soc. bot. Fr. 59: 40. 1912.
Colonies growing moderately slowly, appearing lanose, powdery or funiculose, rarely forming synnemata,
white or yellowish, occasionally pinkish. Aerial hyphae hyaline, smooth- and thin-walled, loose or sometimes
fasciculate. Conidiogenous cells arising from short, often one-celled, more or less swollen stalk cells, often in
dense clusters, or scattered or in whorls from undifferentiated hyphae; they consist of a globose to fusiform
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basal part, and a geniculate, denticulate rachis. Conidia one-celled, hyaline, smooth, thin-walled, globose to
ellipsoidal. No chlamydospores or perfect states were observed.
Descriptions are based on colonies growing on oatmeal agar at 20 C.
Type species: Beauveria bassiana (Bals.) Vuill.
Key to the species

1a.
1b.
Conidiogenous cells mostly in whorls along ascendent hyphae

Conidiogenous cells scattered or in irregular clusters on swollen stalk cells
2a.
Conidia globose or subglobose, in pure culture sometimes broadly ellipsoidal; stalk

cells and globose to flask-shaped conidiogenous cells mostly forming dense
clusters
Conidia mostly ellipsoidal, sometimes subglobose; stalk cells, if present, and
conidiogenous cells rather slender and rarely clustered
2b.
B. alba
2
B. bassiana
B. brongniartii
1. Beauveria bassiana (Bals.) Vuill. - Fig. 2 and 3

Sporotrichum densum Link - Mag. Ges. naturf. Freunde, Berlin 3: 13. 1809; per Pers. - Mycol. eur. 1: 76. 1822 [non
Sporotrichum densum (Ditm. per Pers.) Fr. - Syst. mycol. 3: 459. 1832] = Isaria densa (Link per Pers.) Giard - C. r.
hebd. Sanc. Acad. Sci., Paris 113: [p. 5] 270. 1891 = Spicaria densa (Link per Pers.) Vuill. - Bull. Sanc. Soc. Sci.
Nancy 11: 1S3. 1910 = Beauveria densa (Link per Pers.) Picard - Annls c. natn. Agric. Montpellier 13: 200. 1914 =
Penicillium (Spicaria) densum (Link per Pers.) Biourge - Cellule 33: 102. 1923.
Botrytis paradoxa Bals. - Gazetta di Milano, Suppl., 19 June 18 (nomen provisorium) = Botrytis bassiana Bals. Linnaea 10: 611. 1835 = Stachylidium bassianum (Bals.) Mont. - Syll. Gen. Spec. crypt. p. 301. 1856 = Spicaria
bassiana (Bals.) Vuill. - Bull. Sanc. Soc. Sci. Nancy 11 : 15 3. 1910 = Beauveria bassiana (Bals.) Vuill. - Bull. Soc. bot.
Fr. 59: 40. 1912 = Penicillium bassianum (Bals.) Biourge - Cellule 33: 101. 1923 = Beauveria paradoxa (Bals.) Ramsb. Trans. Br. mycol. Soc. 25:436.1941 (incidentally mentioned).
Sporotrichum larvatum Peck - Rep. N. Y. St. Mus. 32: 44. 1879 = Sporotrichum larvicolum Peck - Bull. N. Y. St.
Mus. 1: 18. 1887 (substitute name).
Sporotrichum globuliferum Speg. - An. Soc. cient. argent. 10: 278. 1880 = Beauveria globulifera (Speg.) Picard Annls c. natn. Agric. Montpellier 13: 203. 1914.
Sporotrichum minimum Speg. - An. Soc. cient. argent. ri: 123. 1881; Pettit in Bull. Cornell Univ. agric. Exp. Stn 97:
362. 1895 = Trichoderma minimum (Speg. sensu Pettit) G. Mller - Wiss. Z. Humboldt - Univ. Berlin, math.-nat. R. 14:
775. 1965
Botrytis bassiana (Bals.) Vuill. subsp. tenella Sacc. - Michelia 2: 544. 1882 = Botrytis tenella (Sacc.) Delacr. - J.
Agric. prat., Paris 1: 164. 1891 = Isaria tenella (Sacc.) Giard - C. r. hebd. Sanc. Acad. Sci., Paris 112: 1272. 1891
(incidentally mentioned) = Beauveria tenella (Sacc.) Siemaszko - Archwm Nauk biol., Tow. Nauk Warszawa 6: 35. 1937
(incidentally mentioned) = Beauveria tenella (Sacc.) MacLeod - Can. J. Bot. 32: 8231954.
Botrytis brongniartii Sacc. subsp. delacroixii Sacc. - Syll. Fung. 20: 540. 1892 = Botrytis delacroixii (Sacc.) Delacr. Bull. Soc. mycol. Fr. 9: 183. 1893 = Spicaria delacroixii (Sacc.) Vuill. - Bull. Sanc. Soc. Sci. Nancy : 153. 1910 =
Penicillium delacroixii (Sacc.) Biourge - Cellule 33: 102. 1923 = Beauveria delacroixii (Sacc.) Petch - Trans. Br. mycol.
Soc. 10: 249. 1924.
Isaria vexans Pettit - Bull. Cornell Univ. agric. Exp. Stn 97: 399. 1895.
Isaria citrinula Speg. - An. Mus. nac. Hist. nat. B. Aires 20: 449. 1910.
Botrytis effusa Beauv. - Rapp. Lab. tud. Soie, Lyon 14: 25. 1911 [non Botrytis effusa Grev. - Fl. Edin. p. 468. 1824)
= Beauveria effusa (Beauv.) Vuill. - Bull. Soc. bot. Fr. 59: 40. 1912.
Botrytis necans Massee - Bull. misc. Inf. R. bot. Gdns Kew 1914: 159.
Botrytis stephanoderis Bally in Friederichs & Bally - Meded. KoffiebessenboeboekFonds 6: 106. 1923 = Beauveria
stephanoderis (Bally) Petch - Trans. Br. mycol. Soc. 10: 249. 1924.
Sporotrichum sulfurescens van Beyma - Verb. K. ned. Akad. Wet., Afd. Nat. 26: 16. 1928 = Beauveria sulfurescens
(van Beyma) J. J. Taylor - Mycologia 62: 820. 1970.
Beauveria laxa Petch - Trans. Br. mycol. Soc. 16: 58. 1931.
Isaria shiotae Kuru - Jap. J. med. Sci. Trans. Abstr., Ser. 9, 2: 351. 1931 = Beauveria shiotae (Kuru) Langer. in Brumpt Prcis Parasitol., ed. 5 p. 1839. 1936 = Tritirachium shiotae (Kuru) Langer. - Annls Parasit. hum. comp. 22: 98. 1947.
Beauveria doryphorae Poisson & Patay - C. r. hebd. Sanc. Acad. Sci., Paris 200: 961. 1935.
Beauveria bassiana Bals. var. lunzinensis Szilvinyi - Zentbl. Bakt. ParasitKde, Abt. 2, 103: 178. 1941. [p. 6]
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Fig. 2. Beauveria bassiana on natural substrate, conidial structures.
Fig. 3. Beauveria bassiana on oatmeal agar. a. conidial structures; b. conidiogenous cells; c. conidia. [p. 5]
Colonies in vitro attaining a diameter of 6-23 mm in 8 days, appearing lanose, floccose, velvety to powdery,
sometimes funiculose, rarely forming synnemata (Rhizomorphen, Blunck, 1939), up to 5 mm. high; at first
white, later becoming yellowish, occasionally reddish. Reverse uncoloured or yellowish to pinkish. Exudate
rarely produced, odour absent. Submerged hyphae hyaline, smooth-walled, 1.5-3 m wide. Hyphae of the aerial
mycelium hyaline, smooth-walled, 1-2 m wide, creeping or ascendent; [p.7] they bear groups of swollen lateral
cells, mostly 3-6 x 3-5 m, which by further branching give rise to smaller swollen cells or 1-5 conidiogenous
cells in the first or second order. In fresh isolates the conidial apparatus is tightly clustered; in older strains
branching becomes less tight, the conidiogenous cells occur in small groups or solitarily on ellipsoidal to
subcylindrical lateral cells (measuring up to 15 x 6 m), or arise directly from the hyphae. Conidiogenous cells
consisting of a globose to flask-shaped, sometimes elongate basal part, mostly 3-6 x 2.5-3.5 m, terminal cells
mostly more slender, and a well developed rachis, up to 20 m long or even longer and rather constantly 1 m
wide, geniculate or irregularly bent, denticulate, denticles as wide as the rachis, up to 1 m long. Conidia hyaline
or rarely yellowish, smooth, globose to broadly ellipsoidal, sometimes with an apiculate base, (1.5-) 2-3 (-4) x
(1.5-) 2-2.5 (-3) m. No chlamydospores were observed.
Mycelium on natural substrate typically cushion-like, granular-pulverulent, sometimes funiculose or rarely
producing synnemata, yellowish, rarely reddish. Hyphae of the aerial mycelium bearing a conidial apparatus as
described above, which is tightly clustered, the conidiogenous cells occuring in dense, globose heads only.
Basal parts of the conidiogenous cells globose, subglobose or somewhat flask-shaped, mostly 2.5-4 x 2-3 m.
A perfect state without name was reported by Schaerffenberg (1955).
Material examined
Herbarium specimens
Sporotrichum densum in herb. B (type), on insect, Rostock, Germany.
Botrytis bassiana subsp. tenella in herb. PAD, on Vespa sp., Padova, Italy, leg. O. Alexis.
Sporotrichum larvatum in herb. NYS (type), on dead larva under alder bushes, Adirondack, Massachusetts, USA.
Sporotrichum globuliferum in herb. LPS: 21680 (type), on Coleoptera indet., San Jose de Flores, Argentina, 188o;
21669, on Acridium sp.; 21671, on Coleoptera indet. and surrounding decaying wood; 21675, on coccid; 21683, on
Coleoptera indet.
Sporotrichum minimum in herb. LPS, 21686 (type), on Formica sp., Buenos Aires, Argentina, July 1881.
Isaria citrinula in herb. LPS, 11215 (type), 2 specimens on Lepidoptera indet., Tucumn, Argentina, April 1906.
Isaria vexans in herb. G. F. Atkinson (CUP), on larva of Coleoptera indet., probably authentic material, which fits Pettits
description.
Botrytis necans in herb. K (type), on Brachartonia catoxantha, - Singapore, leg. S. H. Burkill, 1914.
Beauveria laxa in herb. Fetch (K): R-114 (type), on insect larva on Indigofera endecaphylla, Kirimettia, Thailand, January
1927; R-333, R-371, R-374 and R-474, on insect, cocoons of caterpillar, cocoons of unidentified insect and cocoons of
Lepidoptera indet., respectively, Nuwara Eliya, Ceylon, 1927/28; R-401, on unidentified chrysalid, Wellaway, GreatBritain, January 1928.
Beauveria bassiana in herb. CBS, on Coccinella sp., Hoge Veluwe, leg. G. A. de Vries, 1971.
Living strains
CBS 110.25 isolated from Oecophylla smaragdina, Ceylon, sent in 1925 by T. Petch as Beauveria globulifera. [p. 8]
CBS 119.26, type culture of Botrytis stephanoderis, isolated from Stephanoderis hampei, Java, 1919.
CBS 209.27 (= ATCC 7159), type culture of Sporotrichum sulfurescens, isolated as culture contaminant.
CBS 118.30 (= ATCC 9453) isolated from Bombyx mori, sent in 1930 by A. Guillermond as Beauveria effusa.
CBS 337.32, type culture of Isaria shiotae, isolated from human thorax abcess, Japan.
CBS 124.33 isolated from Brachyderes incanus, sent in 1933 by H. van Vloten.
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CBS 127.35, type culture of Beauveria doryphorae, isolated by R. Patay from Leptinotarsa decemlineata, France.
CBS 121.36 isolated from Hylobius abietis; CBS 122.36 from Lophyrus pini; CBS 123.36 from Leptura sp.; CBS 124.36
from Acanthocinus aedilis; CBS 125.36 from Hylurgops palliatus; CBS 126.36 from Orthotomicus laricis; CBS 127.36
from Blastophagus piniperda; CBS 128.36 from Ips typographus; CBS 129.36 from unidentified Curculionid; CBS 130.36
and CBS 132.36 from Carpocapsa pomonella; CBS 131.36 from Cossus cossus; CBS 133.36 from Pentatoma rufipes;
all strains isolated in Poland, sent in 1936 by W. Siemaszko.
CBS 120.50 (= LCF 351) isolated from Bombyx mori, Brazil, May 1935, sent in 1950 by J. B. Marchionatto.
CBS 212.61 (= IMB 415) isolated from soil, N. France, sent by G. Mller as Sporotrichum epigaeum var. terrestre
Daszewska, from Laboratoire de Cryptogamie, Paris. CBS 324.67 and 325.67 isolated from Eurygaster sp., France, and
Cleonus punctiventris, Yugoslavia, both sent in 1967 by P. Ferron.
CBS 639.68 isolated from insect on leaf of Vicia sp., and CBS 640.68 isolated from Laodelphax striatellus, Rehovot,
Isral, both sent in 1968 by R. Kenneth, under No. 2400 and No. 2491, respectively.
CBS 465.70 (= IMI 147,691) isolated from pupa of Thaumatopoea wilkinsonii, Isral, sent in 1970 by R. Kenneth under
No. 2660.
CBS 650.71 isolated by A. Kahanp, Helsinki, Finland.
CBS 651.71 and 652.71 isolated by R. A. Samson from insects collected by J. Daams in Ankeveen and Kortenhoef, May
and July 1971.
CBS 653.71 isolated by R. A. Samson from insect, Bilthoven, March 1971.
CBS 654.71 isolated by W. Gams from the basidiomycete Cyphellopsis anomala, Baarn, February 1971.
CBS 715.71 isolated by J. Halperin from Phalera bucephaloides, Ilanot, Isral, sent in 1971 by R. Kenneth under No.
2643.
CBS 723.71 isolated from human sputum, Groningen, sent in 1971 by A. Kikstra.
Discussion
The species is a facultative but highly virulent parasite on insects, occurring rarely on other animals
(MacLeod, 1954) and man (Kuru, 1931).
On artificial media most of the strains lose their pathogenicity and degenerate. The capacity to form
clustered fertile branches decreases rather soon. Moreover the conidiogenous cells tend to elongate and
become narrower. The shape of the conidia in each strain is rather constant, though their size may vary
considerably in strains in which sporulation has decreased.
Most of the names considered here to be synonymous with B. bassiana are discussed below in
chronological order.
Sporotrichum densum. The species was described by Link in 1809. The type collection appeared to be the
present species. In 1817 Ditmar described a Botrytis densa Ditm., the type of which (B) is of doubtful identity.
[p. 9]
Both species were validated by Persoon (1822). Fries (1832) transferred Ditmars species to Sporotrichum. He
thought that Links species belonged to Isaria, but did not actually make the combination. This was done by
Giard (1891), who revalidated the species while studying the old literature on entomogenous fungi in order to
find the correct epithet for the red muscardine sent for identification by L. le Moult. Although he saw differences
in the shape of the conidia between his material and Links species, he followed Bresadola (in Saccardo, 1882)
in paying no attention to it. Because of the synnematous growth he made the combination Isaria densa and
placed Botrytis tenella in the synonymy. On the other hand he retained B. bassiana to include strains without
synnemata. So the epithet densum Link has been used for two Beauveria species, and has to be rejected
according to Art. 69 of the International Code of Botanical Nomenclature as a nomen confusum.
Botrytis bassiana. This was the best known name for the Beauveria species with globose conidia. It was
published first in a newspaper as Botrytis paradoxa, but as the author wanted to dedicate the species to its
discoverer, he changed it into B. bassiana. The latter name was subsequently frequently used in the literature
and taken over in Saccardos Sylloge Fungorum (1886).
Sporotrichum globuliferum. Many authors mentioned collections under this name; MacLeod (1954) was the
first to regard it as synonymous with Beauveria bassiana.
Sporotrichum minimum. Examination of the type material proved its identity with Beauveria bassiana. A
second collection of this species was mentioned by Pettit (1895). He gave rather vague illustrations, on the
basis of which Mller (1965) thought that Spegazzini and Pettit had described different species. Mller
transferred Pettits strain to Trichoderma. Gsswald (1939) mentioned S. minimum as a synonym of S.
minutulum Speg. The type of the latter is in bad condition. According to some figures on the envelope, possibly
an Oidiodendron species was meant.
Botrytis bassiana subsp. tenella was described as having globose conidia, measuring 1.5 m in diameter,
whereas in the typical subspecies they were about 3 m in diameter. Both in Michelia (1882) and in Sylloge
Fungorum (1886) Saccardo mentioned three original collections. One of these was depicted in Fungi Italici
(1879) and is therefore considered as the lectotype. The second collection in the Saccardo Herbarium
appeared to be Beauveria bassiana. The third was collected by Bresadola and had oval conidia. Bresadola
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sent part of his material to Delacroix for comparison with L. le Moults red muscardine. That is why the
description by Delacroix (1891) of his Botrytis tenella with ellipsoidal conidia differed considerably from
Saccardos. The combination with Beauveria was mentioned by Siemaszko (1937); however, he only described
B. bassiana, B. globulifera and B. densa as independent species. MacLeod (1954) actually made the
combination Beauveria tenella.
Botrytis brongniartii subsp. delacroixii. During his stay in Algeria Brongniart (1891a and b) collected several
parasitic fungi from the migratory [p. 10] locust. The first of these was a Fusarium species (Delacroix, 1893). It
was sent to Trabut, who described it as Botrytis acridiorum. The second was a Beauveria species with globose
conidia, illegitimately published as Botrytis acridiorum Brongniart & Delacr. (later homonym). A species with
ellipsoidal conidia, which strongly resembled the red muscardine found in France, was described without a
name. Saccardo (1892) named it Botrytis brongniartii. He also redescribed the species with globose conidia,
which he validated by naming it Botrytis brongniartii subsp. delacroixii.
Botrytis effusa. No material is preserved in PC, and the CBS culture of the type was lost. According to the
description, important criteria for separation from B. bassiana were the floccose to funiculose mycelium, and a
stable, wine-red pigment produced when the fungus was cultivated on potato. Microscopically it did not differ
from B. bassiana. This conclusion was also drawn by F. H. van Beyma in an unpublished study of the type
strain.
Botrytis stephanoderis. This species was originally distinguished by 1-2 cm high synnemata, which may be
branched, and by a yellow reverse. The value of these characters was already doubted by Schwarz (1924). In
the type strain CBS 119.26 both characters had almost disappeared. Under the name B. stephanoderis f.
macroconidiana Averna-Sacc (1930), as cited in the Index of Fungi (1971), the chlamydospores of
Lachnidium acridiorum Giard [= Fusarium solani (Martius) Sacc.] were erroneously described.
Beauveria laxa. Collections of this species in K contained mostly Paecilomyces farinosus (Holm. per Fr.)
Brown & G. Smith. In some capsules, among which the type, Beauveria bassiana could be found. According to
Petchs description the species should differ from B. bassiana mainly by its angular conidia, which were formed
in crowded masses on the rachis. However, in our observations the conidia were all globose.
Isaria shiotae. The type strain CBS 337.32 is in rather bad condition, but from Kurus (1931) detailed
descriptions and illustrations the synonymy is obvious.
Botrytis bassiana var. lunzinensis. This variety, of which no type material is left, would differ from the typical
variety by its yellow and reddish pigments.
2. Beauveria brongniartii (Sacc.) Petch - Fig. 4
? Sporotrichum epigaeum Brunaud - Annls Soc. Sci. natn. Rochelle 1888; sensu Aschieri - Atti Ist. hot. Univ. Lab.
crittogam. Pavia, Ser. 4, 4: 207. 1929 = Sporotrichum schenckii Matt. var. fioccoi Dodge - Med. Mycol. p. 808. 1935
(substitute name) = Beauveria epigaea (Brunaud) Langer. in Brumpt - Prcis Parasitol., ed. 5 p. 18391936 =
Sporotrichum beurmannii Matr. & Ramond var. fioccoi (Dodge) Red. & Cif. - Tratt. Micopat. umana 5: 481. 1942 =
Tritirachium epigaeum (Brunaud) Langer. Annls Parasit. hum. comp. 22: 98. 1947.
Botrytis brongniartii Sacc. - Syll. Fung. ro: 540. 1892 = Beauveria brongniartii (Sacc.) Petch - Trans. Br. mycol. Soc.
1o: 249. 1924.
Botrytis melolonthae Sacc. - Annls mycol. 10: 320. 1912 = Beauveria melolonthae [p. 11] (Sacc.) Cif. - Publnes
Estac. agron. Moca, Ser. B, 14: 169. 1929.
Isaria kogane Hasegawa & Koyama - Bull. Govt. Forest Exp. Stn Meguro 4. 74. 1941
Isaria sp., Briggs, Fergus & Shannon - Tetrahedon 8, suppl. 1: 269. 1966.
Misapplied names
Botrytis tenella (Sacc.) Delacr. - J. Agric. prat., Paris 2: 164. 1891 = Beauveria tenella (Sacc.) MacLeod - Can. J.
Bot. 32: 823. 1954.
Isaria densa (Link per Pers.) Giard - C. r. hebd. Sanc. Acad. Sci., Paris 113: 270. 1891.
Fig. 4. Beauveria brongniartii, CBS 109.24. a. conidial structures on oatmeal agar; b. conidial structures on 2/o malt
agar; c. conidia.
Colonies in vitro attaining a diameter of 10-16 mm in 8 days, appearing lanose, floccose, velvety to
powdery, sometimes funiculose, up to 5 mm high; at first white, later often becoming yellowish to pinkish, rarely
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reddish to purple. Reverse uncoloured or yellowish to orange, rarely red or purple. Exudate and odour absent.
Submerged hyphae hyaline, smoothwalled, 1.5-3 m wide. Hyphae of the aerial mycelium hyaline,
smoothwalled, 1-4 m wide, creeping or ascendent, bearing small groups of orthotropic or slightly plagiotropic
lateral cells, up to 9 x 5 m, which by further branching may give rise to smaller, subglobose cells or 1-3 (-4)
conidiogenous cells in the first, or sometimes in the second order. Occasionally the conidial apparatus is
clustered; more often the conidiogenous cells are arranged in small groups or solitarily along the hyphae.
Conidiogenous [p. 12] cells consisting of a subglobose or flask-shaped, sometimes subcylindrical
basal part, (3-) 4-15 (-28) x (1.5-) 2-3.5 (-4) m, and a well developed rachis, up to 25 m long and mostly 1-1.5
m wide, geniculate or irregularly bent, denticulate, denticles mostly as wide as the rachis, up to 1.5 m long.
Conidia hyaline, smooth, ellipsoidal (rarely subglobose), widest near or below the middle, sometimes with a
pointed or apiculate base, (2-) 2.5-4.5 (-6) x (1.5-) 2-2.5 (-3) m. No chlamydospores were observed.
Mycelium on natural substrate lanose to somewhat pulverulent, rarely funiculose, yellowish, sometimes
reddish. Hyphae of the aerial mycelium bearing a conidial apparatus, which is often more clustered than in pure
culture. Perfect state unknown.
Material examined
Herbarium specimen
Botrytis melolonthae in herb. PAD (type), on Melolontha melolontha, Pavia, Italy, 1898.
Living strains
CBS 106.22 (= ATCC 9452) isolated from mummified larva, Java, Indonesia, sent in 1924 by L. le Moult.
CBS 109.24 isolated by R. Ciferri from insect, sent in 1924 by L. Montemartini.
CBS 111.25 isolated from Mantis sp., and CBS 120.26 from unidentified Salticide, both sent by National Collection of
Type Cultures, Kew.
CBS 410.34 (= ATCC 7145) isolated from man, sent in 1934 by G. Pollacci, described by E. Aschieri (1929).
CBS 112.42 isolated from Melolontha melolontha, Kiel, Germany, sent in 1942 by H. Blunck.
CBS 128.53 isolated from larva of Balaninus sp., Turkey, sent in 1953 by M. Gbelez.
CBS 223.53 (= IFO 5299), type culture of Isaria kogane.
CBS 326.67 isolated from Costelytia zeelandica, and CBS 327.67 from Melolontha melolontha, both sent by P. Ferron,
under No. 13 and 6 respectively.
CBS 722.71 (= PDD 6363) sent in 1971 by D. W. Roberts as Isaria sp., strain producing isarolide described by Briggs &
al. (1966).
Discussion
Recently the species has mostly been referred to as Beauveria tenella or B. densa; the type specimens of
both, however, appeared to be B. bassiana. The following names are considered here to be synonyms of B.
brongniartii:
Sporotrichum epigaeum. No type material was available in PC. From Brunauds description it is, however,
likely that he meant a Beauveria species. The oldest strain available is the one described by Aschieri (1929). On
this strain all later combinations are based. Aschieri recognized the relationship with Beauveria, but did not
change the name. Dodge (1935) made a new name and mentioned in the synonymy Sporotrichum epigaeum
Aschieri not Brunaud, though Aschieri (1929) had not excluded the type. This misunderstanding was taken over
by MacLeod (1954). The combination with Tritirachium was made by Langeron (1947) following Limber (1940),
who mentioned the geniculate rachids of the conidiogenous [p. 13] cells as a genus character. Langeron (1947)
transferred almost all Beauveria species to this genus. Our observations on the strain do not exactly agree with
Aschieris. No chlamydospores were found, and the conidia were obviously ellipsoidal, (2-) 2.5-3.5 (-4) x (1.5-) 2
(-3) m, while Aschieri (1929) called them rundlich oder leicht lnglich (Mller, 1964). No clusters of
conidiogenous cells were reported.
Botrytis brongniartii, see discussion of Beauveria bassiana, under B. brongniartii var. delacroixii.
3. Beauveria alba (Limber) Saccas - Fig. 5
Tritirachium album Limber - Mycologia 32: 27. 1940 = Beauveria alba (Limber) Saccas - Revue Mycol. 13: 64. 1948.
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Fig. 5. Beauveria alba, CBS 570.71. a, b. conidial structures; c. conidia.
Colonies in vitro attaining a diameter of 14-16 mm in 8 days, appearing lanose to floccose, up to 2 mm

high, sometimes zonate, purely hyaline. Reverse ochraceous buff or uncoloured. Exudate and odour absent.
Submerged hyphae and hyphae of the aerial mycelium hyaline, smooth-walled, 1-2.5 m wide. Fertile hyphae
mostly slightly differentiated, 2-4 m wide, ascendent to procumbent, somewhat stiff, apically dichotomously
branched, bearing conidiogenous cells in whorls of 1-3 at wide, often right angles; conidiogenous cells
occasionally scattered or with two on a lateral stalk [p. 14] cell which is cylindrical, 6-12 x 2-3 m, or rarely
swollen up to 5 m.
Conidiogenous cells consisting of an elongate to subcylindrical tapering basal part, (7-) 10-24 (-30) x 1.5-2.5
m, sometimes slightly swollen up to 3.5 m, and a well developed rachis, up to 3 5 m long and rather
constantly 1 m wide, geniculate, denticulate, denticles as wide as the rachis and up to 1 m long. Conidia
hyaline, smooth, globose to subglobose, sometimes with an apiculate base, (1.5-) 2-3 (-3.5) x (1.5-) 1.5-2.5 (-3)
m. No chlamydospores were observed.
Mycelium on natural substrate thin, lanose to arachnoid, purely white. Perfect state unknown.
Material examined
Herbarium specimen
Tritirachium album in herb. BPI (type), on cover of a book, Woods Hole, Massachusetts, USA, March 1939, leg. O. W.
Richards.
Tritirachium album in herb. BPI, dried culture isolated from human sputum, Duke, USA,
N. F. Conant.
Living strains
CBS 348.55 isolated by G. A. de Vries from skin of a woman with eczema vesiculosum, Netherlands.
CBS S70.71 sent in 1971 by R. T. de Jongh, Curaao, under No. 8296-1970.
CBS 836.71 isolated from Pisum sp., sent in 1971 by R. Kenneth under No. 2343.
Discussion
The zig-zag shape of the rachis and the verticillate ramification make the species reminiscent of
Tritirachium. Reasons for classification in Beauveria are the denticulate rachis and the absence of pigment in
the aerial mycelium. Ramification is not always strictly verticillate: poorly developed subcultures sometimes can
be distinguished from B. brongniartii only by the shape of their conidia.
Isaria Fr.
Isaria Fr. - Syst. mycol. 3:270. 1832.
Colonies growing moderately slowly, appearing velvety, soon forming synnemata, up to 70 mm long, white
or yellowish. Aerial hyphae hyaline, smooth- and thin-walled, loose or fasciculate. Conidiogenous cells
scattered, arising directly from undifferentiated hyphae or from one-celled, sometimes swollen side branches,
occasionally in dense clusters; they consist of a swollen basal part, not elongating, and a short and narrow,
irregular, [p. 15] denticulate conidiiferous portion. Conidia one-celled, hyaline, smooth, thin-walled, subglobose
to subcylindrical or kidney-shaped. No chlamydospores or perfect states were observed.
Descriptions are based on colonies growing on oatmeal agar at 20 C.
Lectotype species: Isaria felina (DC. per Fr.) Fr.
Discussion
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The name Isaria was first used by Hill (1751). He mentioned three species, which probably were Calocera
viscosa Fr., Ceratiomyxa fruticulosa (Mller) MacBride and a Puccinia species respectively (Petch, 1934).
Persoon (1794) introduced the name Isaria again for the description of two species, i.e. I. mucida (=
Ceratiomyxa fruticulosa) and I. agaricina [= Tilachlidium brachiatum (Batsch per Fr.) Petch]. In 1821 Fries cited
the name Isaria Pers. in the introduction of his Systema Mycologicum, without mentioning any species. In 1832
he did not refer to Persoon (1794) but to Hill (1751). None of Hills species, however, was mentioned, while both
original species of Persoon were listed. Isaria terrestris Fr. was the first species mentioned by Fries (1832). It
was chosen as the lectotype of Isaria by Petch (1934). Examination of the type material of this species (herb.
UPS) showed, however, that the fungus consisted of sterile hyphae, probably belonging to an immature
basidiomycete.
Also Gray (1821) referred to Persoon, though none of Persoons species are explicitly mentioned. He listed
I. velutipes Link, I. crassa Pers., which are both identical with Paecilomyces farinosus (Dicks. per Fr.) Brown &
G. Smith according to the type material (L), and I. eleutheratorum Nees. Isaria farinosa Dicks. per Fr. was
proposed as lectotype of Isaria by Clements and Shear (1931). If this were accepted, the younger genus
Paecilomyces would become a synonym of Isaria.
In order to provide a generic name for synnematous Beauveria-like fungi, von Arx (1970) chose Isaria felina
as lectotype of Isaria. In this sense the genus at present contains two species.
Key to the species

1a.
1b
Conidiogenous cells flask-shaped, often curved; conidia broadly ellipsoidal

Conidiogenous cells globose; conidia subcylindrical to kidney-shaped
I. felina
I. orthopterorum
4. Isaria felina (DC. per Fr.) Fr. - Fig. 6

Clavaria (?) felina DC. - Fl. fr. 6:30. 1815; per Fr. - Syst. mycol. 1: 496. 1821 = Fibrillaria felina (DC. per Fr.) Pers.
- Mycol. eur. 1:53. 1822 = Isaria felina (DC. per [p. 16] Fr.) Fr. - Syst. mycol. 3: 271. 1832 = Penicillium felinum (DC.
per Fr.) Biourge - Cellule 33: 102. 1923.
Isaria felina (DC. per Fr.) Fr. var. suina Sacc. - Michelia 2: 561. 1882.
Isaria felina (DC. per Fr.) Fr. var. aviaria Sacc. - Michelia 2: 561. 1882.
Isaria edwalliana P. Henn. - Hedwigia 43: 209. 1904.
Isaria felina (DC. per Fr.) Fr. var. domestica Speg. - An. Mus. nac. Hist. nat. B. Aires 23: 121. 1912.
Isaria felina (DC. per Fr.) Fr. var. cuniculina Ferr. - Fl. ital. crypt., Fasc. 6, Hyphales p. 152. 1910.
Isaria felina (DC. per Fr.) Fr. var. pirina El. Marchai & Et. Marchai - Bull. Soc. R. bot. Belg. S4: 134. 1921.
Isaria fimicola Sternon - Bull. Soc. R. bot. Belg. 55: 145. 1923.
Isaria cretacea van Beyma - Zentbl. Bakt. ParasitKde, Abt. 2, 91: 350. 1935 = Beauveria cretacea (van Beyma)
Matsushima - Microf. Solomon Isl., Papua-New Guinea p. 7. 1971.
Fig. 6. Isaria felina. a. conidial structures; b. conidiogenous cells; c. conidia.
Colonies in vitro attaining a diameter of 8-14 mm in 8 days, forming in fresh isolates a dense felt, from
which several white, positively phototropic synnemata arise (Taber & Vining, 1963), up to 70 mm high and
constantly 1 mm wide, terete, tomentose, usually unbranched, or occasionally branched at the apex, in old
strains appearing lanose to floccose; at first white, later becoming yellowish. Reverse uncoloured or yellowish
to yellow brown. Exudate rarely produced, odour absent. Submerged hyphae hyaline, smooth-walled, 1-3 m
wide. Hyphae of the aerial mycelium hyaline, smooth-walled, 1-4.5 m wide, creeping, ascendent or fasciculate,
bearing orthotropic conidiogenous cells, solitarily or in small groups; sometimes 1-2 conidiogenous cells are
supported by a slightly swollen stalk [p. 17] cell. Conidiogenous cells not elongating, consisting of a swollen,
flask shaped or curved, occasionally elongate basal part, mostly 3-8.5 x 2-2.5 m, and a short, irregular
conidiiferous portion, mostly comprising 2-4 denticles; in age a short geniculate rachis may be formed. Conidia
hyaline, smooth, subglobose, ellipsoidal or ovoidal, sometimes with a pointed base, (2.5-) 3-4 (-4.5) x (2-) 2.5-3
(-3.5) m. No chlamydospores were observed.
Mycelium on natural substrate forming a thin layer of hyaline hyphae, from which several white,
erect synnemata arise, up to 80 mm high and constantly 0.5-1 mm wide, usually unbranched.
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Perfect state unknown.

Material examined
Herbarium specimens
Isaria felina in herb. Fries (UPS), probably collected by F. Chevallier.
Fibrillaria felina in herb. Persoon (L), Nos. 910.262.101 and 910.262.106, leg. F. Chevallier.
Isaria felina in herb. PC under No. 1534; ex herbarium Durieux de Maisonneuve, leg. L. Motelay, 1879.
Isaria fimicola in herb. BPI (type), cultures on carrot.
Isaria fimicola in herb. Shear (BPI), leg. P. Marchal, Gembloux, Belgium, 1924.
Isaria felina var. suina Sacc. in herb. Bresadola (BPI), type, on dung, Torino, Italy.
Isaria felina in herb. Bommer & Rousseau (B), on rabbit dung, 1882.
Isaria edwalliana in herb. S, on carnivore dung, Sao Paulo, Brazil, 1903.
Isaria felina ex herb. Sydow (S), on rat dung, Leipzig, Germany, leg. B. Auerswald.
Living strains
CBS 110.08 sent in 1908 by A. F. Blakeslee.
CBS 250.34, type culture of Isaria cretacea, Epsom, Great Britain, sent in 1934 by D. Hutchinson.
CBS 235.36 isolated by H. A. Diddens from pupa of Anaitis efformata, Soest, sent in 1936 by R. Tolman.
CBS 236.36 isolated from rabbit dung, Erlangen, Germany, sent in 1936 by H. Greis.
CBS 217.37 isolated from mouldy leaves, sent in 1936 by D. Hellinga.
CBS 312.50 isolated from rabbit dung, sent in 1950 by K. B. Boedijn under No. BR 17/50.
CBS 313.50 isolated as culture contaminant, sent in 1950 by Lab. Microbiology, Delft.
CBS 229.58 sent in 1958 by O. Verona.
CBS 648.66 sent in 1966 by F. J. Herrero, Buenos Aires, Argentina.
CBS 173.71 isolated from porcupine dung, Ontario, Canada, sent in 1969 by D. Malloch under No. TRTC 45685.
5. Isaria orthopterorum Petch - Fig. 7

Isaria (Beauveria) orthopterorum Petch - Trans. Br. mycol. Soc. 18: 70. 1933.
Mycelium on natural substrate forming a thin layer of hyaline hyphae, from which several yellowish
synnemata arise, up to 8 mm high; stalk up to 0.5 mm wide, terete or laterally compressed, when young
tomentose, later glabrous and horny, red brown, outer surface irregular, mostly [p. 18] branched, apically often
swollen up to 2 mm. Hyphae of the aerial mycelium hyaline to yellowish, smooth-walled, 2.5-3 m wide. Fertile
hyphae mostly arising from the heads of the synnemata only, hyaline, smoothwalled, 2.5-3.5 m wide, bearing
groups of subglobose to ellipsoidal lateral cells, 3-4.5 x 3-4 m or pointed if terminal, connected with a
constricted base to the hyphae, giving rise to (1-) 3-6 (-7) conidiogenous cells in the first, rarely in the second
order. The conidial apparatus is rather tightly clustered. Conidiogenous cells not elongating, consisting of a
globose to ubglobose, rarely pear-shaped basal part, mostly 2.5-3.5 m in diameter, with a constricted base,
and a short conidiiferous portion, mostly comprising 3-6 denticles; sometimes a short geniculate rachis is
present, up to 8 m long and 3 m wide. Conidia hyaline, smooth, ellipsoidal to subcylindrical, flattened on one
side to kidney-shaped, narrower at the base than at the broadly rounded tip, rarely somewhat apiculate, (2.5-)
3.5-4.5 (-5.5) x 2-3 m. No chlamydospores were observed. Perfect state unknown.
Fig. 7. Isaria orthopterorum, from type collection. a, b. conidial structures; c. conidia.
Material examined
Isaria orthopterorum in herb. Petch (K), type, on Orthoptera indet., Kotmake, Ceylon, leg. N. K. Jardine, 1920.
Discussion
This species is only known as dried herbarium material. It was collected by T. Petch from several locations
in Ceylon and once in Thailand. Some [p. 19] Ceylonese specimens, including the type, were described briefly
and illustrated (1924a) as Isaria sp. and said to occur on grasshoppers and probably also on Lepidoptera,
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though the latter was not mentioned in the subsequent description in 1933. The conidia were said to be formed
in chains, but in 1941 Petch corrected his error.
The description of Beauveria chiromensis Sawada (1959) is reminiscent of B. orthopterorum; without
examination of the type material the identity can, however, not be established. Sawada did not give a Latin
diagnosis, so the species is not validly published.
Tritirachium Limber
Tritirachium Limber - Mycologia 32: 26. Feb. 1940.
Spirotrichum Saito - J. Ferment. Technol., Osaka 27: 1. 1939 (without Latin diagnosis); ex van Beyma - Antonie van
Leeuwenhoek 6: 286. Sept. 1940.
Colonies growing rather slowly, appearing velvety to powdery, variously coloured. Conidiophores brownish,
lighter towards the base, smooth-walled, ascendent, imperceptably merging into the vegetative mycelium,
bearing in the upper part several whorls of conidiogenous cells or verticillate side branches. Conidiogenous cells
consisting of an elongate basal part, slightly swollen at the base and tapering towards the tip, and a well
developed, regularly geniculate, cicatrized rachis. Conidia one-celled, hyaline, smooth, thin-walled, subglobose
to ellipsoidal. No chlamydospores or perfect states were observed.
Descriptions are based on colonies growing on 2% malt agar at 20 C.
Type species: Tritirachium dependens Limber
Discussion
The genus was separated from Verticillium by Limber (1940) on account of the sympodial conidiogenous
cells. In order to provide a better delimitation of the genera Tritirachium and Beauveria, more emphasis is laid
on the mode of the proliferation of the conidiogenous cells. In Beauveria, Acrodontium and Hansfordia the
rachis bears distinct denticles, in the latter two genera the rachis is not or hardly geniculate. The denticles may
be very small or absent in Geniculosporium and Nodulisporium. In these genera, however, the conidiogenous
cells are cylindrical when a geniculate rachis is present, and the conidia mostly have a truncate or apiculate
base.
Key to the species

1a.
1b.
Conidiophores bearing several whorls of side branches, each supporting [p. 20] a
whorl of conidiogenous cells; colonies ochraceous
Conidiophores rarely branched, mostly supporting whorls of conidiogenous cells
only; colonies purplish
T. dependens
T. oryzae
6. Tritirachium dependens Limber - Fig. 8 a-d

Tritirachium dependens Limber - Mycologia 32: 26. 1940 = Beauveria dependens (Limber) Saccas - Revue Mycol.
13: 64. 1948.
Living cultures were not available.

Mycelium on natural substrate appearing velvety, ochreous (17 to 19"i) (Colour names with symbols refer
to Rayner (1970)).
Conidiophores hyaline at the base, golden brown in the upper part, smoothand somewhat thick-walled,
ascendent to suberect, imperceptably merging into the vegetative mycelium, regular, stiff, up to 1 mm long,
3-4.5 m wide, slightly narrowed at the base, tapering towards the tip, bearing 4-6 (-10) whorls of 3-4 (-5) side
branches or conidiogenous cells; branches 2-3 m wide, bearing conidiogenous cells in 1-2 whorls of 2-4;
branches of second order sometimes present; conidial apparatus pyramidal in outline, up to 150 m wide and
220 m high. Conidiogenous cells consisting of an elongate basal part, slightly swollen below the middle,
tapering towards the tip, mostly 13-35 x 2-3.5 m pale golden brown, and a well developed, geniculate (intervals
2 m cicatrized rachis, up to 40 m long and 1-1.5 m wide, often somewhat tapering towards the tip. Conidia
hyaline to slightly yellowish, smooth, thin-walled, globose to ellipsoidal, (2-) 2.5-3.5 (-4) x (2-) 2-3 (-3.5) m. No
chlamydospores were observed. Perfect state unknown.
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Material examined
Herbarium specimens
Tritirachium dependens in herb. BPI (type), on dead root of Yucca tremuleana, leg. D. P. Limber, May 1935.
Further collection in BPI: New York 762188, on root of Kudzu (Legume), China, 1941, det. D. P. Limber; 37-2059 (San
Francisco 27474), on bulbs of Lilium sp., China, 1950, det. P. R. Frink & A. J. Watson; 37-2601 (San Francisco 27433),
on bulbs of Lilium sp., China, 1950, det. A. J. Watson; 37-1609 (San Francisco 27285), on bulbs of Lilium eruba, Japan,
1950, det. P. R. Frink & A. J. Watson; 37-3614 (San Francisco 27357), on bulbs of Lilium giganteum, Japan, 1950, det.
P. R. Frink & A. J. Watson; 37-1845 (San Francisco 27437), on bulbs of Lycornis radiata, Japan, 1950, det. A. J.
Watson; San Francisco 29077, on Pueraria thunbergiana, China, 1952, det. P. R. Frink & A. J. Watson; 37-2811
(Hoboken 15494), on Watsonia rosea, Stellenbosch, South-Africa, 1951, leg. and det. L. M. Fenner & D. P. Limber. [p.
21]
Fig. 8. a-d Tritirachium dependens, from type collection. a. conidial structures; b. conidiogenous cells; c. conidiophore; d.
conidia. - e-h. Tritirachium oryzae. e. conidial structures; f. conidiophore; g. conidiogenous cells; h. conidia. [p. 22]
7. Tritirachium oryzae (Vincens) de Hoog, comb. nov. - Fig. 8 e-h

? Botrytis rosea Link - Mag. Ges. naturf. Freunde, Berlin 7: 36. 1816; per Pers - Mycol. eur. 1: 35. 1822 [non Botrytis
rosea DC. - Fl. fr. z: 71. 1815].
Beauveria oryzae Vincens - Revue Path. vg. Ent. agric. Fr. 10: 122. 1910 (basionym).
Beauveria brumptii Langer. & Lichaa - Bull. Acad. Md., Paris 111: 133. 1934 = Tritirachium brumptii (Langer. &
Lichaa) Langer. - Annls Parasit. hum. comp. 22: 94. 1947.
Spirotrichum purpureum Saito - J. Ferment. Technol., Osaka 27: 1. 1939 (without Latin diagnosis) = Tritirachium
purpureum (Saito) van Beyma - Antonie van Leeuwenhoek 8: 117. 1942.
Spirotrichum musae van Beyma - Antonie van Leeuwenhoek 6: 286. 1940 = Tritirachium musae (van Beyma) van
Beyma - Antonie van Leeuwenhoek 8: 117. 1942.
Tritirachium roseum van Beyma - Antonie van Leeuwenhoek 8: 1I8. 1942.
Misapplied name
Tritirachium dependens Limber sensu Batista & al. - Publoes Inst. Micol. Recife 447: 16. 1965.
Colonies in vitro growing slowly, attaining a diameter of 3-5 mm in 8 days, appearing velvety to lanose, at
first rosy vinaceous (7"f) to rosy buff (11"d), when older lilac (69") or cinnamon (13"b). Reverse dark brick (9"m)
and towards the margin lighter, or brown vinaceous (5"m). Exudate and odour absent. Submerged hyphae
hyaline to pale brownish, smoothwalled, 1-2 m wide, often somewhat stromatic. Conidiophores reddish brown,
lighter towards the base, smooth- and somewhat thick-walled, ascendent, imperceptably merging into the
vegetative mycelium, 200-1500 m long, 1.5-3 m wide, slightly tapering towards the tip, bearing mainly on the
upper part (3-) 5-12 (-30) whorls of 1-3 (-5) conidiogenous cells and occasionally a side branch which bears 1-3
conidiogenous cells. Conidiogenous cells consisting of an elongate basal part, slightly swollen below the middle,
tapering towards the tip, (7-) 9-20 (-25) x 1.5-3 m and a well developed, geniculate (intervals 1-2 m cicatrized
rachis, mostly 25 m long, up to 165 m, and I m wide. Conidia hyaline, smooth, thin-walled, globose to
ellipsoidal, rarely with a slightly apiculate base, (1.5-) 2-3 (-3.5) x 1.5-3 m. No chlamydospores were observed.
Material examined
Herbarium specimen
Botrytis rosea in herb. Sydow (B) under No. 841, on twigs of Salix sp., Brandenburg, Germany, July 1909.
Living strains
CBS 388.39, type culture of Spirotrichum purpureum, sent in 1939 by K. Saito, Japan.
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CBS 175.40, type culture of Spirotrichum musae, isolated from leaf spots of Musa sapientum, Surinam, sent in 1934 by
G. Stahel as Verticillium sp.
CBS 183.42, type culture of Tritirachium roseum, sent in 1942 under No. 1361 as Beauveria sp., Kiel, Germany. [p. 23]
CBS 164.67 isolated as culture contaminant, Recife, Brazil, 1960, sent in 1967 by A. C. Batista as Tritirachium
dependens.
CBS 442.70 isolated from packing cartoon, Abidjan, Ghana, sent in 1970 by J. Nicot under No. PC 2042.
CBS 896.70 isolated as culture contaminant, Nijmegen, sent in 1970 by G. van den Ende. CBS 837.71 isolated by K.
Martin as culture contaminant, Rehovot, Isral, sent in 1971 by R. Kenneth under No. 2330.
Discussion
No type material of Botrytis rosea was available in B; only a secondary collection from the Sydow
herbarium (B), which appeared to be Tritirachium oryzae, could be examined.
The type of Beauveria oryzae seems to be lost. Vincens (1923), who isolated several strains of the species
in Indo-China as contaminants, gave good descriptions and illustrations. Especially the Forme did not differ
in any respect from CBS 896.70, the most typical strain on which the description is based.
No type material is available of Beauveria brumptii. The description by Langeron (1934) and some
unpublished drawings by van Beyma of a CBS subculture of the type, which is lost, make its identity with
Tritirachium oryzae very likely.
Van Beymas species (1940, 1942) are mainly based on cultural characters, which, because of their variability
when the fungus is grown on different media, are of little use as taxonomic criteria.
Acrodontium de Hoog, gen. nov.

Coloniae fere lente crescunt, velutinae vel pulverulentae, varie coloratae. Hyphae aeriae subhyalinae, brunneolae vel
olivaceae, leves, fere tenui-tunicatae, septatae, ramosae, singulae vel fasciculatae. Conidiophora, si adsunt, orthotropica
vel plagiotropica e hyphis repentibus oriuntur, erecta vel procumbentia, nonnumquam crassi-tunicata et deorsum fusca,
sursum clariora, verticillata vel dichotoma. Cellulae conidiogenae solitares, orthotropicae e hyphis, vel plagiotropicae,
plerumque verticillatae e cellulis fertilibus oriuntur; e parte basilari lageniformi vel elongata sursum attenuata et rachide
sympodialiter elongata, denticulata, recta vel flexuosa constant. Conidia simplicia, hyalina vel pigmentata, levia,
subglobosa vel fusiformia, basi apiculata. Chlamydosporae absunt.
Species typica: Chloridium crateriforme van Beyma
Etym.: Gr. akrs = apical, odn = tooth.
Colonies growing rather slowly, appearing velvety to powdery, variously coloured. Aerial hyphae subhyaline,
brownish or olivaceous, smooth- and rather thin-walled, septate, branched, loose or fasciculate. Conidiophores,
when present, arise orthotropically, or plagiotropically when little differentiated, from creeping hyphae, erect or
procumbent, sometimes thickwalled and dark brown at the base, lighter towards the apex, branched verticillately
or dichotomously. Conidiogenous cells solitary, arising orthotropically [p. 24] from undifferentiated hyphae,
often also in plagiotropic whorls on more or less differentiated conidiophores; they consist of a flask-shaped or
elongate basal part, tapering towards the tip, and a sympodial denticulate rachis, straight or somewhat flexuous.
Conidia one-celled, hyaline or pigmented, smooth, subglobose to fusiform, with an apiculate base. No
chlamydospores were observed.
Descriptions are based on colonies growing on 2/o malt agar at 20 C.
Discussion
The genus Acrodontium is separated from Tritirachium on account of the morphology of the conidiiferous
rachis: it is straight or only slightly flexuous, bearing at regular intervals lateral, alternating denticles.
Acrodontium can be subdivided into two sections. Sect. Grisea is similar to Hansfordia Hughes, from which
it differs mainly by the differentiation of the conidiogenous cells into a basal part, tapering towards the tip, and a
long and slender rachis. In Hansfordia the rachis is mostly very short, of equal width as the basal part of the
conidiogenous cell, which is cylindrical, of variable length. The conidia are hyaline, hardly apiculate when
fusiform, and mostly considerably larger than in Acrodontium. Sect. Acrodontium is somewhat similar to
Rhinocladiella Nannf. non Kamyschko.
Many genera with one-celled conidia borne on sympodial conidiogenous cells are known. The following
genera are most similar to Acrodontium: Dicyma Boul. and Gonytrichella Emoto & Tubaki differ by having
conidia with truncate base, and by the conidiogenous cells which are of variable shape and mostly have a short,
wide or irregular rachis; Phaeoisaria Hhnel, Dematophora Hartig and Tharoopama Subram. differ i.a. by the
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presence of distinct synnemata; Geniculosporium Chesters & Greenhalgh and Nodulisporium Preuss have
more or less cylindrical conidiogenous cells with cicatrized rachids, and Rhinocladiella has almost
imperceptable denticles and pleomorphic conidium formation (Schol-Schwarz, 1968).
The names Chloridium Link and Psilobotrys Sacc. cannot be used for this group of fungi. Hughes (1958)
proved their type species to be congeneric with Bisporomyces van Beyma and to produce phialoconidia.
According to the same author, Haplaria Link is a synonym of Botrytis Pers. per Fr., and Trichosporium Fr.,
which was used for several species with sympodial conidiogenous cells by Arnaud (1953) is a nomen
illegitimum.
Rhinotrichella Arnaud (not validly published) seems to be closely related. The original material of the type
species R. grisea (Sacc.) Arnaud (= Rhinotrichum griseum Sacc.) in PAD, however, was too scanty to trace the
mode of conidial production. Both other species of the genus, i.e. R. (?) globulifera Arnaud and R. (?) pilulifera
Arnaud, are possibly more closely related to Rhinotrichum Corda.
Criteria used for the distinction of both sections of Acrodontium are the presence of more or less
differentiated conidiophores, the shape of the conidiiferous rachis and the pigmentation of the conidia. The
appearance [p 25] of the type species of the sections may differ considerably. However, in A. hydnicola the
conidiophores are often integrated in the vegetative mycelium, whereas in A. crateriforme, verticillately
branched fertile hyphae are sometimes present. For this reason it is thought to be preferable to maintain all
species within the same genus.
Key to the species

1a.
1b.
2a.
2b.
3a.
3b.
4a.
4b.
5a.
5b.
6a.
6b.
Conidia guttuliform to fusiform, hyaline; rachis with small, sharp denticles and
often not differentiated from the basal part of the conidiogenous cell (section
Acrodontium)
Conidia globose to guttuliform, with an apiculate base, pigmented; rachis with
blunt denticles and always distinctly differentiated from the basal part of the
conidiogenous cell (section Grisea)
Colonies pink; conidiogenous cells subhyaline, of about the same colour as
the hyphae
Colonies dark olive or gray brown; conidiogenous cells pigmented, darker
than the hyphae
Mature colonies dark olive; conidia smooth
Mature colonies gray brown; conidia often finely verrucose
4
A. salmoneum
3
A. crateriforme
Acrodontium-state of
Ascocorticium
anomalum
Conidiophores integrated in the vegetative mycelium, branched over the

whole length
Conidiophores differentiated, branched verticillately in the upper part only
Conidiogenous cells elongate, widest near the base, tapering towards the tip
Conidiogenous cells swollen, widest somewhat below the middle, mostly
flask-shaped or somewhat elongate, in the latter case dark olive
Conidiophores rather thin-walled over the whole length, light brown,
somewhat flexuous
Conidiophores in the lower part thick-walled, very dark brown, stiff
A. hydnicola
5
A. simplex
6
A. virellum
A. griseum
Acrodontium section Acrodontium

Conidiogenous cells arising orthotropically from undifferentiated hyphae, consisting of an elongate or
subcylindrical basal part, tapering towards the tip, and a somewhat flexuous rachis, which is often not distinctly
separated from the basal part, and bears a series of small, sharp denticles, less than half as wide as the rachis.
Conidia hyaline, guttuliform to lageniform, with an apiculate base. [p. 26]
Acrodontium crateriforme (van Beyma) de Hoog, comb. nov. - Fig. 9
Chloridium crateriforme van Beyma - Zentbl. Bakt. ParasitKde, Abt. 2, 89: 241. 1933 (basionym).
Misapplied name
Chloridium minutum (Sacc.) Sacc. sensu Tubaki - J. Hattori but. Lab. 20: 152. 1958.
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Fig. 9. Acrodontium crateriforme, CBS 144.33. a. conidial structures; b. conidiogenous cells; c. conidia.
Colonies in vitro growing rather slowly, attaining a diameter of 8-15 mm in 8 days, appearing finely floccose,
velvety or lanose, often forming radially furrowed and zonate, vulcano-shaped stromata, gray olivaceous (21"").
Reverse at first subhyaline or herbage green (25i), soon becoming olivaceous black (25""k). Exudate and odour
absent. Submerged hyphae hyaline or nearly so, smooth-walled, 1-2 (-3) m wide. Hyphae of the aerial
mycelium hyaline to pale olive, smooth-walled, 1.5-2.5 m wide, creeping or ascendent, often strongly
fasciculate. Conidiogenous cells scattered, [p. 27] arising orthotropically from undifferentiated hyphae or in
groups of 1-2 (-3) from the tip of a subtending cell, consisting of a subulate basal part, pale olive, widest at the
basal septum or somewhat above, tapering towards the tip, (19-) 13-28 (-38) m long and 2-3 m wide, rarely
with a transverse septum, and a well developed, denticulate rachis, which is often not distinctly differentiated
from the basal part, straight or flexuous up to 45 m long and constantly 1 m wide, with small, sharp denticles
at intervals of 0.3-2.5 m. Conidia hyaline, smooth, guttuliform to somewhat ellipsoidal, sometimes laterally
flattened, with an apiculate base, (3-) 3.5-4.5 (-5) x (1.5-) 2-3 (-4) m. No chlamydospores were observed.
Material examined
CBS 144.33 (= ATCC 15679), type culture of Chloridium crateriforme, isolated as culture contaminant, sent in 1933 by H.
A. Diddens.
CBS 151.58 isolated by G. A. de Vries from sputum of man.
CBS 985.70 isolated from living leaflet of Fraxinus excelsior, Meathrop Wood, Westmoreland, Great-Britain, sent in 1958
by J. C. Frankland.
CBS 839.71 isolated by W. Gams from leaves of Lazuli silvatica, East Lyn River, Devon, Great-Britain, September 1971.
CBS 840.71 isolated from foodstuffs, Bilthoven, sent in 1971 by M. van Schothorst. CBS 841.71 isolated by W. Gams
from rust on Carex sp., near Boxtel, April 1968.
CBS 842.71 isolated by W. Gams from Citrus leaves, Tjibodas Hortus Botanicus, Java, Indonesia, sent in 1969 by J. H.
van Emden.
CBS 843.71 isolated by W. Gams from leaves of Molinia caerulea, near Boatel, April 1968. CBS 844.71 isolated by W.
Gams from dead spider under pine bark.
9. Acrodontium-state of Ascocorticium anomalum (Ellis & Harkn.) Earle - Fig. 10

Rhinotrichella sp., Oberwinkler & al. - Nova Hedwigia 14: 287. 1967.
Colonies in vitro growing very slowly, attaining a diameter of less than 1 mm in 8 days, appearing velvety,
forming a dense stroma, vinaceous buff (17"d). Reverse olivaceous (17"m). Exudate and odour absent.
Submerged hyphae hyaline or nearly so, smooth-walled, 1.5-2 m wide. Hyphae of the aerial mycelium
subhyaline to brown, smooth-walled, 1-2.5 m wide, creeping. Conidiogenous cells scattered, arising
orthotropically from undifferentiated hyphae or in groups of 1-2 from the tip of a subtending cell, consisting of
an acicular, subulate or ventricose basal part, often with a lateral swelling near the base, widest at the base or
somewhat below the middle, tapering towards the tip, sometimes branched, often not separated from the hypha
by a septum, or attached laterally to the hypha, pale brown, (9-) 13-26 (-30) m long and (1.5-) 3-6 (-7.5) m
wide, and a [p. 28] rather irregular, denticulate rachis, which is often not differentiated from the basal part,
straight or flexuous, up to 35 m and 1-2 m wide, with sharp or blunt denticles at intervals of 0.5-3 m. Conidia
hyaline, smooth or finely verrucose, subglobose or guttuliform to ellipsoidal, with a scar at the base, (3-) 3.5-5
(-6) x (2.5-) 3-3.5 (-4N) m. Sometimes irregular, more or less globose chlamydospore-like structures are
present. Perfect state Ascocorticium anomalum, reported by Oberwinkler and al. (1967).
Material examined
CBS 874.71 isolated by F. Casagrande from bark of Pinus sylvestris, Bad Reichenhall, W. Germany, September 1965,
leg. F. Oberwinkler, sent in 1971 by E. Mller under No. ETH-M 7090.
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Fig. 10. Acrodontium-state of Ascocorticium anomalum, CBS 874.71. a. conidial structures; b. conidiogenous cells; c.
conidia. [p. 29]
10. Acrodontium salmoneum de Hoog, sp. nov. - Fig. 11

Coloniae in vitro post 8 dies 6-10 mm diametro, floccosae vel farinosae vel pulverulentae, primo laete persicinae,
deinde obscuriores; reverso luteolo vel luteo. Non olet. Hyphae submersae hyalinae, 1-1.5 m, crassae, saepe dense
intricatae. Hyphae aeriae hyalinae vel pallide salmoneae, leves, 1.5-2.5 m crassae, repentes vel adscendentes, saepe
fasciculatae. Cellulae conidiogenae sparsae, orthotropicae e hyphis vegetativis vel ex apice cellularum lateralium singulae
vel ternae (quaternae) oriuntur, e parte basilari fere cylindrica, levi, subhyalina, sursum modice attenuata, saepe curvata,
plerumque 15-40 x 1.5-2.5 m et rachide tenuiter denticulata, recta vel flexuosa, 1.5 m crassa, ad 35 m longa, a parte
basilari vix delimitata constant. Conidia hyalina, levia, ovoidea, fusiformia vel ellipsoidea, basi apiculata, (3.5-) 4.5-5.5 (-7)
x (1.5-) 2-3 (-3.s) m. Cellulae fere inflatae, globosae, tenui-tunicatae, ad 5 m diametro, chlamydosporarum similes in
mycelio submerso adsunt. Status perfectus ignotus.
Typus: CBS 847.71, isolatus e sputo humano, Groningen, ab A. Kikstra.
Colonies in vitro growing rather slowly, attaining a diameter of 6-10 mm in 8 days, appearing floccose or
farinose to powdery, at first peach (7f), when older up to salmon (9d). Reverse pale luteous (19d) or luteous
(17). Exudate and odour absent. Submerged hyphae hyaline, 1-1.5 m wide, often somewhat stromatic.
Hyphae of the aerial mycelium hyaline to pale salmon, smooth-walled, 1.5-2.5 m wide, creeping or ascendent,
often fasciculate. Conidiogenous cells scattered, arising orthotropically from undifferentiated hyphae or in
groups of 1-3 (-4) from the tip of a short subtending cell, consisting of a subcylindrical basal part, subhyaline,
widest near the base or around the middle, slightly tapering towards the tip, rarely somewhat flexuous, mostly
15-40 m long and 1.5-2.5 m wide, rarely with a transverse septum, and a well developed, denticulate rachis,
which is mostly not differentiated from the basal part, straight or somewhat flexuous, up to 35 m long and 1.5
m wide, slightly tapering towards the tip, with rather sharp denticles at irregular intervals. Conidia hyaline,
smooth, fusiform to ovoidal, with an apiculate base, (3.5-) 4.5-5.5 (-7) x (1.5-) 2-3 (-3.5) m. Hyaline, globose
(about 5 m in diameter), solitary, rather thin-walled chlamydospore-like structures are very rare in the
submerged mycelium. No perfect state was observed.
Material examined
CBS 847.71 (type) isolated from human sputum, sent in 1971 by A. Kikstra under No. 2.
CBS 580.67 isolated as culture contaminant, Houston, Texas, USA, sent in 1967 by C. L. Keswami as Cephalosporium
sp.
CBS 846.71 (= NRRL 5290) sent in 1971 by D. I. Fennell under No. A-13810.
CBS 848.71 isolated from soil of beech forest, Soiling, W. Germany, sent in 1971 by A. von Klopotek under No. R-10.
Discussion
The species is somewhat reminiscent of some conidial states of Xylariaceae, [p. 29] which are mostly
known as Nodulisporium or Geniculosporium. In these genera the conidiogenous cells are either not elongating,
producing conidia at their swollen tips, or the rachis is geniculate and cicatrized, producing more or less truncate
conidia. Some representatives of the heterogeneous pink-coloured group of Nodulisporium are morphologically
intermediate between the latter genus and Acrodontium sect. Acrodontium, e.g. the conidial state of
Anthostomella spartii Berl. & Vogl. In this species the conidia may be to 4-celled and the rachis is rather short,
with the denticles placed at random, not alternating. The present species is described in Acrodontium mainly
because of the long and slender rachis and the constant shape of the conidia and conidiogenous cells.
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Fig. 11. Acrodontium salmoneum, CBS 847.71. a, b. conidial structures; c. conidia.
Acrodontium section Grisea de Hoog, sect. nov.

Conidiophora composita plerumque orthotropica e hyphis repentibus oriuntur, seu [p. 31] erecta crassi-tunicata et
deorsum fusca, sursum clariora, seu procumbentia in hyphas vegetativas transeuntia, verticillata vel simplicia. Cellulae
conidiogenae e hyphis fertilibus vel e hyphis aeriis vegetativis oriuntur, e parte basilari lageniformi vel elongata et rachide
angustiore recta denticulis obtusis obtecta constant. Conidia modice pigmentata vel subhyalina, subglobosa vel
guttuliformia, basi apiculata.
Species typica: Tritirachium heimii (Saccas) Langer. var. griseum Fassatiov.
Differentiated conidiophores mostly arising from creeping hyphae, either erect, thick-walled and dark brown
in the lower part, lighter towards the apex, or procumbent and sometimes not clearly distinguishable from the
vegetative mycelium; with verticillate or simple branching. Conidiogenous cells as distal branches of
conidiophores or arising orthotropically from undifferentiated hyphae, consisting of a flask-shaped or elongate
basal part, tapering towards the tip, and a straight rachis, which is differentiated from the basal part and bears a
series of blunt denticles, the latter being about as wide as the rachis. Conidia somewhat pigmented to
subhyaline, globose or guttuliform, with an apiculate base.
11. Acrodontium hydnicola (Peck) de Hoog, comb. nov. - Fig. 12
Virgaria hydnicola Peck - Rep. N. Y. St. Mus. 42: 32. 1889 (basionym) = Tritirachium hydnicola (Peck) Hughes Can. J. Bot. 31: 604. 1953.
? Clonostachys dichotoma Bayliss Elliott - Trans. Br. mycol. Soc. 6: 56. 1917.
Colonies in vitro growing slowly, attaining a diameter of 5 mm in 8 days, appearing powdery-velvety, at first
rosy buff (17d), when older cinnamon (15"i). Reverse cinnamon (15"i). Exudate and odour absent. Submerged
hyphae hyaline, 1.5-3.5 m wide, often forming a compact stroma. Hyphae of the aerial mycelium hyaline to
brownish, 1.5-2.5 m wide. Conidiophores arising orthotropically, or plagiotropically when little differentiated,
from creeping hyphae, procumbent, hyaline in the lower part, brown towards the apex, smooth- and rather
thin-walled, stalk up to 175 m high and 2-3 m wide, branched verticillately or dichotomously over the whole
length at intervals of 10-20 m.; often one of the branches forms a conidiogenous cell while the other ramifies
again. Conidiogenous cells arising plagiotropically as a part of compound, more or less differentiated
conidiophores, often also orthotropically from undifferentiated hyphae, consisting of an elongate basal part,
pale brown, slightly swollen at the base and tapering towards the tip, (15-) 18-30 (-42) m long, (1.5-) 1.5-2.5 (-3)
m wide, and a well developed, denticulate rachis, up to 25 m long and mostly 1 m wide, straight or slightly
flexuous, with blunt denticles about 1 m long, at intervals of 0.5-1.5 m. Conidia subhyaline, smooth,
thin-walled, globose to subglobose, with slightly apiculate base, (1.5-) 2-3 (-3.5) x (2-) 2-2.5 (-3) m. No
chlamydospores were observed. Perfect state unknown. [p. 32]
Fig. 12. Acrodontium hydnicola. a. conidial structures in culture; b, c. from type collection, conidial structures on natural
substrate; d. conidial structures in culture; e. conidiogenous cells; f. conidia.
Material examined
Herbarium specimen
Virgaria hydnicola in herb. NYS (type), on resupinate basidiomycete on wood, North Elba, USA.
Living strain
CBS 349.55 isolated as contaminant on keratin, sent in 1955 by F. Blank.
Discussion
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The conidiophores are often little differentiated from the vegetative mycelium. In poorly developed
subcultures on PDA they may be indistinguishable, [p. 33] more or less irregularly branched.
No material of Clonostachys dichotoma was available in K and BPI; probably it is lost. According to the
original description and illustration there is much similarity with A. hydnicola.
Isaria acaricida Pat. is very closely related. It is rather dark and firm and its forms small synnemata. Without
living cultures its identity cannot be established. No type material could be examined, only a secondary
collection from the same substrate and locality (B) was studied.
12. Acrodontium simplex (Mangenot) de Hoog, comb. nov. - Fig. 13
Beauveria simplex Mangenot - Rech. method. champ. certains bois dcomp. p. 40.1952 (basionym); Revue gen. Bot.
63: 454. 1952.
Fig. 13. Acrodontium simplex, CBS 127.53. a, b. conidial structures; c. conidiogenous cells; d. conidia.
Colonies in vitro growing slowly, attaining a diameter of 4 mm in 8 days, appearing velvety, at first
vinaceous buff (17"f), when older hazel (17"). Reverse fuscous (13 ""k), paler towards the margin. Exudate and
odour absent. Submerged hyphae hyaline, smooth- and thin-walled, 1.5-3 m wide. Hyphae of the aerial
mycelium hyaline to brownish, smooth-walled, 1-2 m wide. Conidiophores arising orthotropically from creeping
hyphae, suberect, sometimes not markedly differentiated, brown, smooth and slightly thick-walled, stiff, stalk up
to 110 m high and 1.5-3 m wide, the upper part bearing 1-2 (3) whorls of 1-2 (3) side branches or 1-4
conidiogenous cells. Conidiogenous cells arising plagiotropically as a part of compound conidiophores, often
also orthotropically from undifferentiated hyphae, consisting of an elongate basal part, sometimes swollen at the
base up to 3 m tapering towards the tip, 12-28 m long, and a well developed, denticulate rachis, up to 30 m
long and mostly 1 m wide, straight or slightly flexuous, tapering towards the tip, with blunt denticles about 1 m
long at intervals of 0.5-1 m. Conidia brownish, smooth, slightly thickwalled, subglobose to guttuliform, with an
apiculate base, (1.5-) 1.5-3 (-3.5) x (1-) 1.5-2.5 (-3) m No chlamydospores were observed. Perfect state
unknown.
Material examined
CBS 127.53, type culture of Beauveria simplex, isolated from decaying trunk of Fraxinus excelsior, France, sent by F.
Mangenot under No. 1084.
CBS 845.71 isolated by W. Gams from Oligoporus rennyi on pine wood, Kampina heath, near Boxtel, September 1971.
13. Acrodontium virellum (Fr.) de Hoog, comb. nov. - Fig. 14

Botrytis virella Fr. - Summa Veg. Scand. p. 491. 1827 (basionym) Beauveria virella (Fr.) Mangenot - Rech. method.
champ. certains bois dcomp. P. 36. 1952; Revue gen. Bot. 63: 450. 1952.
Colonies in vitro growing slowly, attaining a diameter of 5 mm in 8 days, appearing velvety to somewhat
powdery, at first rosy buff (15"f), when older up to sepia (13"k). Reverse sepia 17"k). Exudate and odour
absent. Submerged hyphae hyaline, 1-3.5 m wide, often forming a compact stroma. Hyphae of the aerial
mycelium hyaline, or pale brownish when bearing conidiophores, 1.5-4 m wide. Conidiophores arising
orthotropically from creeping hyphae, ascendent to suberect, not markedly differentiated, brown, paler near the
base, hyaline towards the tip, smoothand somewhat thick-walled, with thin transverse septa, stalk up to 650 m
long and (2-) 3-4.5 (-6) m wide, the upper part bearing 2-5 (7) whorls of 1-3 side branches or conidiogenous
cells; side branches bearing 1-3 (5) whorls of conidiogenous cells or branches of second order, hyaline or nearly
so. Conidiogenous cells arising plagiotropically as a part of compound conidiophores, often also orthotropically
from undifferentiated hyphae, consisting of a flask-shaped to elongate basal part, tapering towards the tip and
sometimes constricted in the upper part, subhyaline, mostly 4-8 x 2-3 m, and a wel developed, denticulate
rachis, up to 5 m long and 1-2 m wide, straight or slightly flexuous, with blunt denticles [p. 35] about 1 m long
at intervals of 0.5-1 m. Conidia light brown, globose to subglobose, sometimes guttuliform, with an apiculate
base, (2-) 2.5-3.5 (-3,5) x (1.5-) 2-3 (-3) m. No chlamydospores were observed. Perfect state unknown.
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Fig. 14. Acrodontium virellum, CBS 129.53. a, b. conidial structures; c. conidia.
Material examined
Herbarium specimen
Beauveria heimii Saccas in herb. PC, on bark of Pinus halepensis, Mallorca, Spain, leg. G. Malenon, November 1970.
Living strain
CBS 129.53 isolated from decaying wood, Argonne, France, 1952, sent in 1953 by F. Mangenot as Beauveria virella.
Discussion
No type material is left by Fries in UPS. The characters given in the original description are hardly sufficient to
ascertain its identity. The present description is based on strain CBS 129.53 [p. 36].
14. Acrodontium griseum (Fassatiov) de Hoog, comb. nov. - Fig. 15
Tritirachium heimii (Saccas) Langer. var. griseum Fassatiov - Cesk Mykol. 25: 116. 1972 (basionym).
Fig. 15. Acrodontium griseum, CBS 671.70. a, b. conidial structures; c. conidiogenous cells; d. conidia
Colonies in vitro growing rather slowly, attaining a diameter of 4-7 mm in 8 days, appearing powdery to
somewhat velvety, often forming small sporodochium-like cushions, at first vinaceous buff (17 b), when older
herbage green (23m). Reverse herbage green (23m) to olivaceous black (21""m). Exudate absent, odour very
slightly sourish. Submerged hyphae hyaline, smooth- and thin-walled, 1-3.5 m wide, closely septate. Hyphae of
the aerial mycelium hyaline or brownish, smooth-walled, 1.5-3 m wide. Conidiophores arising orthotropically
from creeping hyphae, erect, markedly differentiated, brown, dark brown to red brown towards the base,
smooth- and thick-walled, stiff, stalk (20-) 50-310 (-450) x 3-4.5 m the upper part bearing 2-4 (5) whorls of 2-3
(4) side branches or conidiogenous cells; branches of second or third order often present. Conidiogenous cells
arising plagiotropically as a part of compound conidiophores, often also orthotropically from undifferentiated
hyphae, consisting of a flask-shaped or elongate basal part, tapering towards the tip, pale brown, mostly 6-12
m [p. 37] long, swollen up to 5 m below the middle, and a well developed, denticulate rachis, up to 3 5 m
long and mostly 1 m wide, straight or somewhat flexuous, with blunt denticles about 1.5 m long at intervals of
0.5-1 m. Conidia pale olive brownish, smooth, globose, subglobose, guttuliform or fusiform, with an apiculate
base, (1.5-) 2-4.5 (-5.5) x 2-3 m No chlamydospores were observed. Perfect state unknown.
Material examined
CBS 671.70, type culture of Tritirachium heimii var. griseum, isolated from wood, Bohemia, Czechoslovakia, sent under
No. 1169.
CBS 849.71 isolated from peat, Oldenburg, W. Germany, July 1970, sent in 1971 by A. von Klopotek under No. T-23.
Doubtful or excluded species of Beauveria, Tritirachium and similar genera

Tritirachium cinnamomeum van Beyma - Antonie van Leeuwenhoek 8: 116. 1942.
The conidiophores are subhyaline, suberect and irregularly verticillate. The conidiogenous cells are
cylindrical, bearing an irregularly geniculate rachis. It is reminiscent of the genus Nodulisporium and should
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better be referred to as Nodulisporium cinnamomeum (van Beyma) de Hoog, comb. nov.

Gonatorrhodiella coccorum Petch - Trans. Br. mycol. Soc. lo: 181. 1924 = Beauveria coccorum (Petch)
Linder - Lloydia 5: 206. 1942.
Rhinotrichum album Petch - Trans. Br. mycol. Soc. II: 258. 1925.
Petch (1924b, 1925) described the same species twice, under different names. In 1931 he discovered his
error and maintained the oldest name. The fungus has smooth, hyaline, creeping hyphae of variable width, with
numerous short, blunt denticles. These give rise to guttuliform conidia. On the dry specimens (K) it could not be
decided whether the conidia were born in chains as described by Petch (1931). The fungus is congeneric with
Oospora meliolae Hansf.
When Linder (1942) made the combination Beauveria coccorum, he gave a description which does not
correspond with the type material. Somewhat further he mentioned the name Beauveria coccospora (Petch)
Linder, which is presumably a misprint.
Racodium (?) entomogenum Pers. - Mycol. eur. I: 72. 1822.
Persoon (1822) thought this species to be related to Sporotrichum densum Link. The type material (L),
however, did not allow to recognize the fungus. [p. 38]
Beauveria cretacea (van Beyma) Matsushima - Microf. Solomon Isl., Papua-New Guinea p. 7. 1971.
= Isaria felina (DC. per Fr.) Fr.
Beauveria heimii Saccas - Revue Mycol. 13: 63. 1948 = Tritirachium heimii (Saccas) Langer. -Revue
Mycol. I4: 13 5. 1949.
No material is left in PC. The species was published without a Latin diagnosis. Most likely an Acrodontium
species, such as A. griseum or A. simplex, was concerned.
Sporotrichum paranense Marchionatto - Physis, B. Aires II: 348. 1933 = Beauveria paranensis
(Marchionatto) Gsswald - Arb. biol. BundAnst. Land- u. Forstw. 22: 434. 1939 = Paecilomyces paranensis
(Marchionatto) G. Mller - Wiss. Z. Humboldt-Univ. Berlin, math.-nat. R. I4: 780. 1965.
According to the type material (CMI) the species has phialides.
Beauveria petelotii Vincens -Bull. Soc. bot. Fr. 61: 132. 193 S.
No material is left in PC. The species was described from various insects as being very variable. Possibly
different Beauveria species were described under the same name (Fassatiov, 1966).
Botrytis rileyi Farlow in Riley - Rep. U. S. Dep. Agric. 1883: 121 = Beauveria rileyi (Farlow) Gsswald - Arb.
biol. BundAnst. Land- u. Forstw. 22: 434. 1939 = Spicaria rileyi (Farlow) Charles - Mycologia 28: 398. 1936.
According to the type material (FH) the species has phialides.
Verticillium rubrum Baquis - Annali Ottal. Clin. ocul. 34: 947. 1905 = Beauveria rubra (Baquis) Langer. - Bull.
Acad. Med., Paris III: 133.1934 = Tritirachium rubrum (Baquis) Langer. - Annls Parasit. hum. comp. 22: 98.
1947.
No material was left in PAD and PC. The original description was very scanty. Doubtful species.
Sporotrichum flavicans Fr. var. spicatum Ferr. in Ferr. & Massa - Annls mycol. 10: 295. 1912 = Tritirachium
spicatum (Ferr.) Limber - Mycologia 32: 29. 1940 = Beauveria spicata (Ferr.) Saccas - Revue Mycol. 13: 64.
1948.
No material left in PAD. Possibly a Nodulisporium-like fungus was meant.
Trichophyton viannai de Mello - Indian J. med. Res. 5: 228. 1917 = Beauveria viannai (de Mello) Langer. Bull. Acad. Md., Paris III: 133. 1934 = Tritirachium viannai (de Mello) Langer. - Annls Parasit. hum. comp. 22:
98. 1947.
No material is left in HCIO and PC. According to the description and illustrations a species reminiscent of
Tritirachium oryzae was meant. [p. 39]
Acknowledgements
The author is indebted to Dr J. A. von Arx for his valuable suggestions and to Dr M. A. Donk for his advice
concerning nomenclature. I am much obliged to Dr W. Gams for his stimulating help in preparing the manuscript
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and much editorial work and to Drs R. A. Samson for his helpful cooperation and the inspiring discussions on
the subject. Mr D. Yarrow and Dr K. M. Old are acknowledged for correcting the English text.
References
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De HOOG 1972 TThe Genera Beauveria, Isaria, Tritirachium and Acrodontium

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THE GENERA BEAUVERIA, ISARIA, TRITIRACHIUM AND ACR...

Studies in Mycology, No. 1

The genera Beauveria, Isaria, Tritirachium and Acrodontium gen. nov

THE GENERA BEAUVERIA, ISARIA, TRITIRACHIUM AND ACR...

Key to the genera and sections discussed in the present paper

Conidia globose, ellipsoidal or subcylindrical, mostly with a rounded base,

THE GENERA BEAUVERIA, ISARIA, TRITIRACHIUM AND ACR...

Key to the species

Conidiogenous cells mostly in whorls along ascendent hyphae

Conidia globose or subglobose, in pure culture sometimes broadly ellipsoidal; stalk

1. Beauveria bassiana (Bals.) Vuill. - Fig. 2 and 3

THE GENERA BEAUVERIA, ISARIA, TRITIRACHIUM AND ACR...

Fig. 2. Beauveria bassiana on natural substrate, conidial structures.

THE GENERA BEAUVERIA, ISARIA, TRITIRACHIUM AND ACR...

THE GENERA BEAUVERIA, ISARIA, TRITIRACHIUM AND ACR...

THE GENERA BEAUVERIA, ISARIA, TRITIRACHIUM AND ACR...

THE GENERA BEAUVERIA, ISARIA, TRITIRACHIUM AND ACR...

Fig. 5. Beauveria alba, CBS 570.71. a, b. conidial structures; c. conidia.

Colonies in vitro attaining a diameter of 14-16 mm in 8 days, appearing lanose to floccose, up to 2 mm

THE GENERA BEAUVERIA, ISARIA, TRITIRACHIUM AND ACR...

Key to the species

Conidiogenous cells flask-shaped, often curved; conidia broadly ellipsoidal

4. Isaria felina (DC. per Fr.) Fr. - Fig. 6

Fig. 6. Isaria felina. a. conidial structures; b. conidiogenous cells; c. conidia.

THE GENERA BEAUVERIA, ISARIA, TRITIRACHIUM AND ACR...

Perfect state unknown.

5. Isaria orthopterorum Petch - Fig. 7

Fig. 7. Isaria orthopterorum, from type collection. a, b. conidial structures; c. conidia.

THE GENERA BEAUVERIA, ISARIA, TRITIRACHIUM AND ACR...

Key to the species

6. Tritirachium dependens Limber - Fig. 8 a-d

Living cultures were not available.

THE GENERA BEAUVERIA, ISARIA, TRITIRACHIUM AND ACR...

7. Tritirachium oryzae (Vincens) de Hoog, comb. nov. - Fig. 8 e-h

THE GENERA BEAUVERIA, ISARIA, TRITIRACHIUM AND ACR...

Acrodontium de Hoog, gen. nov.

THE GENERA BEAUVERIA, ISARIA, TRITIRACHIUM AND ACR...

Key to the species

Conidiophores integrated in the vegetative mycelium, branched over the

Acrodontium section Acrodontium

THE GENERA BEAUVERIA, ISARIA, TRITIRACHIUM AND ACR...

9. Acrodontium-state of Ascocorticium anomalum (Ellis & Harkn.) Earle - Fig. 10

THE GENERA BEAUVERIA, ISARIA, TRITIRACHIUM AND ACR...

10. Acrodontium salmoneum de Hoog, sp. nov. - Fig. 11

THE GENERA BEAUVERIA, ISARIA, TRITIRACHIUM AND ACR...

Fig. 11. Acrodontium salmoneum, CBS 847.71. a, b. conidial structures; c. conidia.

Acrodontium section Grisea de Hoog, sect. nov.

THE GENERA BEAUVERIA, ISARIA, TRITIRACHIUM AND ACR...

13. Acrodontium virellum (Fr.) de Hoog, comb. nov. - Fig. 14

THE GENERA BEAUVERIA, ISARIA, TRITIRACHIUM AND ACR...

Fig. 14. Acrodontium virellum, CBS 129.53. a, b. conidial structures; c. conidia.

Doubtful or excluded species of Beauveria, Tritirachium and similar genera

THE GENERA BEAUVERIA, ISARIA, TRITIRACHIUM AND ACR...

better be referred to as Nodulisporium cinnamomeum (van Beyma) de Hoog, comb. nov.

THE GENERA BEAUVERIA, ISARIA, TRITIRACHIUM AND ACR...

THE GENERA BEAUVERIA, ISARIA, TRITIRACHIUM AND ACR...

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