Full text of DOI:10.1016/S0953-7562(08)61976-0

1123
Mycol. Res. 105 (9): 1123–1133 (September 2001). Printed in the United Kingdom.
New and interesting Biscogniauxia taxa, with a key to the
world species
Yu-Ming JU1 and Jack D. ROGERS2
"
Institute of Botany, Academia Sinica, Nankang, Taipei 115, Taiwan.
Department of Plant Pathology, Washington State University, Pullman, Washington 99164-6430, USA.
#
E-mail: yumingju!gate.sinica.edu.tw
Received 9 February 2001; accepted 24 April 2001.
Six new taxa of Biscogniauxia are described from Taiwan, including four new species: B. ambiens, B. cylindrispora, B. formosana, and B.
latirima; and two new varieties: B. albosticta var. orientalis and B. formosana var. kentingensis. Their cultural and anamorphic data are
also provided. A description is provided for B. doidgeae based on its type. Cultural and anamorphic data are given for B. philippinensis
var. microspora. A key to the 57 known species of Biscogniauxia in the world is also given.
INTRODUCTION
terminating with 3–5 conidiogenous cells arising in whorls
from the main axis and\or from major branches. Conidiogenous
cells cylindrical, hyaline to yellowish, finely roughened,
8–16i2n5–3n5 µm, bearing poroid conidial secession scars on
apical region, with apices becoming distorted from production
of numerous conidia. Conidia produced holoblastically in
sympodial sequence, hyaline, smooth, ellipsoid to obovoid,
5–7i2n5–3 µm, with flattened base indicating former point
of attachment to conidiogenous cell.
Notes: The typical variety of B. albosticta is known only
from A. P. Morgan’s collections from Preston, Ohio, USA.
Var. albosticta is much like var. orientalis except for having
larger ascospores 15n5–19n5i6n5–8n5 µm and larger ascal
apical rings. They can be distinguished from the other
Biscogniauxia taxa by having bilateral ascospore germ slits,
punctate ostioles filled with white material, and narrow-ended
ascospores.
Biscogniauxia was recently monographed by Ju et al. (1998).
Forty-nine taxa were included. Six new taxa from Taiwan are
added to the genus; they are described from collections made
by Y.-M. J. and his colleagues. Additional information is given
on two previously described taxa. Cultural data are from
cultures grown on 2% Difco oat meal Agar (OMA). Colour
designations are from Rayner (1970). Information on allo-
physes, a newly described hamathecial element, is to be found
in Rogers et al. (2000). Cultures are deposited in the
American Type Culture Collection (ATCC) and the Culture
Collection and Research Center (CCRC) in Taiwan.
TAXONOMY
Biscogniauxia albosticta var. orientalis Y.-M. Ju & J. D.
Rogers, var. nov.
(Figs 1–5, 44)
A varietate typica differt in ascosporis 13–16i5–7 µm et in
annulo apicali asci 2 µm altoi3n5 µm lato.
Typus: Taiwan: Nan-tou Co.: Yu-chee, Lien-hwa-chee, on bark, 12
May 2000, Y.-M. Ju 89051204 [cultured] (HAST – holotypus; WSP
70162 – isotypus).
Biscogniauxia ambiens Y.-M. Ju & J. D. Rogers, sp. nov.
(Figs 6–10, 45)
Etym.: Refers to the germ slit running around the ascospore
longitudinally.
Differs from the typical variety in having ascospores
13–16i5–7 µm and the ascal apical ring 2 µm highi3n5 µm
broad.
Stromata applanata, 0n7–4n5 cm longai0n4–1n4 cm latai0n3 mm
crassa; stratum externum dehiscens fuscum tenue; superficies matura
viridifusca; textura sub superficie et inter perithecia carbonacea;
textura sub peritheciis inconspicua. Perithecia globosa, 0n2–0n3 mm
diam, singularis vel aliquot per canalem communem ostiolarem
emittentibus. Ostiola umbilicata et saepe substantia alba repleta. Asci
80–100 µm longitudine totai5–6 µm crassi, partibus sporiferis
75–90 µm longitudine, stipitibus 5–15 µm longitudine, annulo apicali
in liquore iodata Melzeri cyanescente, discoideo, 0n5 µm altoi2 µm
lato. Ascosporae brunneae vel fuscae, unicellulares, ellipsoideae fere
equilaterales apicibus latis, leves, 10–13i4–5(k6) µm, rima
germinativa cingenti praeditae. Paraphyses et allophyses inclusae.
Culture and anamorph: Colonies on OMA at ca 20 mC and
under 12 h fluorescent light reaching the edge of 9 cm Petri
dish in 2–2n5 wk, whitish, felty, azonate, with diffuse margins,
becoming Olivaceous Buff (89) and cottony. Reverse Dull
Green (70). Sporulating regions near inoculation point, whitish.
Conidiogenous structure with Periconiella-like branching pattern
as defined in Ju & Rogers (1996). Conidiophores brownish,
roughened, composed of a main axis, 3n5–4n5 µm broad, and
sometimes one or more major branches, branches mostly

Biscogniauxia
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Figs 1–17. Biscogniauxia albosticta var. orientalis, B. ambiens, B. cylindrispora. Figs 1–5. B. albosticta var. orientalis (holotype). Figs 1–2.
Stromata. Fig. 3. Stromatal surface with ostiolar openings. Fig. 4. Amyloid ascal apical ring and ascospores. Fig. 5. Ascospore showing
the germ slit. Figs 6–10. B. ambiens (holotype). Figs 6–7. Stromata. Fig. 8. Stromatal surface with ostiolar openings. Fig. 9. Amyloid
ascal apical ring and ascospores. Fig. 10. Ascospores showing the germ slits. Figs 11–17. B. cylindrispora (holotype). Figs 11–12.
Stromata. Fig. 13. Stromatal surface with ostiolar openings. Fig. 14. Multiple laterally compressed perithecia with covering stromatal
tissue fractured away showing common ostiole (arrow head) which is overlain with white material. Fig. 15. Amyloid ascal apical ring.
Fig. 16. Ascospore showing the germ slit. Fig. 17. Ascospores. Bars in Figs 1, 6, 11 l 0n5 cm; Figs 2, 7, 12 l 1 mm;
Figs 3, 8, 13–14 l 0n1 mm; Figs 4–5, 9–10, 15–17 l 10 µm.
Typus: Taiwan: Nan-tou Co.: Yu-chee, Lien-hwa-chee, on bark, 12
broadi0n3 mm thick; outer dehiscing layer dull brown, thin;
May 2000, Y.-M. Ju 89051212 [cultured] (HAST – holotypus; WSP
70163 – isotypus).
mature surface dark brown with olivaceous tinge; car-
bonaceous immediately beneath surface and between
Stromata applanate, 0n7–4n5 cm longi0n4–1n4 cm perithecia; tissue beneath perithecia inconspicuous. Perithecia

Y.-M. Ju and J. D. Rogers
1125
Figs 18–43. Biscogniauxia doidgeae, B. formosana, B. formosana var. kentingensis, B. latirima, B. philippinensis var. microspora. Figs 18–22. B.
doidgeae (lectotype). Fig. 18. Stroma. Fig. 19. Stromatal surface with ostiolar opening (arrowhead). Fig. 20. Vertical section of a stroma
showing perithecia and ostiolar canals. Arrowheads point to the carbonaceous tissue encasing each individual ostiole. Fig. 21. Amyloid
ascal apical ring and immature ascospores. Fig. 22. Ascospores. Figs 23–28. B. formosana (holotype). Figs 23, 24. Stromata. Fig. 25.
Stromatal surface with ostiolar openings. Fig. 26. Amyloid ascal apical ring and ascospore. Fig. 27. Ascospore showing the germ slit.
Fig. 28. Ascospores. Figs 29–34. B. formosana var. kentingensis (holotype). Figs 29–30. Stromata. Fig. 31. Stromatal surface with
ostiolar openings. Fig. 32. Amyloid ascal apical ring and ascospores. Fig. 33. Ascospores showing the germ slits. Fig. 34. Ascospore.
Figs 35–40. B. latirima (holotype). Fig. 35. Stroma. Fig. 36. Stromatal surface with ostiolar openings. Fig. 37. Vertical section of a
stroma showing perithecia sharing an ostiolar canal. Fig. 38. Amyloid ascal apical ring and ascospores. Fig. 39. Ascospore showing the
germ slit. Fig. 40. Ascospores. Figs 41–43. B. philippinensis var. microspora. Fig. 41. Amyloid ascal apical ring and ascospore. Fig. 42.
Ascospore showing the germ slit. Fig. 43. Ascospores. Bars in Fig. 18 l 2 mm; Figs 23, 29 l 0n5 cm; Figs 24, 30, 35 l 1 mm;
Figs 19, 20, 25, 31, 36, 37 l 0n1 mm; Figs 21, 22, 26–28, 32–34, 38–43 l 10 µm.

Biscogniauxia
1126
45
44
46
47
Figs 44–47. Conidiogenous structures and conidia. Fig. 44. B. albosticta var. orientalis (holotype). Fig. 45. B. ambiens (holotype).
Fig. 46. B. cylindrispora (holotype). Fig. 47. B. philippinensis var. microspora (89041101). Bar l 10 µm.

Y.-M. Ju and J. D. Rogers
1127
48
50
49
Figs 48–50. Conidiogenous structures and conidia. Fig. 48. B. formosana (holotype). Fig. 49. B. formosana var. kentingensis (holotype).
Fig. 50. B. latirima (holotype). Bar l 10 µm.

Biscogniauxia
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Typus: Taiwan: Taipei Co.: Sun-shei, on bark of Cinnamomum, 27
Sep. 2000, Y.-M. Ju 89092701 [cultured] (HAST – holotypus; WSP
70164 – isotypus).
spherical, 0n2–0n3 mm diam, with one to several individuals
discharging through a single ostiolar canal. Ostioles lower than
stromatal surface, with openings punctate and often filled with
white material. Asci 80–100 µm total lengthi5–6 µm broad,
the spore-bearing parts 75–90 µm long, the stipes 5–15 µm
long, with apical ring bluing in Melzer’s iodine reagent,
discoid, 0n5 µm highi2 µm broad. Ascospores brown to dark
brown, unicellular, ellipsoid, nearly equilateral, with broadly
rounded ends, smooth, 10–13i4–5(k6) µm, with straight
germ slit spore-length on both sides. Both paraphyses and
allophyses present.
Culture and anamorph: Colonies on OMA at ca 20 m and
under 12 h fluorescent light reaching the edge of 9 cm Petri
dish in 2n5–3 wk, whitish, velvety, azonate, with fimbriate
margins, becoming Isabelline (65) to Olivaceous (48), with
Amber (47) to Citrine (13) pigments diffused into medium.
Reverse Greenish Olivaceous (90) to Dull Green (70).
Sporulating regions scattered over entire surface of colony,
whitish. Conidiogenous structure with Nodulisporium-like
branching pattern as defined in Ju & Rogers (1996).
Conidiophores yellowish to pale brown, roughened, di- or
trichotomously branched, sometimes with additional branches
arising from the first level of conidiogenous regions, 3n5–4 µm
diam, with 2–3 conidiogenous cells arising from each terminus.
Conidiogenous cells cylindrical, hyaline, finely roughened,
10–24i2n5–3n5 µm, bearing poroid conidial secession scars
on apical region, with apices becoming distorted from
production of numerous conidia. Conidia produced holo-
blastically in sympodial sequence, hyaline, smooth, obovoid
to clavate, 7–10i2n5–3 µm, with flattened base indicating
former point of attachment to conidiogenous cell.
Stromata
applanate,
0n3–2 cm
longi0n3–1n5 cm
broadi0n6–0n8 mm thick; outer dehiscing layer dull brown,
thin; mature surface dull black; carbonaceous immediately
beneath surface and between perithecia; tissue beneath
perithecia inconspicuous. Perithecia obovoid to tubular,
0n05–0n1 mm diami0n5–0n6 mm high, laterally compressed,
with numerous individuals arranged in rosettes and dis-
charging through a single ostiolar canal. Ostioles lower than
stromatal surface, with openings punctate, often surrounded
with numerous radiating furrows. Asci 170–205 µm total
lengthi15–18 µm broad, the spore-bearing parts 160–
185 µm long, the stipes 10–20 µm long, with apical ring
bluing in Melzer’s iodine reagent, discoid, 3–3n5 µm highi6–
7 µm broad. Ascospores blackish brown, unicellular, cylindrical
to, less frequently, ellipsoid, with broadly rounded ends,
smooth, 20–26i10–14(–16) µm, with straight germ slit
spore-length or nearly so. Paraphyses and allophyses present.
Culture and anamorph: Colonies on OMA at ca 20 m and
under 12 h fluorescent light reaching the edge of 9 cm Petri
dish in 2 wk, whitish, felty, azonate, with diffuse margins,
becoming Olivaceous (48) to Dull Green (70) from centre
outwards. Reverse Greenish Olivaceous (90). Sporulating
regions scattered over entire surface of colony, whitish.
Conidiogenous structure with Periconiella-like branching pattern
as defined in Ju & Rogers (1996). Conidiophores hyaline to
yellowish, finely roughened, composed of a main axis,
3n5–5 µm broad, and sometimes one or more major branches,
branches mostly terminating with 3–5 conidiogenous cells
arising in whorls from the main axis and\or from major
branches. Conidiogenous cells cylindrical, hyaline to yellowish,
smooth to finely roughened, 8–19i3n5–4n5 µm, bearing
poroid conidial secession scars on apical region, with apices
becoming distorted from production of numerous conidia.
Conidia produced holoblastically in sympodial sequence,
hyaline, smooth, ellipsoid, obovoid to clavate, 5n5–
9i2n5–3(k3n5) µm, with flattened base indicating former
point of attachment to conidiogenous cell.
Notes: Seven other Biscogniauxia species also have the germ
slit on both sides of the ascospore: B. albosticta, B. albosticta
var. orientalis, B. cinereolilacina, B. nawawii, B. querna, B. repanda,
and B. schweinitzii (Ju et al. 1998, Whalley et al. 2000). B.
ambiens can be separated from the species with bilateral germ
slit by the combination of the following characteristics:
applanate stromata with a dark brown surface tinged
olivaceous, carbonaceous stromatal tissue homogeneously
distributed between perithecia, punctate ostiolar openings,
and ellipsoid ascospores not laterally compressed.
Notes: This fungus, like B. communapertura and B. plana, is
characterized by having numerous perithecia arranged in
rosettes and served by a single ostiolar canal (Fig. 14). In other
Biscogniauxia taxa, a single ostiolar canal serves one single
perithecium or is shared by several perithecia. Biscogniauxia
communapertura and B. plana can readily be separated from B.
cylindrispora by their smaller, paler, ellipsoid ascospores.
Furthermore, Biscogniauxia communapertura differs from B.
cylindrispora in having a much less than spore-length ascospore
germ-slit and in having coarsely papillate ostiolar openings,
whereas B. plana differs in having ostiolar openings punctate
or slightly papillate rather than exclusively punctate. Ju &
Rogers (1999: 365) reported an unnamed Biscogniauxia species
from Taiwan which is based on a depauperate specimen, Ju
80091405. It appears to be B. cylindrispora; the larger
perithecial diameter reported for that specimen is seemingly
resulted from destruction of the thin, fragile stromatal tissue
between the perithecia in a rosette.
Biscogniauxia cylindrispora Y.-M. Ju & J. D. Rogers, sp.
nov.
(Figs 11–17, 46)
Etym.: After the ascospore shape.
Stromata applanata, 0n3–2 cm longai0n3–1n5 cm latai0n6–
0n8 mm crassa; stratum externum dehiscens fuscum tenue; superficies
matura atra; textura proxime sub superficie et inter perithecia
carbonaceae; textura sub peritheciis inconspicua. Perithecia
obovoidea vel tubularia, 0n05–0n1 mm diami0n5–0n6 mm alta,
compressa, multa in circula deposita et per canalem communem
ostiolarem emittentibus. Ostiola umbilicata, saepe sulcis patulis prae-
dita. Asci 170–205 µm longitudine totai15–18 µm crassi, partibus
sporiferis 160–185 µm longitudine, stipitibus 10–20 µm longitudine,
annulo apicali in liquore iodato Melzeri cyanescente, discoideo,
3–3n5 µm altoi6–7 µm lato. Ascosporae fuscae, unicellulares,
cylindricae vel infrequenter ellipsoideae apicibus latis, leves, 20–
26i10–14(–16) µm, rima germinativa longa recta praeditae. Para-
physes et allophyses inclusae.

Y.-M. Ju and J. D. Rogers
1129
long, with apical ring bluing in Melzer’s iodine reagent,
discoid, 2–3 µm highi4n5–6 µm broad. Ascospores dark
brown, unicellular, fusoid- to ellipsoid-inequilateral, with
narrowly rounded ends frequently pinched, smooth,
(18–)19–24(–25)i8n5–10 µm, with straight or slightly
oblique germ slit spore-length or nearly so on less convex
side.
Biscogniauxia doidgeae (J. H. Miller) Whalley & Lœssøe, in
Whalley, Læssøe & Kile, Mycol. Res. 94: 239 (1990).
(Figs 18–22)
Hypoxylon doidgeae J. H. Miller, Monogr. Hypoxylon: 110
(1961); as ‘doidgei’.
Typus: South Africa: Natal, on wood, 1936, Morgan, K. E. &
Doidge, E. M. [Union of South Africa Herb. 30256] (BPI
588595 – lectotypus, hic designatus).
Culture and anamorph: Colonies on OMA at ca 20 m and
under 12 h fluorescent light reaching the edge of 9 cm Petri
dish in 2 wk, whitish, felty, azonate, with diffuse margins,
becoming Olivaceous Buff (89) to Greenish Olivaceous (90).
Reverse Dull Green (70). Sporulating regions scattered over
entire surface of colony, whitish. Conidiogenous structure with
Periconiella-like branching pattern as defined in Ju & Rogers
(1996). Conidiophores yellowish to brownish, coarsely
roughened, composed of a main axis, 3n5–5 µm broad, and
sometimes one or more major branches, branches mostly
terminating with 3–5 conidiogenous cells arising in whorls
from the main axis and\or from major branches. Conidiogenous
cells cylindrical to barrel-shaped, yellowish to brownish, finely
roughened, 5–10(k13)i3–5 µm, bearing poroid conidial
secession scars on apical region, with apices becoming
distorted from production of numerous conidia. Conidia
produced holoblastically in sympodial sequence. Conidia
hyaline, smooth, obovoid, 4n5–5n5i2n5–3n2 µm, with
flattened base indicating former point of attachment to
conidiogenous cell.
Notes: Biscogniauxia formosana is likely to be confused with
B. capnodes var. rumpens which has a similar ascospore size
range (14k)15–22n5(k24)i8–11(k12) µm. Two types of
ostiolar openings are always found on stromata of B. capnodes
var. rumpens and other varieties of B. capnodes: slightly
papillate and punctate. Ostiolar openings in B. formosana are
exclusively punctate. In addition, ascospores of the B. capnodes
complex are more or less equilateral, whereas those of B.
formosana are conspicuously inequilateral.
Stroma pulvinate, 1n7 cm diami2 mm thick; outer
dehiscing layer unknown; mature surface dull ochraceous
brown, plane; dark brown, woody immediately beneath
surface and between perithecia, with carbonaceous tissue
encasing each individual ostiole; tissue beneath perithecia
inconspicuous. Perithecia tubular, 0n4 mm diami1n7 mm high.
Ostioles lower than stromatal surface, with openings umbilicate,
often overlain with white substance. Asci fragmentary, with
apical ring bluing in Melzer’s iodine reagent, discoid, 1 µm
highi3–3n5 µm broad. Ascospores light brown to brown,
ellipsoid-inequilateral, with one end narrowly rounded and the
other bearing a hyaline to pale brown cellular appendage
1n5–2 µm longi2n5–3 µm broad, smooth, 8–11n5 (not
including appendage)i4–5n5 µm, apparently lacking a germ-
slit.
Notes: Biscogniauxia doidgeae is like B. arima and B. waitpela
in lacking an ascospore germ slit but can be readily separated
from the latter two taxa in having a cellular appendage on one
end of the ascospore. The stromatal tissue between perithecia
is largely dark brown but with individual ostioles encased
with reduced carbonaceous tissue (Fig. 20). This feature also
occurs in B. arima, B. cinereolilacina, B. marginata, B. repanda,
and B. simplicior (Ju et al. 1998).
Miller (1961) designated specimen no. 30256 at PREM as
the (holo)type of Hypoxylon doidgeae, which, unfortunately,
was not located by us. There is a specimen bearing the same
number at BPI and was labeled as type by Miller; we
accordingly select this as the lectotype. This fungus is known
only from South Africa.
Biscogniauxia formosana var. kentingensis Y.-M. Ju &
J. D. Rogers, var. nov.
(Figs 29–34, 49)
Etym.: After the locality where the collection was made.
Biscogniauxia formosana Y.-M. Ju & J. D. Rogers, sp. nov.
(Figs 23–28, 48)
A varietate typica differt in ascosporis angustioribus dilutioribus
17–23n5i7–8 µm et in conidiis 6–12i3–4n5 µm.
Typus: Taiwan: Ping-tung Co.: Hen-chuan, Ken-ting, on bark, 16
Etym.: After Taiwan.
Sep. 2000, Y.-M. Ju & H.-M. Hsieh 89091613 [cultured]
(HAST – holotypus; WSP 70166 – isotypus).
A Biscogniauxia capnode var. rumpenti differt in ascosporis valde
inequilateralis et in ostiolis solum umbilicatis.
Typus: Taiwan: Taipei City: Nankang Dist., on bark, 22 Mar.
2000, Y.-M. Ju 89032201 [cultured] (HAST – holotypus; WSP
70165 – isotypus).
Var. kentingensis differs from var. formosana in the narrower
paler ascospores 17–23n5i7–8 µm, and in the larger conidia
6–12i3–4n5 µm.
Stromata
applanate,
0n4–3 cm
longi0n4–2n5 cm
Culture and anamorph: Colonies on OMA at ca 20 m and
broadi0n6–0n7 mm thick; outer dehiscing layer dull brown, under 12 h fluorescent light much as in the typical variety.
thin; mature surface dark brown with grayish tinge, plane; Conidiogenous structure with Periconiella-like branching pattern
carbonaceous immediately beneath surface and between as defined in Ju & Rogers (1996). Conidiophores hyaline,
perithecia; tissue beneath perithecia inconspicuous. Perithecia roughened, frequently flexuous, composed of a main axis,
obovoid to tubular, 0n1–0n2 mm diami0n3–0n6 mm high, 3–4 µm broad, and sometimes one or more major branches,
with several individuals discharging through a single ostiolar branches mostly terminating with 3–5 conidiogenous cells
canal. Ostioles lower than stromatal surface, with openings arising in whorls from the main axis and\or from major
punctate. Asci 160–200 µm total lengthi9–12 µm broad, the branches. Conidiogenous cells cylindrical, hyaline, finely
spore-bearing parts 150–170 µm long, the stipes 10–30 µm roughened, 8–15i2n5–4n5 µm, bearing poroid conidial se-

Biscogniauxia
1130
cession scars on apical region, with apices becoming distorted Conidiophores pale brownish, roughened, composed of a main
from production of numerous conidia. Conidia produced axis, 3n5–5n5 µm broad, and sometimes one or more major
holoblastically in sympodial sequence, hyaline, smooth, branches, branches mostly terminating with 3–5 conidio-
subglobose to obovoid to clavate, 6–12i3–4n5 µm, with genous cells arising in whorls from the main axis and\or from
flattened base indicating former point of attachment to major branches or from additional subordinate branches;
conidiogenous cell.
additional branches infrequently produced. Conidiogenous cells
Notes: The differences between the teleomorphs and cultures cylindrical, hyaline to yellowish, smooth to finely roughened,
of var. kentingensis and var. formosana are small. Their 8–18i3n5–5 µm, bearing poroid conidial secession scars on
differences in the anamorphs are, however, substantial: while apical region, with apices becoming distorted from production
the conidia of the typical variety are small and regular in shape of numerous conidia. Conidia produced holoblastically in
(obovoid) and have a narrow length range, 4n5–5n5 µm, those sympodial sequence, hyaline, smooth, ellipsoid, 7–
of var. kentingensis are larger and quite variable in shape 9(k10n5)i4–5 µm, with flattened base indicating former
(subglobose to obovoid to clavate) and have a fairly broad point of attachment to conidiogenous cell.
length range, 6–12 µm.
Notes: Biscogniauxia laterima is unique in having beaked
ascospores with a broad germ slit. Broad ascospore germ slits
are uncommon in Biscogniauxia; they are also found in B.
philippinensis and its var. microspora, and B. citriformis.
Biscogniauxia latirima Y.-M. Ju & J. D. Rogers, sp. nov.
(Figs 35–40, 50) Nonetheless, the former two taxa have a cellular appendage
on the ascospore and the latter has none and much smaller
Etym.: Refers to the broad ascospore germ-slit.
ascospores 10n5–12n5i5n5–7n5 µm and smaller conidia 5n5–
7n5i1n5–2 µm (Ju et al. 1998). It should be noted that B.
citriformis var. macrospora has a conspicuous, narrow ascospore
germ slit (Ju et al. 1998). Moreover, B. citriformis varieties have
notably sticky hamathecia owing to abundant allophyses
(Rogers et al. 2000), a feature absent in B. laterima.
Stromata applanata, 0n3–2n5 cm latai0n6–0n8 mm crassa; stratum
externum dehiscens brunneum tenue; superficies matura fumeofusca;
textura sub superficie et inter perithecia carbonacea; textura sub
peritheciis inconspicua. Perithecia tubularia, 0n2 mm diami0n4–
0n6 mm alta, singularis vel aliquot per canalem communem ostiolarem
emittentibus. Ostiola umbilicata. Asci 155–180 µm longitudine
totai12–14 µm crassi, partibus sporiferis 140–160 µm longitudine,
stipitibus 10–20 µm longitudine, annulo apicali in liquore iodata
Melzeri cyanescente, discoideo, 4–5 µm altoi5–6 µm lato.
Ascosporae fuscae vel furcae, unicellulares, ellipsoideo-inequilaterales,
uno extremo lato, altero extremo angustatis vel rostratis, leves,
(16k)18–21n5i10–12 µm, rima germinativa longa recta, 2 µm
crassa, in latere convexa praeditae. Paraphyses inclusae.
Biscogniauxia philippinensis var microspora Y.-M. Ju &
J. D. Rogers, in Ju, Rogers, San Martın & Granmo,
Mycotaxon 66: 46 (1998).
(Figs 41–43, 47)
The teleomorph is as described in Ju et al. (1998).
Culture and anamorph: Colonies on OMA at ca 20 m and
under 12 h fluorescent light reaching the edge of 9 cm Petri
dish in 1n5 wk, whitish, felty, azonate, with diffuse margins,
becoming cottony and overlain with Buff (45) to Honey (64)
patches. Reverse Dull Green (70). Sporulating regions near
inoculation point, whitish. Conidiogenous structure with Peri-
coniella-like branching pattern as defined in Ju & Rogers
(1996). Conidiophores yellowish to brownish, coarsely
roughened, composed of a main axis, 6–7n5 µm broad, and
sometimes one or more major branches, branches mostly
terminating with 3–5 conidiogenous cells arising in whorls
from the main axis and\or from major branches. Conidiogenous
cells cylindrical, yellowish to brownish, finely roughened,
7–15i3–4n5 µm, bearing poroid conidial secession scars on
apical region, with apices becoming distorted from production
of numerous conidia. Conidia produced holoblastically in sym-
podial sequence, hyaline, smooth, obovoid, 5n5–7i3–4 µm,
with flattened base indicating former point of attachment
to conidiogenous cell.
Typus: Taiwan: I-lan Co.: Fu-shan, on bark, 11 Oct. 2000, Y.-M.
Ju 89101101 [cultured] (HAST – holotypus; WSP 70167 – isotypus).
Stromata applanate, 0n3–2n5 cm diami0n6–0n8 mm thick;
outer dehiscing layer brown, thin; mature surface dark grayish
brown, plane; carbonaceous immediately beneath surface and
between perithecia; tissue beneath perithecia inconspicuous.
Perithecia tubular, 0n2 mm diami0n4–0n6 mm high, with one
or several individuals discharging through a single ostiolar
canal. Ostioles lower than stromatal surface, with openings
punctate. Asci 155–180 µm total lengthi12–14 µm broad,
the spore-bearing parts 140–160 µm long, the stipes 10–20 µm
long, with apical ring bluing in Melzer’s iodine reagent,
discoid, 4–5 µm highi5–6 µm broad. Ascospores dark brown
to blackish brown, unicellular, ellipsoid-inequilateral, with one
end broadly rounded and the other narrowly rounded and
often beaked, smooth, (16k)18–21n5i10–12 µm, with
straight, 2-µm-broad germ slit slightly less than spore-length
on more convex side.
Culture and anamorph: Colonies on OMA at ca 20 m and
under 12 h fluorescent light reaching the edge of 9 cm Petri
dish in 2 wk, whitish, felty, azonate, with diffuse margins,
becoming cottony at places and Isabelline (65) to Fawn (87).
Reverse Citrine (13) to Greenish Olivaceous (90). Sporulating
regions scattered over entire surface of colony, whitish to
light gray. Conidiogenous structure with Periconiella-like
Notes: The Taiwan collection is much like the holotype
material from Guyana. The anamorph of the typical variety of
B. philippinensis differs primarily in having larger conidia
6n5–9i3n5–4 µm and in having a less robust conidiophore
main axis (Ju et al. 1998).
Specimen examined: Taiwan: Ping-tung Co.: Hen-chuan, Shei-tin
Park, on bark, 11 Apr. 2000, Y.-M. Ju & H.-M. Hsieh 89041101
branching pattern as defined in Ju & Rogers (1996). [cultured] (HAST; WSP 70168).

Y.-M. Ju and J. D. Rogers
1131
Key to taxa of Biscogniauxia
This key is modified from Ju et al. (1998) with addition of the new Biscogniauxia taxa described in the present study and those from Whalley
et al. (2000) and Rogers et al. (2000). Note that ascospore size ranges given for taxa with appendaged ascospores do not include the
appendage.
1
Mature ascospores bearing a cellular appendage or with one flattened end indicating appendage loss .
.
.
.
.
.
.
.
.
.
2
18
Mature ascospores lacking a cellular appendage and\or a conspicuously flattened end
[Note: B. fuscella bears a minute appendage on some mature ascospores.]
.
.
2(1) Ascospores ornamented with striations .
Ascospores not ornamented
.
.
.
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.
3
4
.
.
3(2) Ascospores (22k)25–28(k30)i(8k)9n5–10n5(k12) µm
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
. weldenii
. weldenii var. microspora
Ascospores (17k)18–21i(7k)8–10(k11) µm
.
.
4(2) Ascospores apparently lacking a germ slit
Ascospores with a germ slit
.
.
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5
6
.
.
5(4) Mature stromatal surface light orange brown; ascospores pale brown to light brown, 6–7n5i3n5–4 µm
Mature stromatal surface dull ochraceous brown; ascospores light brown to brown, 8–11n5i4–5n5 µm
.
.
.
.
. waitpela
. doidgeae
6(4) Ostiolar openings coarsely papillate
Ostiolar openings slightly papillate or umbilicate
.
.
.
.
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.
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.
.
.
7
8
7(6) Ascospores (12k)13–14n5(k15)i7–8 µm
Ascospores 22–33i12n5–15n5 µm
.
.
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.
.
.
.nothofagi
. pithodes
.
8(6) Ascospores mostly remaining hyaline when mature, 10–13i7–9(k10) µm
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
anceps
9
Ascospores mostly with larger cell darkened when mature
.
.
.
.
9(8) Ascospores with a faint, broad germ slit on the more convex side .
.
.
.
.
.
.
Ascospores with a conspicuous, narrow germ slit either on the more convex or on the less convex side
.
.
.
.
.
.
.
.
10
11
10(9) Ascospores 22–27i11n5–13 µm .
Ascospores 13n5–19i8–10 µm
.
.
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.
.
. philippinensis
. philippinensis var. microspora
.
11(9) Ascospores mostly greater than 15 µm long .
Ascospores mostly less than 15 µm long
.
.
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.
.
12
13
.
12(11) Ascospores (14k)16–21i5–6n5 µm; known only from New Zealand
Ascospores 15–22i6–8 µm; known only from the Americas .
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
. zelandica
divergens
.
13(11) Mature stromatal surface dark grey; ascospores 12–14n5i6–7 µm; known only from Hawaiian Islands
Mature stromatal surface other than grey
.
.
.
.
. plumbea
. 14
.
.
.
.
.
.
.
.
.
.
.
.
14(13) Hamathecial tissue composed of paraphyses intermixed with copious, sticky allophyses; ascospores (9k)10–12i4n5–6 µm
viscosicentra
15
Hamathecial tissue mostly composed of paraphyses
.
.
.
.
.
.
.
.
.
.
.
.
.
15(14) Mature stromatal surface brown to dark brown; ascospores 12–16i(5k)5n5–7n5 µm, with dark cell narrowly rounded or
pinched at end .
.
.
Mature stromatal surface black
.
.
.
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.
.
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.
.
grenadensis
16
.
.
16(15) Ascospores (9k)10–14i5–7 µm, with germ-slit spore-length or slightly less than spore-length; appendage 2–2n5 µm long
uniapiculata
17
Ascospores with germ-slit less than spore-length
.
.
.
.
.
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.
.
.
17(16) Ascospores 10–12i6–7n5(k8n5) µm; appendage 2n5–4 µm long
.
.
.
.
.
.
.
.
.
.
.
.
uniapiculata var. indica
Ascospores 13–15n5i7n5–10 µm; appendage 2–3 µm long
.
uniapiculata var. macrospora
18(1) Ascospores ornamented with longitudinal ridges or shallow reticulations .
.
.
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.
.
19
20
Ascospores smooth
.
.
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.
.
19(18) Ascospores 26–30i7n5–10 µm, ornamented with longitudinal ridges
.
.
.
.
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.
.
.
.
.
.
.
.
.
.
.
. reticulospora
sinuosa
Ascospores 27–34i14–15n5 µm, ornamented with reticulations
.
20(18) Ascospores 7n5–9i3–4 µm, apparently lacking a germ slit
.
.
.
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.
.
.
.
.
.
.
arima
.
Ascospores larger, with a germ slit
.
.
.
.
21
21(20) Germ-slit on both sides of the ascospore .
Germ-slit only on one side of the ascospore .
.
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22
29
22(21) Ascospores laterally compressed .
Ascospores not laterally compressed
.
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.
23
24
23(22) Ostiolar openings punctate; ascospores 13–14n5i8n5–11n5i7–7n5 µm
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
schweinitzii
.
Ostiolar openings coarsely papillate; ascospores 17–22i11–14i9–10 µm
querna
24(22) Stromatal tissue between perithecia entirely composed of black, carbonaceous tissue .
.
.
.
.
.
.
.
25
28
Stromatal tissue between perithecia composed of greyish brown to dark brown, woody tissue and black, carbonaceous tissue
encasing individual ostioles reduced .
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.

Biscogniauxia
1132
25(24) Ascospores subglobose, 7n5–8n5i6–7 µm
.
Ascospores ellipsoid, greater than 9 µm long .
.
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.
.
. nawawii
.
.
26
. ambiens
27
.albosticta
26(25) Mature stromatal surface dark brown with olivaceous tinge; ascospores 10–13i4–5(k6) µm
.
.
.
.
.
.
.
.
Mature stromatal surface not tinged olivaceous; ascospores greater than 13 µm long
.
.
.
27(26) Ascospores 15n5–19n5i6n5–8n5 µm
Ascospores 13–16i5–7 µm
.
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.
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.
. albosticta var. orientalis
28(24) Stromata applanate; ascospores 12–16(k17n5)i(4n5–)5n5–8 µm
Stromata raised-discoid; ascospores (8k)9–14i(4k)4n5–6 µm
.
.
.
.
.
.
.
.
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.
.
.
.
.
.
.
. cinereolilacina
.
. repanda
29(21) Stromata raised-discoid or short-cylindrical; ascospores 13–16n5i9n5–13n5 µm, with a sigmoid germ-slit
.
.
.
.
marginata
30
Stromata various; ascospores with a straight germ-slit
.
.
.
.
.
.
.
.
.
.
.
30(29) Stromata raised-discoid, usually with conspicuously raised margins, with circular outlines
Stromata applanate, lacking raised margins, usually with irregular outlines
.
.
.
.
.
.
.
.
.
.
.
.
.
.
31
36
.
.
31(30) Ostiolar openings papillate to coarsely papillate
Ostiolar openings punctate .
.
.
.
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.
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.
32
34
.
.
.
32(31) Ascospores light brown to brown, 10–13i4n5–5n5(k6) µm .
.
.
.
Ascospores dark brown to blackish brown, greater than 13 µm long and 6 µm broad
.
.
.
.
.
.
.
.
.
.
.
. africana
33
.
.
33(32) Perithecia 0n3–0n5 mm diam; ascospores 15n5–21i10n5–13n5 µm; known only from Europe
Perithecia 0n1–0n3 mm diam; ascospores 14–16n5i9n5–11n5 µm; known only from Australia
.
.
.
.
.
.
.
.
. dennisii
baileyi
.
34(31) Stromatal tissue between perithecia carbonaceous; ascospores (9n5k)10–12(k13)i5–6 µm
.
.
.
.
.
.
. kalchbrenneri
35
Stromatal tissue between perithecia essentially dark brown, woody .
.
.
.
.
.
.
.
35(34) Ostioles discretely encased by carbonaceous stromatal tissue; ascospores 13–17n5i5–6n5 µm; on Rhamnus in Europe simplicior
Ostioles not encased by carbonaceous stromatal tissue; ascospores 10n5–14n5i5–7 µm; on Prunus in Europe
[Note: This fungus was referred to as ‘B. pruni’, a provisional name, in Ju et al. (1998).]
granmo
36(30) Mature stromatal surface white
.
.
Mature stromatal surface other than white
.
.
.
.
.
.
.
.
.
.
.
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.
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.
.
.
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.
.
37
39
37(36) Ascospores (20k)23–30i11n5–14n5 µm
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
atropunctata
38
Ascospores smaller
.
.
.
.
.
.
38(37) Ascospores 16–22i7n5–11 µm
Ascospores 12–15n5i7–8 µm
.
.
.
.
.
.
.
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.
.
.
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.
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.
.
.
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.
.
.
atropunctata var. intermedia
atropunctata var. maritima
39(36) Ascospores strongly inequilateral .
Ascospores more or less equilateral
.
.
.
.
.
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.
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.
.
40
44
40(39) Ascospores greater than 18 µm long
Ascospores less than 18 µm long .
.
.
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.
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.
.
.
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.
.
41
43
41(40) Ascospores (16k)18–21n5i10–12 µm, with one end broadly rounded and the other narrowly rounded and often beaked, with
a 2-µm-broad germ-slit on more convex side
.
.
.
.
.
.
.
.
.
.
Ascospores with both ends narrowly rounded and frequently pinched, with a narrow germ-slit on less convex side
.
.
.
.
latirima
.
42
42(41) Ascospores (18k)19–24(k25)i8n5–10 µm; conidia 4n5–5n5i2n5–3n2 µm
.
.
.
.
.
.
.
.
.
formosana var. kentingensis
.
.
formosana
Ascospores 17–23n5i7–8 µm; conidia 6–12i3–4n5 µm
.
.
.
43(40) Ascospores 10n5–12n5i5n5–7n5 µm
Ascospores (12–)13n5–17n5i6n5–9 µm
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
citriformis
citriformis var. macrospora
44(39) Ascospores 8n5–10n5i5–5n5 µm, surrounded by a hyaline sheath
Ascospores lacking a hyaline sheath
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
. mucigera
45
.
.
.
.
.
.
.
45(44) Outer dehiscing layer white; ascospores 13–16n5i6n5–8 µm, sometimes bearing a minute cellular appendage
Outer dehiscing layer fawn, brown, or dark brown; ascospores not bearing a cellular appendage
.
.
.
.
fuscella
.
.
.
46
46(45) Stromatal tissue between perithecia composed entirely of gray, grayish brown, or dark brown, woody tissue
Stromatal tissue between perithecia composed entirely of carbonaceous tissue
.
.
.
.
.
.
47
48
.
.
.
.
.
.
47(46) Ascospores 17n5–22(k25)i(8–)9–11(k12) µm
Ascospores 10–13(k14)i(6n5k)7n5–8n5 µm
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
. bartholomaei
nummularia
.
48(46) Ascospores 8–11n5i4n5–6 µm, with a much less than spore-length germ-slit; a number of perithecia usually sharing one ostiole
.
.
.
.
.
.
.
.
.
.
.
.
.
Ascospores with a spore-length germ-slit; perithecia seldom or frequently sharing a common ostiole .
.
.
.
.
.
.
.
.
communapertura
49
.
.
.
49(48) Ostiolar openings coarsely papillate
.
.
Ostiolar openings slightly papillate or punctate
.
.
.
.
.
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.
.
.
50
52
50(49) Ascospores 20–26i11n5–13 µm .
Ascospores smaller
.
.
.
.
.
.
.
.
.
.
.
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.
.
.
.
.
.
. mediterranea var. macrospora
.
.
.
.
.
.
.
.
.
51
51(50) Ascospores 15n5–21i(6n5k)7–10 µm .
Ascospores 12–17i5n5–7n5 µm .
.
.
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.
.
.
.
.
.
.
.
.
.
. mediterranea
. mediterranea var. microspora
.

Y.-M. Ju and J. D. Rogers
1133
52(49) Perithecia very narrow, ca 0n05–0n1 mm, with numerous individuals arranged in rosettes and sharing a single ostiolar canal . 53
Perithecia more than 0n2 mm diam, usually not arranged in rosettes and usually with individual ostioles
.
.
.
.
54
53(52) Ascospores 11–14i5n5–7n5 µm .
Ascospores 20–26i10–14(k16) µm
.
.
.
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.
.
.
.
.
.
. cylindrispora
plana
54(52) Ascospores lemon-shaped, 11n5–14i6–7n5 µm
Ascospores other than lemon-shaped
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
capnodes var. limoniispora
55
.
.
.
.
.
.
.
55(54) Ascospores (14k)15–22n5(k24)i8–11(k12) µm
Ascospores smaller
.
.
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.
capnodes var. rumpens
.
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.
.
56
56(55) Ascospores brown to dark brown, 8n5–15i5–7n5 µm
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
. capnodes
Ascospores light brown, 9n5–13n5i4n5–5 µm
.
. capnodes var. theissenii
Ju, Y.-M., Rogers, J. D., San Martın Gonzalez, F. & Granmo, A. (1998) The
genus Biscogniauxia. Mycotaxon 66: 1–98.
ACKNOWLEDGEMENTS
This study was supported by National Science Council of R.O.C. Grant
NSC89-2311-B-001-126 to Y.-M.J. and by US National Science Foundation
Grant DEB-9813304 to J.D.R. The curators of BPI are thanked for the loan
of type material of Hypoxylon doidgeae.
Miller, J. H. (1961) A Monograph of the World Species of Hypoxylon. University
Georgia Press, Athens.
Rayner, R. W. (1970) A Mycological Colour Chart. Commonwealth My-
cological Institute, Kew.
Rogers, J. D., San Martın, F., Ju, Y.-M. & Hansen, K. (2000) Venezuelan fungi:
Biscogniauxia viscosicentra sp. nov. and the anamorph of Camillea cyclops.
Nova Hedwigia 71: 431–437.
REFERENCES
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Corresponding Editor: D. L. Hawksworth