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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
Fassatiová, 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, some- what
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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Cornell Univ. 1931 = Beauveria shiotae (Kuru) Langer.Rapp. Soc.Zentbl.) Delacr. ed. in Brumpt Précis Parasitol. math. Sporotrichum globuliferum Speg.Univ.Fig.. 1923 = Beauveria paradoxa (Bals. Séanc.. Stn 97: 399.J.Bull. Beauveria doryphorae Poisson & Patay . Br.Fl. . Suppl.Cellule 33: 101.) Biourge . 1931.Wiss. . Beauveria bassiana Bals. Sci.Syll. 1912.Bull. 6] 04/03/2015 16:32 . . Edin.Gazetta di Milano. .) Delacr. 10: 249.Syst.. Cornell Univ. Conidia one-celled. 16: 58. Bakt. natn.htm basal part. Berlin. and a geniculate. hebd. hebd. comp. sometimes subglobose. Abstr. and conidiogenous cells rather slender and rarely clustered 2b. Soc. Agric. Sporotrichum sulfurescens van Beyma . stalk cells. Hist. subsp. per Pers. tenella Sacc. 32: 8231954. brongniartii 1. 1935. 3: 459.) Vuill. 1914 = Penicillium (Spicaria) densum (Link per Pers. Paris 113: [p. Beauveria bassiana (Bals. No chlamydospores or perfect states were observed. Conidiogenous cells mostly in whorls along ascendent hyphae Conidiogenous cells scattered or in irregular clusters on swollen stalk cells 2a. argent. bot.) Mont.) Vuill. . St. Nauk Warszawa 6: 35. crypt. cient. 1832] = Isaria densa (Link per Pers. R. p. Br. eur. 1928 = Beauveria sulfurescens (van Beyma) J. globose to ellipsoidal. . natn. Botrytis paradoxa Bals.) Vuill. Botrytis effusa Beauv. Séanc. Br. r.) Picard Annls Éc. N. 1936 = Tritirachium shiotae (Kuru) Langer.) Ramsb. Aires 20: 449. Müller .Archwm Nauk biol. Nancy : 153. stalk cells and globose to flask-shaped conidiogenous cells mostly forming dense clusters Conidia mostly ellipsoidal. 1923 = Beauveria stephanoderis (Bally) Petch . 1879 = Sporotrichum larvicolum Peck . 26: 16. 9: 183. argent.An. . Séanc.Bull.Michelia 2: 544. 1937 (incidentally mentioned) = Beauveria tenella (Sacc. Freunde. Botrytis necans Massee . Fr.) Vuill. 1839.C.C. misc. 10: 278. Fr. Isaria citrinula Speg. hyaline.An. Wet. Sci. J.. Soc.Syll. Acad.-nat.Bull. 22: 98. Bot. 3 sur 22 http://www.1941 (incidentally mentioned). 10: 249. Ser. Akad.Can. St. 1941. Inf. Séanc. Linnaea 10: 611.) Biourge . 1b. 1881.Rep. 1910 = Beauveria bassiana (Bals. 1: 76. Spec. 1914.Bull. Sci.C. Nancy 11 : 15 3. 1923. 20: 540. var. Botrytis brongniartii Sacc. 2. smooth.Annls Parasit. Sci. N. Fung. Abt. per Pers. Soc. R. 19 June 18 (nomen provisorium) = Botrytis bassiana Bals.) Biourge . Montpellier 13: 200. mycol.Meded. . mycol. 1856 = Spicaria bassiana (Bals. 1947. hum. J. alba 2 B. Montpellier 13: 203. TRITIRACHIUM AND ACR. subsp.Bull. r. B. 1891 = Spicaria densa (Link per Pers. 1891 (incidentally mentioned) = Beauveria tenella (Sacc.Trans. Acad. Nat.) Fr. thin-walled. nat. Gen. Étud. . denticulate rachis. lunzinensis Szilvinyi . 1970. Soc. 2 and 3 Sporotrichum densum Link ..) Giard .) Vuill.Mycol.Cellule 33: 102. 14: 775. Paris 112: 1272. Mus. agric.Mycologia 62: 820. K. ParasitKde. Y. . Trans. . nac. p. 5 p.Bull. cient. Sci. in pure culture sometimes broadly ellipsoidal. Soc.. Soc. 1835 = Stachylidium bassianum (Bals.An. Humboldt . ri: 123. 9. bassiana B. 1824) = Beauveria effusa (Beauv. Lyon 14: 25.knaw.Cellule 33: 102.. Sci. sensu Pettit) G. 1910 = Beauveria densa (Link per Pers. 1892 = Botrytis delacroixii (Sacc. Agric. . 1965 Botrytis bassiana (Bals. delacroixii Sacc. Botrytis stephanoderis Bally in Friederichs & Bally . Trans. mycol. Exp. 1: 18.cbs. Soc. 59: 40. 1911 [non Botrytis effusa Grev. 1880 = Beauveria globulifera (Speg. . Taylor .Trans. Nancy 11: 1S3. Afd. Pettit in Bull.) Siemaszko . Mus. Mus. Beauveria laxa Petch . Paris 200: 961.Mag. Sporotrichum minimum Speg.) MacLeod . . 1924. . 1910. 59: 40. Agric. Sporotrichum larvatum Peck . Soc. Berlin 3: 13.) Vuill. B.. 1895 = Trichoderma minimum (Speg.Trans. Gdns Kew 1914: 159. Lab. Séanc. Bull. 103: 178. Soc. Sci. naturf.) Vuill. J. med. 1822 [non Sporotrichum densum (Ditm. 468. Key to the species 1a. 1887 (substitute name). 1910 = Penicillium delacroixii (Sacc. mycol. 1882 = Botrytis tenella (Sacc. . Isaria vexans Pettit . bot. Br. . mycol. .Verb. KoffiebessenboeboekFonds 6: 106. 1893 = Spicaria delacroixii (Sacc. 1809.Annls Éc. Exp. Conidia globose or subglobose.) Giard . 32: 44. 25:436. Tow. Ges. mycol. 1912 = Penicillium bassianum (Bals. Paris 1: 164. agric. Acad. Séanc. Y. [p. Descriptions are based on colonies growing on oatmeal agar at 20° C. hebd. 5] 270.nl/publications/1001/content_files/content.. Type species: Beauveria bassiana (Bals.) Picard . 1891 = Isaria tenella (Sacc. Z.. if present.Bull. 1895. Soc. prat..) Petch .Jap.THE GENERA BEAUVERIA. Isaria shiotae Kuru . Stn 97: 362. ISARIA. Soc. 2: 351. bot. . Soie. r. ned. 301.) Vuill. 1924. Fr. 1923 = Beauveria delacroixii (Sacc.

Hyphae of the aerial mycelium hyaline. leg. on larva of Coleoptera indet. Beauveria laxa in herb. TRITIRACHIUM AND ACR. H. tenella in herb. Thailand. creeping or ascendent. 21669. 1. Sporotrichum minimum in herb. Beauveria bassiana on oatmeal agar. No chlamydospores were observed. 1927/28. conidia. 21686 (type). in older strains branching becomes less tight.32. Hoge Veluwe. mostly 2. Argentina. Conidia hyaline or rarely yellowish. PAD.7] they bear groups of swollen lateral cells. Botrytis bassiana subsp. 2. b. c. smooth. January 1927. up to 1 µm long. type culture of Botrytis stephanoderis. Java.5-) 2-2. sent in 1933 by H. denticles as wide as the rachis.. up to 20 µm long or even longer and rather constantly 1 µm wide.. sometimes funiculose. [p... sent in 1930 by A. 5] Colonies in vitro attaining a diameter of 6-23 mm in 8 days.nl/publications/1001/content_files/content. conidial structures. Mycelium on natural substrate typically cushion-like. type culture of Isaria shiotae. Buenos Aires. Tucumán. Germany.30 (= ATCC 9453) isolated from Bombyx mori. GreatBritain. 188o. Botrytis necans in herb.cbs.25 isolated from Oecophylla smaragdina. Fig. rarely reddish. Fetch (K): R-114 (type). [p. LPS. on dead larva under alder bushes. April 1906. sometimes funiculose or rarely producing synnemata. 21683. the conidiogenous cells occuring in dense.. isolated from Stephanoderis hampei. Reverse uncoloured or yellowish to pinkish. occasionally reddish. which is tightly clustered.5-3. up to 5 mm. Nuwara Eliya. which fits Pettit’s description. on Coleoptera indet. Conidiogenous cells consisting of a globose to flask-shaped. on insect larva on Indigofera endecaphylla. isolated as culture contaminant. July 1881. USA. high. isolated from human thorax abcess. smooth-walled. 21675. . Adirondack. 11215 (type). Ceylon. rarely forming synnemata (‘Rhizomorphen’. B (type). [p. 4 sur 22 http://www. probably authentic material. Basal parts of the conidiogenous cells globose. (1. Burkill. yellowish.. NYS (type). 21671. on Acridium sp. Sporotrichum globuliferum in herb. Massachusetts. Blunck. leg. CBS 124. Material examined Herbarium specimens Sporotrichum densum in herb. a. LPS: 21680 (type). Isaria vexans in herb. 3. A perfect state without name was reported by Schaerffenberg (1955). Sporotrichum larvatum in herb. 1-2 µm wide. Living strains CBS 110. Isaria citrinula in herb. odour absent. mostly 3-6 x 3-5 µm. S. geniculate or irregularly bent. sometimes elongate basal part. granular-pulverulent. R-374 and R-474. F. on insect. January 1928. type culture of Sporotrichum sulfurescens. velvety to powdery. denticulate.5-4 x 2-3 µm. and a well developed rachis. cocoons of caterpillar. on Coccinella sp. LPS. Ceylon. 1919.htm Fig. or arise directly from the hyphae.5 µm. CBS. floccose..Singapore. van Vloten. Submerged hyphae hyaline. on unidentified chrysalid.THE GENERA BEAUVERIA. Exudate rarely produced. on Coleoptera indet.26. on Coleoptera indet. on Formica sp. CBS 337.5-) 2-3 (-4) x (1. Wellaway. at first white. leg. on insect. Rostock. CBS 118. respectively. K (type). Italy. and surrounding decaying wood. 2 specimens on Lepidoptera indet. Kirimettia.27 (= ATCC 7159). R-371. conidiogenous cells. appearing lanose. In fresh isolates the conidial apparatus is tightly clustered. Petch as Beauveria globulifera. 1939). Guillermond as Beauveria effusa. A.33 isolated from Brachyderes incanus.5 (-3) µm. Padova. 1914. Beauveria bassiana on natural substrate. sometimes with an apiculate base. globose to broadly ellipsoidal. sent in 1925 by T. R-401. globose heads only. G.. on Brachartonia catoxantha. Japan. on Vespa sp. smooth-walled. mostly 3-6 x 2. Atkinson (CUP). terminal cells mostly more slender. the conidiogenous cells occur in small groups or solitarily on ellipsoidal to subcylindrical lateral cells (measuring up to 15 x 6 µm). on coccid. G.. Beauveria bassiana in herb. Alexis. Argentina. 1971.5-3 µm wide. San Jose de Flores. Argentina. 04/03/2015 16:32 . cocoons of unidentified insect and cocoons of Lepidoptera indet. R-333. conidial structures. de Vries.. subglobose or somewhat flask-shaped. 8] CBS 119. CBS 209. O. later becoming yellowish. which by further branching give rise to smaller swollen cells or 1-5 conidiogenous cells in the first or second order. ISARIA. Hyphae of the aerial mycelium bearing a conidial apparatus as described above.knaw.

from Laboratoire de Cryptogamie. Gams from the basidiomycete Cyphellopsis anomala. and has to be rejected according to Art. On artificial media most of the strains lose their pathogenicity and degenerate. sent in 1971 by R.36 from Blastophagus piniperda.5 µm in diameter. CBS 133.68 isolated from Laodelphax striatellus. Because of the synnematous growth he made the combination Isaria densa and placed Botrytis tenella in the synonymy. Israël. N. Many authors mentioned collections under this name. Yugoslavia. CBS 127. 2400 and No.71 isolated from human sputum. So the epithet densum Link has been used for two Beauveria species. This was the best known name for the Beauveria species with globose conidia. bassiana.36 from Orthotomicus laricis. but as the author wanted to dedicate the species to its discoverer. CBS 651. whereas in the typical subspecies they were about 3 µm in diameter. Kenneth.67 isolated from Eurygaster sp. Moreover the conidiogenous cells tend to elongate and become narrower. Groningen. sent in 1970 by R. France. bassiana are discussed below in chronological order. CBS 465. Müller as Sporotrichum epigaeum var. 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.knaw. Sporotrichum globuliferum. Sporotrichum minimum. terrestre Daszewska. Bresadola 04/03/2015 16:32 . CBS 128. Patay from Leptinotarsa decemlineata.68 isolated from insect on leaf of Vicia sp. Israël. all strains isolated in Poland. both sent in 1967 by P.36 from Pentatoma rufipes. Botrytis bassiana. A second collection of this species was mentioned by Pettit (1895). 1931). Finland. 69 of the International Code of Botanical Nomenclature as a nomen confusum.71 and 652. According to some figures on the envelope. Botrytis bassiana subsp. Brazil.71 isolated by A. occurring rarely on other animals (MacLeod. CBS 212. type culture of Beauveria doryphorae.50 (= LCF 351) isolated from Bombyx mori. Kikstra. Ilanot. Sporotrichum densum. and CBS 640. Most of the names considered here to be synonymous with B. Kenneth under No. CBS 131. Discussion The species is a facultative but highly virulent parasite on insects. CBS 715. Kahanpää. though their size may vary considerably in strains in which sporulation has decreased. 5 sur 22 http://www. A. 2660.nl/publications/1001/content_files/content.67 and 325. Rehovot.htm CBS 127. Marchionatto. Daams in Ankeveen and Kortenhoef. The type of the latter is in bad condition. le Moult. sent by G. CBS 123. A.. Samson from insects collected by J. Israël. CBS 120. and Cleonus punctiventris. isolated by R. sent in 1950 by J. he changed it into B. The latter name was subsequently frequently used in the literature and taken over in Saccardo’s Sylloge Fungorum (1886). One of these was depicted in Fungi Italici (1879) and is therefore considered as the lectotype.. Samson from insect. CBS 653. CBS 121. May 1935. February 1971. [p.36 from Carpocapsa pomonella.61 (= IMB 415) isolated from soil..36 and CBS 132. CBS 122. ISARIA. Kenneth under No. March 1971. MacLeod (1954) was the first to regard it as synonymous with Beauveria bassiana. France. Although he saw differences in the shape of the conidia between his material and Link’s species.36 isolated from Hylobius abietis..cbs.70 (= IMI 147. CBS 125. he followed Bresadola (in Saccardo. Baarn.36 from Acanthocinus aedilis. respectively. Siemaszko.36 from Leptura sp.36 from Lophyrus pini. minimum as a synonym of S. CBS 126. CBS 129. In 1817 Ditmar described a Botrytis densa Ditm. CBS 650. 2643. The shape of the conidia in each strain is rather constant. but did not actually make the combination. Examination of the type material proved its identity with Beauveria bassiana. 1882) in paying no attention to it.691) isolated from pupa of Thaumatopoea wilkinsonii. CBS 639. sent in 1971 by A. TRITIRACHIUM AND ACR. bassiana to include strains without synnemata. on the basis of which Müller (1965) thought that Spegazzini and Pettit had described different species. under No. The third was collected by Bresadola and had oval conidia. Halperin from Phalera bucephaloides. 9] Both species were validated by Persoon (1822). tenella was described as having globose conidia. CBS 324. CBS 723. This was done by Giard (1891).71 isolated by R. France. On the other hand he retained B. It was published first in a newspaper as Botrytis paradoxa. CBS 654. He thought that Link’s species belonged to Isaria. Müller transferred Pettit’s strain to Trichoderma.36 from unidentified Curculionid. Paris. 2491. Gösswald (1939) mentioned S. possibly an Oidiodendron species was meant. both sent in 1968 by R. Ferron. Both in Michelia (1882) and in Sylloge Fungorum (1886) Saccardo mentioned three original collections. 1954) and man (Kuru.35. May and July 1971.36 from Cossus cossus. The type collection appeared to be the present species.. the type of which (B) is of doubtful identity. The species was described by Link in 1809.36 from Hylurgops palliatus. Helsinki. B. Bilthoven. The capacity to form clustered fertile branches decreases rather soon.71 isolated by W. sent in 1936 by W. CBS 130. Fries (1832) transferred Ditmar’s species to Sporotrichum.71 isolated by R. He gave rather vague illustrations. measuring 1.36 from Ips typographus.71 isolated by J. CBS 124.THE GENERA BEAUVERIA. minutulum Speg.. The second collection in the Saccardo Herbarium appeared to be Beauveria bassiana.

Séanc. delacroixii. Botrytis effusa. r. 5 p.knaw. among which the type. This species was originally distinguished by 1-2 cm high synnemata. bassiana mainly by its angular conidia. natn. Fung. 4: 207. as cited in the Index of Fungi (1971). Paris 113: 270. In the type strain CBS 119. floccose. In some capsules. Univ. fioccoi Dodge .Syll. densa as independent species. . Mycol. B. suppl. Forest Exp. 2. 4. (later homonym). sometimes funiculose. Agric. agron. & Cif. Bot. 18391936 = Sporotrichum beurmannii Matr.C.nl/publications/1001/content_files/content. per Fr.Bull. Botrytis melolonthae Sacc. CBS 109. conidial structures on oatmeal agar. 14: 169. sensu Aschieri . Br. Beauveria bassiana could be found. & Ramond var. Briggs. which may be branched. lunzinensis.Tratt. 1942 = Tritirachium epigaeum (Brunaud) Langer. crittogam. ro: 540. Ser. hot.) Petch . appearing lanose.. Misapplied names Botrytis tenella (Sacc. During his stay in Algeria Brongniart (1891a and b) collected several parasitic fungi from the migratory [p. hebd.Med. later often becoming yellowish to pinkish.Tetrahedon 8. 1912 = Beauveria melolonthae [p. comp... macroconidiana Averna-Saccá (1930). the chlamydospores of Lachnidium acridiorum Giard [= Fusarium solani (Martius) Sacc. Beauveria laxa. b. wine-red pigment produced when the fungus was cultivated on potato.Trans. but from Kuru’s (1931) detailed descriptions and illustrations the synonymy is obvious. 1924. Saccardo (1892) named it Botrytis brongniartii. in our observations the conidia were all globose. was described without a name. 1o: 249.) Cif. Isaria kogane Hasegawa & Koyama . stephanoderis f. who described it as Botrytis acridiorum. According to the description. 1941 Isaria sp. 1891 = Beauveria tenella (Sacc. Pavia. illegitimately published as Botrytis acridiorum Brongniart & Delacr. Paris 2: 164.Précis Parasitol.) Petch . 1935 (substitute name) = Beauveria epigaea (Brunaud) Langer. he only described B. Beauveria brongniartii. conidia. 1954. . – Annls Parasit. 4 ? Sporotrichum epigaeum Brunaud . 1947. of which no type material is left. 4. Ser. van Beyma in an unpublished study of the type strain. globulifera and B. According to Petch’s description the species should differ from B.Annls Soc. Botrytis brongniartii subsp. Botrytis bassiana var. Botrytis brongniartii Sacc. MacLeod (1954) actually made the combination Beauveria tenella. conidial structures on 2°/o malt agar. c.htm sent part of his material to Delacroix for comparison with L. a. bassiana. The value of these characters was already doubted by Schwarz (1924). It was sent to Trabut.) Delacr. ISARIA. 22: 98. which strongly resembled the ‘red muscardine’ found in France..) MacLeod . H. 10] locust. 11] (Sacc. The first of these was a Fusarium species (Delacroix. Soc. A species with ellipsoidal conidia.) Brown & G. prat. bassiana were the floccose to funiculose mycelium. fioccoi (Dodge) Red.cbs. le Moult’s ‘red muscardine’. Govt. 1891. Botrytis stephanoderis. would differ from the typical variety by its yellow and reddish pigments. . Beauveria brongniartii (Sacc. which were formed in crowded masses on the rachis. 1: 269. Microscopically it did not differ from B. Lab. B..26 both characters had almost disappeared. and by a yellow reverse. var. TRITIRACHIUM AND ACR. The combination with Beauveria was mentioned by Siemaszko (1937). Sci. He also redescribed the species with globose conidia. Isaria shiotae.. umana 5: 481. However. Micopat.Fig. This conclusion was also drawn by F. That is why the description by Delacroix (1891) of his Botrytis tenella with ellipsoidal conidia differed considerably from Saccardo’s. however. Colonies in vitro attaining a diameter of 10-16 mm in 8 days. and the CBS culture of the type was lost. delacroixii. 10: 320. Smith. . and a stable. . This variety.Annls mycol. bassiana. at first white. 808.24. The type strain CBS 337. Sci.32 is in rather bad condition. p. 1966. Rochelle 1888.Atti Ist. 1929. up to 5 mm high.THE GENERA BEAUVERIA. velvety to powdery.Can. Collections of this species in K contained mostly Paecilomyces farinosus (Holm. Under the name B.J. Fig.) Giard .] were erroneously described. No material is preserved in PC. 1892 = Beauveria brongniartii (Sacc. The second was a Beauveria species with globose conidia. which he validated by naming it Botrytis brongniartii subsp. important criteria for separation from B.Publnes Estac. Isaria densa (Link per Pers. in Brumpt . Acad. Fergus & Shannon . 32: 823. 1929 = Sporotrichum schenckii Matt. 1893). 74. ed. Moca. hum. 6 sur 22 http://www. mycol. J. rarely 04/03/2015 16:32 . Stn Meguro 4.

and the conidia were obviously ellipsoidal. which is often more clustered than in pure culture. subglobose cells or 1-3 (-4) conidiogenous cells in the first.Mycologia 32: 27. but did not change the name.cbs. Kew.nl/publications/1001/content_files/content. No type material was available in PC. more often the conidiogenous cells are arranged in small groups or solitarily along the hyphae. and CBS 327. From Brunaud’s description it is. rarely funiculose. who mentioned the geniculate rachids of the conidiogenous [p. Beauveria alba (Limber) Saccas . Pollacci.25 isolated from Mantis sp. and a well developed rachis. rarely red or purple.34 (= ATCC 7145) isolated from man.24 isolated by R. though Aschieri (1929) had not excluded the type. Material examined Herbarium specimen Botrytis melolonthae in herb. Our observations on the strain do not exactly agree with Aschieri’s.5-) 2 (-3) µm.42 isolated from Melolontha melolontha.5-) 2-2.26 from unidentified Salticide.Revue Mycol. CBS 109. on Melolontha melolontha. le Moult. Kiel. 1-4 µm wide.67 from Melolontha melolontha.5 (-6) x (1.5 (-4) µm. sent in 1942 by H. CBS 223.htm reddish to purple. 1940 = Beauveria alba (Limber) Saccas . Aschieri recognized the relationship with Beauveria. 5 Tritirachium album Limber . 04/03/2015 16:32 . Ciferri from insect. see discussion of Beauveria bassiana.5-3 µm wide.Fig. denticles mostly as wide as the rachis. Germany. CBS 112. PAD (type)... delacroixii. 13: 64.5 (-3) µm.THE GENERA BEAUVERIA. sent in 1924 by L.5 µm long. densa.. Submerged hyphae hyaline. bassiana. No chlamydospores were found.5-) 2-3. Turkey. Göbelez.5 µm wide.knaw. however.53 (= IFO 5299). which by further branching may give rise to smaller. type culture of Isaria kogane. Aschieri (1929). CBS 326.5-4. W.67 isolated from Costelytia zeelandica. CBS 722. strain producing ‘isarolide’ described by Briggs & al. appeared to be B. Exudate and odour absent. up to 1. The following names are considered here to be synonyms of B. (2-) 2.. Conidiogenous [p. Ferron. ISARIA. creeping or ascendent. No chlamydospores were observed. denticulate. CBS 111. sent in 1934 by G. 1.22 (= ATCC 9452) isolated from mummified larva. likely that he meant a Beauveria species. Conidia hyaline. Botrytis brongniartii. however. smoothwalled. Hyphae of the aerial mycelium bearing a conidial apparatus.. CBS 410. Langeron (1947) transferred almost all Beauveria species to this genus. (3-) 4-15 (-28) x (1. or sometimes in the second order. sometimes subcylindrical basal part.53 isolated from larva of Balaninus sp. The combination with Tritirachium was made by Langeron (1947) following Limber (1940). Pavia. Dodge (1935) made a new name and mentioned in the synonymy ‘Sporotrichum epigaeum Aschieri not Brunaud’. sent in 1924 by L. Living strains CBS 106. and CBS 120. Perfect state unknown. 12] cells consisting of a subglobose or flask-shaped. both sent by P. 3. Indonesia. TRITIRACHIUM AND ACR. Blunck. 1964). 13 and 6 respectively. Discussion Recently the species has mostly been referred to as Beauveria tenella or B. Italy. Roberts as Isaria sp. Mycelium on natural substrate lanose to somewhat pulverulent. geniculate or irregularly bent. widest near or below the middle. Montemartini. Reverse uncoloured or yellowish to orange. under B. smooth. 1948. up to 9 x 5 µm. bearing small groups of orthotropic or slightly plagiotropic lateral cells. This misunderstanding was taken over by MacLeod (1954). up to 25 µm long and mostly 1-1. yellowish. smoothwalled. ellipsoidal (rarely subglobose). sent in 1953 by M. brongniartii: Sporotrichum epigaeum. (2-) 2. Hyphae of the aerial mycelium hyaline. On this strain all later combinations are based. described by E. 13] cells as a genus character. The oldest strain available is the one described by Aschieri (1929). (1966). brongniartii var. 7 sur 22 http://www. sometimes with a pointed or apiculate base. both sent by National Collection of Type Cultures. Java. 1898.5-3.71 (= PDD 6363) sent in 1971 by D.5 (-4) x (1. Occasionally the conidial apparatus is clustered. the type specimens of both. sometimes reddish. under No. CBS 128. No clusters of conidiogenous cells were reported. while Aschieri (1929) called them ‘rundlich oder leicht länglich’ (Müller.

Reasons for classification in Beauveria are the denticulate rachis and the absence of pigment in the aerial mycelium. denticulate. 15] denticulate conidiiferous portion. Conidia hyaline. bearing conidiogenous cells in whorls of 1-3 at wide. Exudate and odour absent.5-2. up to 70 mm long. 1832. Isaria Fr. Colonies in vitro attaining a diameter of 14-16 mm in 8 days. Isaria Fr. No chlamydospores were observed. A. globose to subglobose.5 (-3) µm.5-2. appearing velvety.Syst. under No. on cover of a book. . they consist of a swollen basal part. somewhat stiff. 5. CBS 570. 8 sur 22 http://www.5 µm. Colonies growing moderately slowly. Perfect state unknown. Conant. March 1939. 3:270. brongniartii only by the shape of their conidia. sometimes with an apiculate base. 2-4 µm wide. Living strains CBS 348. Fertile hyphae mostly slightly differentiated. ascendent to procumbent. de Jongh. sent in 1971 by R.5-) 2-3 (-3. loose or fasciculate. (1. and a well developed rachis. Conidiogenous cells scattered. O. conidial structures. occasionally in dense clusters. BPI. dried culture isolated from human sputum.htm Fig. Discussion The zig-zag shape of the rachis and the verticillate ramification make the species reminiscent of Tritirachium. b. Duke. ISARIA. soon forming synnemata.5 µm. purely white. W. thin-walled. (7-) 10-24 (-30) x 1. Conidiogenous cells consisting of an elongate to subcylindrical tapering basal part. N.. 6-12 x 2-3 µm. smooth. a. smooth-walled.knaw.5 µm wide. irregular. TRITIRACHIUM AND ACR. 1-2.) Fr.71 sent in 1971 by R. 2343. USA. No chlamydospores or perfect states were observed.71 isolated from Pisum sp. mycol. not elongating. lanose to arachnoid. Lectotype species: Isaria felina (DC. Aerial hyphae hyaline. apically dichotomously branched.5) x (1. Tritirachium album in herb. T. Descriptions are based on colonies growing on oatmeal agar at 20° C. Beauveria alba.71. c. or rarely swollen up to 5 µm. per Fr. Reverse ochraceous buff or uncoloured. purely hyaline. up to 3 5 µm long and rather constantly 1 µm wide. de Vries from skin of a woman with eczema vesiculosum. USA. smooth. CBS 836. F. 8296-1970. Ramification is not always strictly verticillate: poorly developed subcultures sometimes can be distinguished from B. smooth. conidia.. Curaçao. Richards. white or yellowish. sometimes zonate. [p. hyaline. denticles as wide as the rachis and up to 1 µm long.55 isolated by G. CBS S70.cbs. Kenneth under No. sometimes slightly swollen up to 3.. Material examined Herbarium specimen Tritirachium album in herb. up to 2 mm high.THE GENERA BEAUVERIA. geniculate. BPI (type). conidiogenous cells occasionally scattered or with two on a lateral stalk [p.5-) 1.and thin-walled. Conidia one-celled. leg. and a short and narrow. 14] cell which is cylindrical. Mycelium on natural substrate thin. Woods Hole. arising directly from undifferentiated hyphae or from one-celled. subglobose to subcylindrical or kidney-shaped.nl/publications/1001/content_files/content. often right angles. Netherlands. Submerged hyphae and hyphae of the aerial mycelium hyaline. Massachusetts. appearing lanose to floccose. Discussion 04/03/2015 16:32 . sometimes swollen side branches.

Submerged hyphae hyaline. pirina El. bearing orthotropic conidiogenous cells. orthopterorum 4. eur. Hyphae of the aerial mycelium hyaline. often curved.) Fr. flask shaped or curved.5 x 2-2. conidia. 1882. Isaria felina (DC. Key to the species 1a. crassa Pers. per Fr. Isaria fimicola Sternon . the younger genus Paecilomyces would become a synonym of Isaria. mostly 3-8.. Abt. 1963). 1923. smooth-walled. Colonies in vitro attaining a diameter of 8-14 mm in 8 days. ellipsoidal or ovoidal. He listed I. 1: 496.nl/publications/1001/content_files/content. 2. per Fr. was proposed as lectotype of Isaria by Clements and Shear (1931). Aires 23: 121. . Solomon Isl. usually unbranched.) Biourge . Papua-New Guinea p. 1821 = Fibrillaria felina (DC. up to 70 mm high and constantly 1 mm wide. eleutheratorum Nees. per Fr. probably belonging to an immature basidiomycete. mostly comprising 2-4 denticles. 16] Fr. without mentioning any species. suina Sacc. was the first species mentioned by Fries (1832). Isaria felina (DC. a. per Fr. 6. In this sense the genus at present contains two species.Bull. Bakt. var.e. 1882.. aviaria Sacc. per [p.) Fr. smooth. velutipes Link. from which several white.Syst. He mentioned three species.Fig. Isaria felina (DC. conidiogenous cells. solitarily or in small groups. sometimes with a pointed base. terete. Isaria felina (DC. I.5) x (2-) 2.5-3 (-3. Fasc.) Fr. Belg. Exudate rarely produced. from which several white. 1832 = Penicillium felinum (DC. which probably were Calocera viscosa Fr. in age a short geniculate rachis may be formed.Michelia 2: 561. von Arx (1970) chose Isaria felina as lectotype of Isaria. per Fr. (2. 1971. I. 1815.) Fr. in old strains appearing lanose to floccose. tomentose.THE GENERA BEAUVERIA. while both original species of Persoon were listed. b. Soc. Conidia hyaline. up to 80 mm high and constantly 0. 04/03/2015 16:32 . Hyphales p. B. 152. cuniculina Ferr. irregular conidiiferous portion.Syst. 1-4. var. In 1821 Fries cited the name Isaria Pers.5 µm. 6. though none of Persoon’s species are explicitly mentioned. . Isaria farinosa Dicks. forming in fresh isolates a dense felt. It was chosen as the lectotype of Isaria by Petch (1934). 1822 = Isaria felina (DC. 1921. var. at first white. ISARIA. per Fr. bot.5-1 mm wide. mycol. however. 1923. mycol. erect synnemata arise. which are both identical with Paecilomyces farinosus (Dicks. Belg. ParasitKde. Isaria cretacea van Beyma . subglobose. Reverse uncoloured or yellowish to yellow brown. Isaria felina (DC.. conidia broadly ellipsoidal Conidiogenous cells globose. Fig.Mycol.An. per Fr. 1935 = Beauveria cretacea (van Beyma) Matsushima . conidia subcylindrical to kidney-shaped I. ascendent or fasciculate. TRITIRACHIUM AND ACR. 17] cell. Isaria felina. odour absent. var. nat. per Fr. and I. 1b Conidiogenous cells flask-shaped. however. UPS) showed. Mus. Marchai & Et. 1-3 µm wide. bot. . 1910.) Fr. Isaria edwalliana P.Bull.) Fr. . mucida (= Ceratiomyxa fruticulosa) and I. 91: 350. Isaria felina (DC. 1904.Fl.) Fr. per Fr. .. Conidiogenous cells not elongating.Cellule 33: 102.5-) 3-4 (-4.) Petch]. 1934). Smith according to the type material (L). None of Hill’s species.. sometimes 1-2 conidiogenous cells are supported by a slightly swollen stalk [p. Ceratiomyxa fruticulosa (Müller) MacBride and a Puccinia species respectively (Petch. Isaria terrestris Fr. conidial structures.Fl. smooth-walled. 1:53.Microf. Persoon (1794) introduced the name Isaria again for the description of two species. per Fr. creeping. 9 sur 22 http://www. Examination of the type material of this species (herb. agaricina [= Tilachlidium brachiatum (Batsch per Fr. ital. crypt. Also Gray (1821) referred to Persoon.Hedwigia 43: 209. In order to provide a generic name for synnematous Beauveria-like fungi. fr. .htm The name Isaria was first used by Hill (1751). felina I. . domestica Speg. 3: 271. usually unbranched. . R. consisting of a swollen.. 7. var. and a short. Soc.cbs. or occasionally branched at the apex. later becoming yellowish. If this were accepted. 55: 145. nac. . in the introduction of his Systema Mycologicum. Hist. S4: 134. positively phototropic synnemata arise (Taber & Vining. Henn. In 1832 he did not refer to Persoon (1794) but to Hill (1751).5 µm wide. Mycelium on natural substrate forming a thin layer of hyaline hyphae.5) µm. that the fungus consisted of sterile hyphae. occasionally elongate basal part. 6:30.) Brown & G. was mentioned. 6 Clavaria (?) felina DC.Michelia 2: 561.Zentbl. .) Pers.knaw. 1912. i. R. per Fr. c. Marchai . No chlamydospores were observed.

on rabbit dung. Sao Paulo. sent in 1936 by R. CBS 250. It was collected by T. F. Living strains CBS 110. type culture of Isaria cretacea.36 isolated by H. connected with a constricted base to the hyphae. leg. sent in 1936 by H. Diddens from pupa of Anaitis efformata. sometimes a short geniculate rachis is present. ISARIA. sent in 1936 by D. 7. N. 2. terete or laterally compressed. Petch from several locations in Ceylon and once in Thailand. mostly comprising 3-6 denticles. type. Fibrillaria felina in herb. in herb. Hyphae of the aerial mycelium hyaline to yellowish. 1920.262.50 isolated from rabbit dung. Greis. Isaria felina in herb.5 mm wide. Mycelium on natural substrate forming a thin layer of hyaline hyphae.htm Perfect state unknown. Isaria edwalliana in herb. 1534. Fries (UPS). CBS 235. 18] branched. giving rise to (1-) 3-6 (-7) conidiogenous cells in the first. Bommer & Rousseau (B). type. later glabrous and horny. b. CBS 648. Great Britain. Chevallier. a. ex herbarium Durieux de Maisonneuve.Trans.knaw. sent in 1950 by Lab. Canada. Some [p. Conidia hyaline. on carnivore dung. No chlamydospores were observed. Kotmake. Ontario. and said to occur on grasshoppers and probably also on Lepidoptera. Discussion This species is only known as dried herbarium material. narrower at the base than at the broadly rounded tip. CBS 312. conidia.. Material examined Herbarium specimens Isaria felina in herb. suina Sacc. bearing groups of subglobose to ellipsoidal lateral cells. TRTC 45685. Isaria orthopterorum Petch . Blakeslee. on Orthoptera indet. outer surface irregular. up to 8 µm long and 3 µm wide. smooth. Tolman. (2. Microbiology. sent in 1950 by K. Auerswald. Malloch under No. CBS 229. flattened on one side to kidney-shaped. leg. Brazil. Petch (K). rarely somewhat apiculate. conidial structures.71 isolated from porcupine dung. and a short conidiiferous portion. B. PC under No. F. Shear (BPI). rarely in the second order.36 isolated from rabbit dung.. Sydow (S). 1933. Isaria fimicola in herb. Fig. Germany. up to 8 mm high. 2. Br. B. 10 sur 22 http://www. 910. Buenos Aires. Isaria orthopterorum. when young tomentose. CBS 173.Fig.66 sent in 1966 by F. Belgium.5 µm wide.101 and 910.cbs. BR 17/50. Material examined Isaria orthopterorum in herb. Isaria felina ex herb. from which several yellowish synnemata arise. c. with a constricted base.5-3 µm wide.5-3.5-) 3. 1882. 1903. Germany. mostly 2. were described briefly and illustrated (1924a) as Isaria sp.5-4. Chevallier.THE GENERA BEAUVERIA. ellipsoidal to subcylindrical.5 µm in diameter. CBS 217. Jardine. The conidial apparatus is rather tightly clustered. Ceylon.262. 04/03/2015 16:32 . L. mycol. leg.. Italy. 1924. mostly [p. from type collection. 18: 70. on rat dung. 5. hyaline. CBS 236. K. Argentina. red brown. P. Verona. Marchal. A. Leipzig. sent in 1969 by D. J. Soc. consisting of a globose to ubglobose.58 sent in 1958 by O. Boedijn under No. Hutchinson. including the type. probably collected by F. Soest. TRITIRACHIUM AND ACR.50 isolated as culture contaminant. Motelay. Perfect state unknown. BPI (type). Isaria fimicola in herb. Epsom.08 sent in 1908 by A.nl/publications/1001/content_files/content. 3-4. CBS 313. Erlangen. smooth-walled. S. Nos.37 isolated from mouldy leaves. 1879. stalk up to 0.5) x 2-3 µm. Isaria felina in herb. Herrero. Bresadola (BPI). Conidiogenous cells not elongating. rarely pear-shaped basal part. leg. Persoon (L). Hellinga. 19] Ceylonese specimens. Torino. smoothwalled.5 (-5. Isaria felina var. 7 Isaria (Beauveria) orthopterorum Petch .5-3. apically often swollen up to 2 mm.34.5 x 3-4 µm or pointed if terminal. Delft. leg. sent in 1934 by D. Gembloux. cultures on carrot. on dung. Fertile hyphae mostly arising from the heads of the synnemata only.106.

slightly swollen below the middle. Conidiophores bearing several whorls of side branches. smooth. No chlamydospores were observed. smooth-walled. tapering towards the tip. TRITIRACHIUM AND ACR. 3-4. hyaline. 1939 (without Latin diagnosis). ISARIA. ascendent to suberect. ochreous (17’ to 19"i) (Colour names with symbols refer to Rayner (1970)). No chlamydospores or perfect states were observed.THE GENERA BEAUVERIA. and a well developed. not be established. bearing conidiogenous cells in 1-2 whorls of 2-4.5 µm wide. stiff.5-3. variously coloured. 04/03/2015 16:32 .htm though the latter was not mentioned in the subsequent description in 1933.nl/publications/1001/content_files/content. lighter towards the base. 20] a whorl of conidiogenous cells. Conidia one-celled. 13: 64. Conidiogenous cells consisting of an elongate basal part. and the conidia mostly have a truncate or apiculate base.Antonie van Leeuwenhoek 6: 286.J.. golden brown in the upper part. and a well developed. Colonies growing rather slowly. Spirotrichum Saito . The conidia were said to be formed in chains. so the species is not validly published. the conidiogenous cells are cylindrical when a geniculate rachis is present. 1940 = Beauveria dependens (Limber) Saccas . up to 150 µm wide and 220 µm high. 1940. Mycelium on natural substrate appearing velvety. branches 2-3 µm wide.5 µm pale golden brown. Tritirachium Limber Tritirachium Limber . up to 40 µm long and 1-1. smooth. (2-) 2. geniculate (intervals 2 µm cicatrized rachis. slightly narrowed at the base. however. Living cultures were not available.5 µm wide. Technol.knaw. bearing 4-6 (-10) whorls of 3-4 (-5) side branches or conidiogenous cells. thin-walled.Mycologia 32: 26. Feb. however. ascendent. 11 sur 22 http://www.. each supporting [p. Tritirachium dependens Limber . Key to the species 1a. tapering towards the tip. oryzae 6. often somewhat tapering towards the tip. subglobose to ellipsoidal. Sawada did not give a Latin diagnosis. Conidiogenous cells consisting of an elongate basal part. 8 a-d Tritirachium dependens Limber . colonies purplish T. The denticles may be very small or absent in Geniculosporium and Nodulisporium. In these genera. regularly geniculate. ex van Beyma . 1b. Acrodontium and Hansfordia the rachis bears distinct denticles. colonies ochraceous Conidiophores rarely branched. dependens T. Descriptions are based on colonies growing on 2% malt agar at 20° C. conidial apparatus pyramidal in outline. Ferment. Type species: Tritirachium dependens Limber Discussion The genus was separated from Verticillium by Limber (1940) on account of the sympodial conidiogenous cells.5) µm. Conidia hyaline to slightly yellowish. up to 1 mm long.Revue Mycol. globose to ellipsoidal. thin-walled. In order to provide a better delimitation of the genera Tritirachium and Beauveria. 1948. Conidiophores brownish.. regular.Fig. mostly 13-35 x 2-3. smoothand somewhat thick-walled. 1940. in the latter two genera the rachis is not or hardly geniculate. In Beauveria. Perfect state unknown. appearing velvety to powdery. imperceptably merging into the vegetative mycelium. but in 1941 Petch corrected his error. imperceptably merging into the vegetative mycelium. slightly swollen at the base and tapering towards the tip.5 (-4) x (2-) 2-3 (-3. bearing in the upper part several whorls of conidiogenous cells or verticillate side branches. cicatrized rachis. branches of second order sometimes present.cbs.Mycologia 32: 26. mostly supporting whorls of conidiogenous cells only. The description of Beauveria chiromensis Sawada (1959) is reminiscent of B. Sept. Conidiophores hyaline at the base. Osaka 27: 1. more emphasis is laid on the mode of the proliferation of the conidiogenous cells. without examination of the type material the identity can. orthopterorum.

. Frink & A. Spirotrichum purpureum Saito . 1942. P. D. ISARIA. Frink & A.Fl. 1815]. 1947. 22] 7. Watson. Recife 447: 16. China. & Lichaa . 12 sur 22 http://www.Bull. Tritirachium oryzae. Tritirachium roseum van Beyma . May 1935. eur. 1942. J. 1950. Limber. comb. Germany. Fr. . vég. on bulbs of Lilium eruba.. a-d Tritirachium dependens. appearing velvety to lanose. 1939 (without Latin diagnosis) = Tritirachium purpureum (Saito) van Beyma . on root of Kudzu (Legume). on bulbs of Lilium sp. 37-1845 (San Francisco 27437). agric. R.5-3 µm wide.e-h. when older lilac (69"‘) or cinnamon (13"b). on bulbs of Lilium giganteum. China.. on bulbs of Lycornis radiata.J. R. Japan.Mycol. L. conidiophore. P. on Watsonia rosea.knaw. 841. 1950. b. smoothwalled.5-3 µm. Fenner & D.and somewhat thick-walled. up to 165 µm. 1822 [non Botrytis rosea DC. ascendent. China. a. f.Antonie van Leeuwenhoek 8: 1I8. 04/03/2015 16:32 . 1940 = Tritirachium musae (van Beyma) van Beyma . slightly tapering towards the tip.nl/publications/1001/content_files/content. No chlamydospores were observed. P. conidial structures. 1950. P. imperceptably merging into the vegetative mycelium. Living strains CBS 388. TRITIRACHIUM AND ACR. leg. thin-walled. leg. Japan. det. South-Africa. Perfect state unknown. on Pueraria thunbergiana. Freunde. (1.THE GENERA BEAUVERIA. Limber.Revue Path. 1950.Annls Parasit. Saito. P.. geniculate (intervals 1-2 µm cicatrized rachis. [p. often somewhat stromatic. J. Tritirachium oryzae (Vincens) de Hoog. Watson. Japan. 37-2601 (San Francisco 27433). San Francisco 29077.Antonie van Leeuwenhoek 6: 286. tapering towards the tip. Watson. Spirotrichum musae van Beyma . J. Japan. per Pers . & Lichaa) Langer. (7-) 9-20 (-25) x 1. Limber. 1934 = Tritirachium brumptii (Langer. M. smooth. on twigs of Salix sp. Sydow (B) under No. from type collection. Watson.htm Material examined Herbarium specimens Tritirachium dependens in herb. 22: 94. hum.5) x 1. d. China. . 37-2059 (San Francisco 27474). 1. conidiophore. at first rosy vinaceous (7"f) to rosy buff (11"d). Watson. 37-3614 (San Francisco 27357). Stellenbosch.. e. J. J.Antonie van Leeuwenhoek 8: 117. Paris 111: 133. and I µm wide. 1950. z: 71. 1816. conidia. nov. Frink & A. 1965. h. and det. P. Material examined Herbarium specimen Botrytis rosea in herb. Conidiogenous cells consisting of an elongate basal part.. A. R. globose to ellipsoidal. Misapplied name Tritirachium dependens Limber sensu Batista & al. Watson. c. Ges. Micol. fr. 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. A. mostly 25 µm long. [p. . 21] Fig. 1910 (basionym). comp. det. Frink & A. Further collection in BPI: New York 762188. or brown vinaceous (5"‘m). D.Antonie van Leeuwenhoek 8: 117. P.Publçoes Inst.39. Ent. .5-3 µm and a well developed. Colonies in vitro growing slowly. Technol.Fig. det. conidial structures. lighter towards the base. J. 1951. Submerged hyphae hyaline to pale brownish. Beauveria oryzae Vincens . type culture of Spirotrichum purpureum. det. 37-2811 (Hoboken 15494). sent in 1939 by K. det. rarely with a slightly apiculate base. on dead root of Yucca tremuleana. 1941.Mag. Ferment. 1-2 µm wide. det. 1952. R. conidia.. 200-1500 µm long. Conidiophores reddish brown. 10: 122.5-) 2-3 (-3. on bulbs of Lilium sp. 8. Méd. Osaka 27: 1. July 1909. smooth. Brandenburg. 1942. Exudate and odour absent. 1: 35. det. g. attaining a diameter of 3-5 mm in 8 days. 37-1609 (San Francisco 27285). slightly swollen below the middle.. naturf. Reverse dark brick (9"m) and towards the margin lighter. conidiogenous cells. Acad. 8 e-h ? Botrytis rosea Link . Conidia hyaline. conidiogenous cells.cbs. Beauveria brumptii Langer. Berlin 7: 36. BPI (type).

make its identity with Tritirachium oryzae very likely. isolated from leaf spots of Musa sapientum. 1960. basi apiculata. which is cylindrical. Ghana. No chlamydospores were observed. and a long and slender rachis. Especially the ‘Forme ß’ did not differ in any respect from CBS 896. CBS 896. by the 04/03/2015 16:32 . van den Ende. Conidiophores. from which it differs mainly by the differentiation of the conidiogenous cells into a basal part. Many genera with one-celled conidia borne on sympodial conidiogenous cells are known. 1361 as Beauveria sp. 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. sent in 1970 by J. plerumque verticillatae e cellulis fertilibus oriuntur. hardly apiculate when fusiform. Martin as culture contaminant. No type material is available of Beauveria brumptii. sent in 1971 by R. sent in 1970 by G. Abidjan. which appeared to be Tritirachium oryzae. Colonies growing rather slowly. of equal width as the basal part of the conidiogenous cell. levia.40. varie coloratae. which is lost.a. Conidiophora. Dematophora Hartig and Tharoopama Subram. when present. Phaeoisaria Höhnel. ramosae. Acrodontium de Hoog. brownish or olivaceous. nov. [p. PC 2042. type culture of Spirotrichum musae. wide or irregular rachis. orthotropica vel plagiotropica e hyphis repentibus oriuntur. nonnumquam crassi-tunicata et deorsum fusca. Chlamydosporae absunt.nl/publications/1001/content_files/content. because of their variability when the fungus is grown on different media.70 isolated from packing cartoon. smooth. they consist of a flask-shaped or elongate basal part.: Gr. often also in plagiotropic whorls on more or less differentiated conidiophores. verticillata vel dichotoma. and mostly considerably larger than in Acrodontium. Nicot under No. Conidia one-celled.htm CBS 175. The following genera are most similar to Acrodontium: Dicyma Boul. Germany. hyalina vel pigmentata. sent in 1967 by A.cbs. 24] from undifferentiated hyphae. Vincens (1923). Kiel. 1942) are mainly based on cultural characters. and a sympodial denticulate rachis.70. Recife. are of little use as taxonomic criteria. Batista as Tritirachium dependens. non Kamyschko. tapering towards the tip. sometimes thickwalled and dark brown at the base. Conidiogenous cells solitary. hyaline or pigmented. subglobosa vel fusiformia. Van Beyma’s species (1940. who isolated several strains of the species in Indo-China as contaminants. Descriptions are based on colonies growing on 2°/o malt agar at 20° C. The type of Beauveria oryzae seems to be lost. odón = tooth. Hyphae aeriae subhyalinae. brunneolae vel olivaceae. or plagiotropically when little differentiated. The description by Langeron (1934) and some unpublished drawings by van Beyma of a CBS subculture of the type. Kenneth under No. Sect. bearing at regular intervals lateral. sent in 1934 by G. lighter towards the apex. recta vel flexuosa constant. si adsunt. branched verticillately or dichotomously. gave good descriptions and illustrations. of variable length. the most typical strain on which the description is based. velutinae vel pulverulentae. Sect. TRITIRACHIUM AND ACR. Acrodontium is somewhat similar to Rhinocladiella Nannf. The conidia are hyaline. singulae vel fasciculatae.and rather thin-walled. Nijmegen. septate. smooth. loose or fasciculate. tapering towards the tip. gen. denticulata. 13 sur 22 http://www. leves. orthotropicae e hyphis. C. In Hansfordia the rachis is mostly very short. akrós = apical. CBS 442. sent in 1942 under No. ISARIA. vel plagiotropicae. Species typica: Chloridium crateriforme van Beyma Etym. Cellulae conidiogenae solitares. fere tenui-tunicatae. e parte basilari lageniformi vel elongata sursum attenuata et rachide sympodialiter elongata.70 isolated as culture contaminant. arise orthotropically. Grisea is similar to Hansfordia Hughes. from creeping hyphae. Discussion The genus Acrodontium is separated from Tritirachium on account of the morphology of the conidiiferous rachis: it is straight or only slightly flexuous. CBS 837... subglobose to fusiform. variously coloured. Discussion No type material of Botrytis rosea was available in B. Brazil.71 isolated by K. arising orthotropically [p. appearing velvety to powdery. Rehovot. with an apiculate base. type culture of Tritirachium roseum.67 isolated as culture contaminant. straight or somewhat flexuous. alternating denticles. Acrodontium can be subdivided into two sections. only a secondary collection from the Sydow herbarium (B). could be examined. Stahel as Verticillium sp.. Israël.42.knaw. Surinam. Coloniae fere lente crescunt. sursum clariora. differ i. 2330. erect or procumbent. erecta vel procumbentia. Conidia simplicia. CBS 183. which. branched. septatae. Aerial hyphae subhyaline.THE GENERA BEAUVERIA. 23] CBS 164.

Hattori but.) in PAD. in the latter case dark olive Conidiophores rather thin-walled over the whole length. sensu Tubaki . simplex 6 A. guttuliform to lageniform. hyaline. crateriforme. cannot be used for this group of fungi. 6a. 1958. (?) globulifera Arnaud and R. 3b. branched over the whole length Conidiophores differentiated. 2b. salmoneum 3 A. The names Chloridium Link and Psilobotrys Sacc. tapering towards the tip Conidiogenous cells swollen. conidiogenous cells pigmented. 4a. tapering towards the tip.cbs. very dark brown. Bakt.. sharp denticles. in A. 3a. Conidia guttuliform to fusiform. 5a.Fig. whereas in A. Conidia hyaline. somewhat flexuous Conidiophores in the lower part thick-walled. and Trichosporium Fr. 4b. 89: 241. Abt. Criteria used for the distinction of both sections of Acrodontium are the presence of more or less differentiated conidiophores. 26] Acrodontium crateriforme (van Beyma) de Hoog. Lab. Both other species of the genus. pigmented. (?) pilulifera Arnaud. with an apiculate base. 2. Rhinotrichella Arnaud (not validly published) seems to be closely related. consisting of an elongate or subcylindrical basal part. conidia often finely verrucose 2 4 A.htm presence of distinct synnemata. which was used for several species with sympodial conidiogenous cells by Arnaud (1953) is a nomen illegitimum. 9 Chloridium crateriforme van Beyma . ParasitKde. comb. 1933 (basionym). Haplaria Link is a synonym of Botrytis Pers. are possibly more closely related to Rhinotrichum Corda. mostly flask-shaped or somewhat elongate. 14 sur 22 http://www. and bears a series of small. which is often not distinctly separated from the basal part. According to the same author. the shape of the conidiiferous rachis and the pigmentation of the conidia. 1968). conidia smooth Mature colonies gray brown. and a somewhat flexuous rachis. . widest somewhat below the middle. 5b.knaw. virellum A. R. was too scanty to trace the mode of conidial production. ISARIA.nl/publications/1001/content_files/content. less than half as wide as the rachis. per Fr. nov. of about the same colour as the hyphae Colonies dark olive or gray brown. i. 1b..e. however. branched verticillately in the upper part only Conidiogenous cells elongate. with an apiculate base.Zentbl. hydnicola the conidiophores are often integrated in the vegetative mycelium.THE GENERA BEAUVERIA. 20: 152. 6b. griseum Acrodontium section Acrodontium Conidiogenous cells arising orthotropically from undifferentiated hyphae. Geniculosporium Chesters & Greenhalgh and Nodulisporium Preuss have more or less cylindrical conidiogenous cells with cicatrized rachids.J. 04/03/2015 16:32 . grisea (Sacc.) Arnaud (= Rhinotrichum griseum Sacc. 2a. The original material of the type species R. However. Misapplied name Chloridium minutum (Sacc. Key to the species 1a.. [p. TRITIRACHIUM AND ACR. For this reason it is thought to be preferable to maintain all species within the same genus. rachis with blunt denticles and always distinctly differentiated from the basal part of the conidiogenous cell (section Grisea) Colonies pink..) Sacc. The appearance [p 25] of the type species of the sections may differ considerably. sharp denticles and often not differentiated from the basal part of the conidiogenous cell (section Acrodontium) Conidia globose to guttuliform. rachis with small. light brown. widest near the base. darker than the hyphae Mature colonies dark olive. verticillately branched fertile hyphae are sometimes present. crateriforme Acrodontium-state of Ascocorticium anomalum Conidiophores integrated in the vegetative mycelium. conidiogenous cells subhyaline. and Rhinocladiella has almost imperceptable denticles and pleomorphic conidium formation (Schol-Schwarz. hydnicola 5 A. Hughes (1958) proved their type species to be congeneric with Bisporomyces van Beyma and to produce phialoconidia. stiff A.

c. arising orthotropically from undifferentiated hyphae or in groups of 1-2 from the tip of a subtending cell.71 isolated by W.5-5 (-6) x (2. Tjibodas Hortus Botanicus. sent in 1933 by H. with a scar at the base. CBS 841. gray olivaceous (21""). which is often not differentiated from the basal part. conidial structures. forming a dense stroma.5 µm.70 isolated from living leaflet of Fraxinus excelsior.71 isolated by W.71 isolated by W. CBS 844.33 (= ATCC 15679). Conidiogenous cells scattered.. reported by Oberwinkler and al.71 isolated by F. Westmoreland.5-) 2-3 (-4) µm. Acrodontium-state of Ascocorticium anomalum (Ellis & Harkn. April 1968. sent in 1971 by E.71 isolated by W. ETH-M 7090. tapering towards the tip. Casagrande from bark of Pinus sylvestris.71 isolated from foodstuffs.5 (-4N) µm. Material examined CBS 144. April 1968. Gams from Citrus leaves. b.knaw. sent in 1958 by J. 28] rather irregular. Diddens. Germany. denticulate rachis. often strongly fasciculate. appearing velvety. sharp denticles at intervals of 0. Java.cbs. often forming radially furrowed and zonate. TRITIRACHIUM AND ACR. Reverse olivaceous (17"‘m). straight or flexuous. often with a lateral swelling near the base. more or less globose chlamydospore-like structures are present. smooth or finely verrucose. Oberwinkler. tapering towards the tip.nl/publications/1001/content_files/content.htm Fig. with small. Hyphae of the aerial mycelium hyaline to pale olive. .Nova Hedwigia 14: 287. (3-) 3. straight or flexuous up to 45 µm long and constantly 1 µm wide. CBS 840. vinaceous buff (17"‘d). C. velvety or lanose. A. Submerged hyphae hyaline or nearly so. a.5 µm wide. CBS 985. F. or attached laterally to the hypha. sent in 1971 by M. consisting of a subulate basal part. 1-2. (1967). Müller under No. (9-) 13-26 (-30) µm long and (1. September 1965. ISARIA. near Boxtel. 9. Exudate and odour absent. soon becoming olivaceous black (25""k). Frankland. Gams from rust on Carex sp. Gams from leaves of Molinia caerulea. conidia. Meathrop Wood. sometimes laterally flattened.Fig. Acrodontium crateriforme.5 (-5) x (1. sent in 1969 by J. creeping or ascendent. 1..5-) 3-6 (-7. and a well developed. Great-Britain. Gams from dead spider under pine bark. smooth-walled. (19-) 13-28 (-38) µm long and 2-3 µm wide. W. pale brown. smooth-walled. A. attaining a diameter of less than 1 mm in 8 days.5-2. 15 sur 22 http://www.3-2.) Earle . September 1971. van Emden. Colonies in vitro growing rather slowly. pale olive. Devon. Perfect state unknown. 04/03/2015 16:32 . widest at the base or somewhat below the middle.5-) 3-3.5-4. appearing finely floccose. subulate or ventricose basal part. consisting of an acicular. Exudate and odour absent. with an apiculate base. Colonies in vitro growing very slowly. rarely with a transverse septum. smooth-walled. 27] arising orthotropically from undifferentiated hyphae or in groups of 1-2 (-3) from the tip of a subtending cell. Conidia hyaline. smooth-walled. Conidiogenous cells scattered. van Schothorst. often not separated from the hypha by a septum. Oberwinkler & al. guttuliform to somewhat ellipsoidal. Bad Reichenhall. H. 1-2 (-3) µm wide. 10 Rhinotrichella sp. attaining a diameter of 8-15 mm in 8 days. near Boatel. which is often not distinctly differentiated from the basal part. [p. smooth. No chlamydospores were observed. Great-Britain. up to 35 µm and 1-2 µm wide.5 µm wide. sometimes branched. Hyphae of the aerial mycelium subhyaline to brown. CBS 839. Reverse at first subhyaline or herbage green (25i). subglobose or guttuliform to ellipsoidal. type culture of Chloridium crateriforme.5-2 µm wide. (3-) 3.. denticulate rachis. Conidia hyaline. de Vries from sputum of man. CBS 151. Material examined CBS 874. East Lyn River. Bilthoven. widest at the basal septum or somewhat above. isolated as culture contaminant. 9. leg. Gams from leaves of Lazuli silvatica. Perfect state Ascocorticium anomalum. vulcano-shaped stromata. CBS 144.33.58 isolated by G. 1. creeping. Indonesia. with sharp or blunt denticles at intervals of 0.5) µm wide. and a [p. Sometimes irregular. 1967.5-3 µm.71 isolated by W..THE GENERA BEAUVERIA. conidiogenous cells. CBS 842. Submerged hyphae hyaline or nearly so. CBS 843.

conidial structures. Conidiogenous cells scattered.. saepe fasciculatae. von Klopotek under No.71 (type) isolated from human sputum. fusiformia vel ellipsoidea.5-2. 1-1. ovoidea.5-) 2-3 (-3. widest near the base or around the middle. globose (about 5 µm in diameter). rarely somewhat flexuous. ISARIA.knaw. saepe dense intricatae. with the denticles placed at random. Cellulae conidiogenae sparsae. smooth-walled. Non olet. (3.cbs. globosae.nl/publications/1001/content_files/content.71 (= NRRL 5290) sent in 1971 by D. Hyphae aeriae hyalinae vel pallide salmoneae. Hyaline. when older up to salmon (9’d). appearing floccose or farinose to powdery. CBS 848. & Vogl. Exudate and odour absent. Material examined CBS 847. Fennell under No. floccosae vel farinosae vel pulverulentae. Colonies in vitro growing rather slowly. Submerged hyphae hyaline. [p. Status perfectus ignotus. levia. and a well developed. rather thin-walled chlamydospore-like structures are very rare in the submerged mycelium. subhyaline.. Acrodontium-state of Ascocorticium anomalum. at first peach (7f). leves. sent in 1971 by A. sp.5 µm wide.Fig. No perfect state was observed.5-) 2-3 (-3. levi. Hyphae of the aerial mycelium hyaline to pale salmon. reverso luteolo vel luteo. Reverse pale luteous (19d) or luteous (17). subhyalina. e.5-5. 1-1. conidiogenous cells. 1.5 (-7) x (1. often somewhat stromatic. isolatus e sputo humano. In these genera the conidiogenous cells are either not elongating. 2. In this species the conidia may be to 4-celled and the rachis is rather short.5 µm crassae.5-) 4. 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. plerumque 15-40 x 1. crassae. CBS 846.5-2. Kikstra under No. ab A. fusiform to ovoidal. USA.71 isolated from soil of beech forest. slightly tapering towards the tip. Keswami as Cephalosporium sp. primo laete persicinae. producing more or less truncate conidia. A-13810. saepe curvata. 29] 10.5 µm et rachide tenuiter denticulata.5-) 4. often fasciculate. deinde obscuriores.5-5. R-10. solitary.5 µm wide. Some representatives of the heterogeneous pink-coloured group of Nodulisporium are morphologically intermediate between the latter genus and Acrodontium sect. 11 Coloniae in vitro post 8 dies 6-10 mm diametro. recta vel flexuosa. with an apiculate base.htm Fig. basi apiculata. arising orthotropically from undifferentiated hyphae or in groups of 1-3 (-4) from the tip of a short subtending cell. tenui-tunicatae.5) µm. Conidia hyaline. creeping or ascendent. producing conidia at their swollen tips. which is mostly not differentiated from the basal part. Germany. 1. ad 5 µm diametro.5-2.THE GENERA BEAUVERIA. repentes vel adscendentes. Texas. W. Conidia hyalina.s) µm. ad 35 µm longa. TRITIRACHIUM AND ACR. rarely with a transverse septum.71.5 (-7) x (1. Typus: CBS 847. [p. a. 16 sur 22 http://www. conidia.67 isolated as culture contaminant.5 µm wide. not alternating. CBS 580. straight or somewhat flexuous. (3. b. Acrodontium salmoneum de Hoog. Houston. Discussion The species is somewhat reminiscent of some conidial states of Xylariaceae. attaining a diameter of 6-10 mm in 8 days.5-2.71. orthotropicae e hyphis vegetativis vel ex apice cellularum lateralium singulae vel ternae (quaternae) oriuntur. slightly tapering towards the tip. c. 04/03/2015 16:32 .5 µm crassa. with rather sharp denticles at irregular intervals. Soiling. consisting of a subcylindrical basal part. Acrodontium. sent in 1971 by A.5 µm. Hyphae submersae hyalinae. Cellulae fere inflatae. mostly 15-40 µm long and 1. I. or the rachis is geniculate and cicatrized. the conidial state of Anthostomella spartii Berl. 29] which are mostly known as Nodulisporium or Geniculosporium. CBS 874. sent in 1967 by C. Kikstra. denticulate rachis. nov. . sursum modice attenuata. L. Groningen. 1. smooth. 10. a parte basilari vix delimitata constant.g. chlamydosporarum similes in mycelio submerso adsunt. e parte basilari fere cylindrica. up to 35 µm long and 1.5 µm wide.

sent in 1955 by F. CBS 847. Colonies in vitro growing slowly. 32] Fig. or procumbent and sometimes not clearly distinguishable from the vegetative mycelium. nov. Material examined Herbarium specimen Virgaria hydnicola in herb. Acrodontium section Grisea de Hoog.5-) 1. at first rosy buff (17’’d). Conidiogenous cells arising plagiotropically as a part of compound. Bot. seu procumbentia in hyphas vegetativas transeuntia. Conidia subhyaline. (15-) 18-30 (-42) µm long. Conidia modice pigmentata vel subhyalina. Submerged hyphae hyaline. a. attaining a diameter of 5 mm in 8 days. conidial structures on natural substrate. St. conidiogenous cells. with slightly apiculate base.cbs.THE GENERA BEAUVERIA. from creeping hyphae.Rep. appearing powdery-velvety. often one of the branches forms a conidiogenous cell while the other ramifies again. conidial structures. Cellulae conidiogenae e hyphis fertilibus vel e hyphis aeriis vegetativis oriuntur. verticillata vel simplicia. var. 1953. N. smooth. pale brown. basi apiculata. 31] erecta crassi-tunicata et deorsum fusca. stalk up to 175 µm high and 2-3 µm wide. seu [p. Species typica: Tritirachium heimii (Saccas) Langer. and a straight rachis. griseum Fassatiová. 11. c. Br.5-) 2-3 (-3.55 isolated as contaminant on keratin. conidial structures in culture. procumbent. c. brown towards the apex. consisting of an elongate basal part.5-1. e parte basilari lageniformi vel elongata et rachide angustiore recta denticulis obtusis obtecta constant. Acrodontium hydnicola (Peck) de Hoog. conidia. 1.5) x (2-) 2-2. Differentiated conidiophores mostly arising from creeping hyphae. mycol. smooth. straight or slightly flexuous.5 µm wide. (1.. 12. which is differentiated from the basal part and bears a series of blunt denticles. and a well developed.Fig. more or less differentiated conidiophores. e. (1. when older cinnamon (15"i). Discussion 04/03/2015 16:32 . 6: 56. or plagiotropically when little differentiated.and rather thin-walled. often also orthotropically from undifferentiated hyphae. Hyphae of the aerial mycelium hyaline to brownish. with blunt denticles about 1 µm long. at intervals of 0. slightly swollen at the base and tapering towards the tip. consisting of a flask-shaped or elongate basal part.5 (-3) µm wide. from type collection.nl/publications/1001/content_files/content. the latter being about as wide as the rachis.5 µm. d. Reverse cinnamon (15"i). Conidia somewhat pigmented to subhyaline. nov. . Soc. NYS (type). often forming a compact stroma. up to 25 µm long and mostly 1 µm wide. sect. denticulate rachis.5-3. No chlamydospores were observed. Mus.71. with verticillate or simple branching. 31: 604. Living strain CBS 349. conidia. f. Y.Trans. 11. Exudate and odour absent. Acrodontium salmoneum. ISARIA.5 (-3) µm. with an apiculate base. globose or guttuliform. tapering towards the tip.. lighter towards the apex. b. thin-walled. Blank. comb. on resupinate basidiomycete on wood. branched verticillately or dichotomously over the whole length at intervals of 10-20 µm. 12 Virgaria hydnicola Peck . 1917. b. 1889 (basionym) = Tritirachium hydnicola (Peck) Hughes Can. hyaline in the lower part. Perfect state unknown. globose to subglobose. either erect. 17 sur 22 http://www. Acrodontium hydnicola. conidial structures in culture.htm Fig. J.knaw. Conidiogenous cells as distal branches of conidiophores or arising orthotropically from undifferentiated hyphae. North Elba.. ? Clonostachys dichotoma Bayliss Elliott . USA. [p.5-2. TRITIRACHIUM AND ACR. thick-walled and dark brown in the lower part.5-2. 42: 32. subglobosa vel guttuliformia. 1.5 µm wide. a. Conidiophora composita plerumque orthotropica e hyphis repentibus oriuntur. Conidiophores arising orthotropically. sursum clariora.

nl/publications/1001/content_files/content. stalk up to 650 µm long and (2-) 3-4. champ. Submerged hyphae hyaline. 1827 (basionym) Beauveria virella (Fr. consisting of an elongate basal part. champ. Colonies in vitro growing slowly. 33] more or less irregularly branched. only a secondary collection from the same substrate and locality (B) was studied.53.Rech.htm The conidiophores are often little differentiated from the vegetative mycelium. up to 30 µm long and mostly 1 µm wide. attaining a diameter of 4 mm in 8 days. the upper part bearing 2-5 (7) whorls of 1-3 side branches or conidiogenous cells. with blunt denticles [p. often also orthotropically from undifferentiated hyphae. 63: 454. . 13 Beauveria simplex Mangenot . and a well developed. certains bois décomp. nov. Bot. with blunt denticles about 1 µm long at intervals of 0. is very closely related. denticulate rachis. certains bois décomp.5 µm wide. 18 sur 22 http://www. method. subglobose to guttuliform. comb. 1-2 µm wide.5) x (1-) 1. Without living cultures its identity cannot be established. 1952. or pale brownish when bearing conidiophores. probably it is lost.Fig.71 isolated by W. suberect. 63: 450. type culture of Beauveria simplex. (2-) 2. smooth. c. . Revue gen. smooth. a.5-) 1.5-3. often also orthotropically from undifferentiated hyphae.1952 (basionym). b. Conidia light brown. Perfect state unknown. . 12-28 µm long. According to the original description and illustration there is much similarity with A. France. often forming a compact stroma. appearing velvety. at first rosy buff (15"f). paler towards the margin.5 (-6) µm wide. It is rather dark and firm and its forms small synnemata. Reverse sepia 17"k). 491. when older hazel (17"‘). Hyphae of the aerial mycelium hyaline to brownish. brown. with an apiculate base. 13. nov. September 1971. ISARIA. with thin transverse septa. Exudate and odour absent. 35] about 1 µm long at intervals of 0. 1. Hyphae of the aerial mycelium hyaline. Gams from Oligoporus rennyi on pine wood. CBS 845. Conidiogenous cells arising plagiotropically as a part of compound conidiophores. comb. the upper part bearing 1-2 (3) whorls of 1-2 (3) side branches or 1-4 conidiogenous cells. Isaria acaricida Pat.5-1 µm. sometimes not markedly differentiated. Kampina heath.5-3 (-3. 14 Botrytis virella Fr. at first vinaceous buff (17"‘f). ascendent to suberect. tapering towards the tip. Conidiophores arising orthotropically from creeping hyphae. tapering towards the tip and sometimes constricted in the upper part. conidia. Perfect state unknown. paler near the base. consisting of a flask-shaped to elongate basal part. and a wel developed. globose to subglobose. 12.5 (-3. Conidia brownish.5-) 2-3 (-3) µm. side branches bearing 1-3 (5) whorls of conidiogenous cells or branches of second order.53.5 (-3) µm No chlamydospores were observed. smoothand somewhat thick-walled.5-2. Scand.5-1 µm. p.knaw. smooth-walled. Conidiogenous cells arising plagiotropically as a part of compound conidiophores. 36. Mangenot under No. hyaline or nearly so. 1-3. sent by F. TRITIRACHIUM AND ACR. d. Acrodontium virellum (Fr. when older up to sepia (13"k).Fig. near Boxtel. p. stiff. [p. 04/03/2015 16:32 . subhyaline. conidial structures. 1084. appearing velvety to somewhat powdery. hydnicola.Summa Veg. straight or slightly flexuous. conidiogenous cells. 13. hyaline towards the tip. No chlamydospores were observed.Rech. P. 1952. isolated from decaying trunk of Fraxinus excelsior. mostly 4-8 x 2-3 µm.cbs.5-3 µm wide. Colonies in vitro growing slowly.5) x (1.) Mangenot . Fig. Exudate and odour absent. CBS 127. not markedly differentiated. Acrodontium simplex (Mangenot) de Hoog. up to 5 µm long and 1-2 µm wide.THE GENERA BEAUVERIA. 40. Conidiophores arising orthotropically from creeping hyphae. Acrodontium simplex. Revue gen.and thin-walled.. method. slightly thickwalled. straight or slightly flexuous. Reverse fuscous (13 ""k).. smooth and slightly thick-walled. No type material could be examined. Submerged hyphae hyaline. sometimes guttuliform.5-3 µm wide.) de Hoog. stalk up to 110 µm high and 1. 1952. attaining a diameter of 5 mm in 8 days. In poorly developed subcultures on PDA they may be indistinguishable. Material examined CBS 127. denticulate rachis. No material of Clonostachys dichotoma was available in K and BPI. brown. sometimes swollen at the base up to 3 µm tapering towards the tip. (1. with an apiculate base. 1.5-4 µm wide. Bot.

knaw. France. brown. Oldenburg. b. sent in 1953 by F. tapering towards the tip. c. smooth-walled. griseum Fassatiová . Exudate absent. attaining a diameter of 4-7 mm in 8 days.5-1 µm. var. with an apiculate base. straight or somewhat flexuous. Bohemia.5-) 2-4.53. Reverse herbage green (23m) to olivaceous black (21""m). 1-3. up to 3 5 µm long and mostly 1 µm wide. Discussion No type material is left by Fries in UPS. Mallorca. July 1970. CBS 129. Fig.5 µm long at intervals of 0.Antonie van Leeuwenhoek 8: 116. CBS 671.Ceská Mykol.5) x 2-3 µm No chlamydospores were observed. Conidia pale olive brownish. ISARIA. 37] long. erect. appearing powdery to somewhat velvety.and thin-walled. Czechoslovakia. isolated from wood. pale brown. conidiogenous cells. Perfect state unknown. swollen up to 5 µm below the middle. conidia Colonies in vitro growing rather slowly. Acrodontium griseum (Fassatiová) de Hoog. d. griseum. dark brown to red brown towards the base. (1. Acrodontium virellum. and a well developed. Material examined Herbarium specimen Beauveria heimii Saccas in herb. nov. a. consisting of a flask-shaped or elongate basal part. Malençon.70. Hyphae of the aerial mycelium hyaline or brownish. a. The conidiogenous cells are cylindrical. conidial structures.htm Fig. It is reminiscent of the genus Nodulisporium and should 04/03/2015 16:32 . Conidiophores arising orthotropically from creeping hyphae.Fig. on bark of Pinus halepensis. stalk (20-) 50-310 (-450) x 3-4. denticulate rachis. mostly 6-12 µm [p. Submerged hyphae hyaline.5 µm the upper part bearing 2-4 (5) whorls of 2-3 (4) side branches or conidiogenous cells. Conidiogenous cells arising plagiotropically as a part of compound conidiophores. sent in 1971 by A.5 µm wide. subglobose. 15 Tritirachium heimii (Saccas) Langer.cbs. 15. CBS 849. 1. Doubtful or excluded species of Beauveria. Argonne. suberect and irregularly verticillate. The characters given in the original description are hardly sufficient to ascertain its identity. 19 sur 22 http://www. leg. stiff. 1942. sent under No. smooth.5-3 µm wide. globose. Material examined CBS 671. conidial structures. comb.53 isolated from decaying wood. PC. TRITIRACHIUM AND ACR. branches of second or third order often present. T-23.nl/publications/1001/content_files/content. often forming small sporodochium-like cushions. 1972 (basionym). b. The conidiophores are subhyaline.71 isolated from peat.and thick-walled. 14. odour very slightly sourish. Acrodontium griseum.5 (-5. c. 1169. 1952. November 1970. Germany. at first vinaceous buff (17’’’ b). W. Living strain CBS 129.. type culture of Tritirachium heimii var. . often also orthotropically from undifferentiated hyphae. Tritirachium and similar genera Tritirachium cinnamomeum van Beyma . 14.53 [p. 25: 116. smooth.70. with blunt denticles about 1. 36]. Mangenot as Beauveria virella. smooth. markedly differentiated. G. guttuliform or fusiform. bearing an irregularly geniculate rachis.THE GENERA BEAUVERIA.. conidia. closely septate. The present description is based on strain CBS 129. Spain. when older herbage green (23m). von Klopotek under No.

1948. The species was published without a Latin diagnosis. 1949. var. 1939 = Spicaria rileyi (Farlow) Charles . The fungus is congeneric with Oospora meliolae Hansf.Arb. BundAnst. Papua-New Guinea p. Méd. which is presumably a misprint. Persoon (1822) thought this species to be related to Sporotrichum densum Link. hum. Most likely an Acrodontium species. 1948 = Tritirachium heimii (Saccas) Langer. 39] Acknowledgements The author is indebted to Dr J. Berlin. Br. 10: 295.) Fr. mycol.Annls mycol. Paris III: 133. 1905 = Beauveria rubra (Baquis) Langer. nov. 1942. I4: 13 5.Microf. 1883: 121 = Beauveria rileyi (Farlow) Gösswald . Res.. According to the description and illustrations a species reminiscent of Tritirachium oryzae was meant. 1924 = Beauveria coccorum (Petch) Linder . Land. biol.) Saccas . 1912 = Tritirachium spicatum (Ferr. hyaline. Rhinotrichum album Petch . 34: 947. Acad.Indian J. 1936. Br. creeping hyphae of variable width. Land. under different names. 1934 = Tritirachium viannai (de Mello) Langer. von Arx for his valuable suggestions and to Dr M.) Limber . The species was described from various insects as being very variable.-nat. I: 72. No material is left in PC.. Forstw. .Rep. . Sporotrichum flavicans Fr. 1925) described the same species twice. he gave a description which does not correspond with the type material. 1965. Forstw. Doubtful species. No material is left in HCIO and PC.u.Trans. 20 sur 22 http://www. Solomon Isl. A. Possibly a Nodulisporium-like fungus was meant. Soc. These give rise to guttuliform conidia. 1947. Trichophyton viannai de Mello .Mycol. [p. A. Fr. Müller .cbs.Physis. R. hum.Annls Parasit. No material left in PAD..Trans. blunt denticles. ISARIA. According to the type material (CMI) the species has phialides.htm better be referred to as Nodulisporium cinnamomeum (van Beyma) de Hoog. 1933 = Beauveria paranensis (Marchionatto) Gösswald . 1971. eur. In 1931 he discovered his error and maintained the oldest name. & Massa . [p. comp. Beauveria petelotii Vincens -Bull. Dep.. however. lo: 181. B. spicatum Ferr.Arb. 22: 434. 22: 98. No material was left in PAD and PC.Wiss. TRITIRACHIUM AND ACR. I am much obliged to Dr W. per Fr. On the dry specimens (K) it could not be decided whether the conidia were born in chains as described by Petch (1931). Possibly different Beauveria species were described under the same name (Fassatiová. comp. math. -Revue Mycol. in Ferr. Gonatorrhodiella coccorum Petch . simplex. did not allow to recognize the fungus. Paris III: 133. was concerned. Agric. S. Gams for his stimulating help in preparing the manuscript 04/03/2015 16:32 . Racodium (?) entomogenum Pers. Petch (1924b. Bull. ocul. griseum or A. mycol.Annls Parasit. When Linder (1942) made the combination Beauveria coccorum. Beauveria heimii Saccas .knaw. 13: 64. 38] Beauveria cretacea (van Beyma) Matsushima . 1822. According to the type material (FH) the species has phialides.u. Verticillium rubrum Baquis . Z. Med.Annali Ottal. Aires II: 348. with numerous short. med.THE GENERA BEAUVERIA. The fungus has smooth. Sporotrichum paranense Marchionatto .. 5: 228. Humboldt-Univ.Revue Mycol. Clin. Donk for his advice concerning nomenclature. BundAnst. The type material (L). 1939 = Paecilomyces paranensis (Marchionatto) G. The original description was very scanty. comb. 1925. Botrytis rileyi Farlow in Riley . bot. 1917 = Beauveria viannai (de Mello) Langer. II: 258.1934 = Tritirachium rubrum (Baquis) Langer. 1966). 1947. Soc. 22: 434. 61: 132.Mycologia 28: 398.Bull. No material is left in PC.Lloydia 5: 206. Soc. . = Isaria felina (DC. 193 S. biol. Somewhat further he mentioned the name Beauveria coccospora (Petch) Linder. 7. 22: 98. such as A. .Revue Mycol.nl/publications/1001/content_files/content. 1940 = Beauveria spicata (Ferr. 13: 63. U. Acad. I4: 780.Mycologia 32: 29.

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K. P. 22 sur 22 http://www. Agric.. 1886 . PflSchutz 62: 544-549. 6: 68-69. R..Revue Path.. . 1955 .cbs.Rhinocladiella..Studies in artificial cultures of entomogenous fungi.Spec. Petch.Bull. . Kew.Sylloge Fungorum 10. H. Cornell Univ. Soc. Schol-Schwarz. bot. Br. Fr. Revue Mycol. le Beauveria heimii sp.) Link.. Taiwan 8: 1-268. R. Ent.. L. sp. B. . ISARIA. 25: 250-265.Fungi veneti vel novi critici. 9: 136-139. its synonym Fonsecaea and its relation to Phialophora. T.Trans. 1912 .Descriptive catalogue of Taiwan (Formosan) fungi. W. M. 19: 34-38.Isaria. . PflPath. Schaerffenberg.Archwm Nauk biol. agric. Saccardo. Soc.) Link and B. Bally and B. . nouveau genre de Verticilliacées.Beauveria. J. A. Sawada. Soc. Buitenz.nl/publications/1001/content_files/content. A. 10: 112-131. 1895 . 29: 34-40. veg. Saccas.A mycological colour chart. nov. 1963 .Die Hauptfruchtform (Ascus-Form) von Beauveria bassiana (Vuill. A..Sylloge Fungorum 4.Studja nad grzybami owadobójczemi Polski. Coll. P.Étude morphologique et biologique d’un nouveau champignon papyricole. A. Rayner. Schwarz. Saccardo... 13: 61-81.Antonie van Leeuwenhoek 34: 119-152.Bull. Vincens.Observations sur le Sclerotium orizae faites en Cochinchine.Fungi italici delineati 1. densa (Vuill. .htm Petch. A. TRITIRACHIUM AND ACR. 1879 . A. W. agric.. P. Jard... C. 1892 . M. bot. mycol. Vuillemin.Trans. . 1923 . 1934 . . B. mycol. Publ. 1959 ..Bull. Part XI . Siemaszko.Commonwealth Mycological Institute. bassiana Bals. 6: 1-83.Botrytis stephanoderis n. Exp.knaw.. V. Fr. 1879 . Microbiol. 04/03/2015 16:32 . Saccardo. . 1968 .Z. T. 1970 . 1924 ...THE GENERA BEAUVERIA.Padova. Pettit. . Mycologia venetae.Can. P. P. Br. .Studies on Isaria cretacea: antibiotic factors produced by strain B.. & Vining. . 1937 .Padova. 1941 . Taber. . Ser.Notes on entomogenous fungi. Stn 97: 339-378. XIII. A. . PflKrankh.. . W.. Saccardo. 1948 .Michelia 2:528-563.Padova. F.