INTRODUCTION

Species of Nectria (Fr.) Fr. sensu lato are characterized

by bright-colored, mainly red to orange perithecia, which

are mostly formed superficially on their substrata and

solitary or aggregated into pustules. Most of the species

have two-celled, hyaline ascospores, which are smooth,

warted or striated. Unitunicate asci, with or without an

apical apparatus, and apical paraphyses place these fungi

in the order Hypocreales. Fungi of Nectria sensu lato have

been segregated from the genus and redistributed among

a number of genera mainly in the families Nectriaceae

and Bionectriaceae (Rossman et al., 1999; Samuels et

al., 2002). These two families are separated primarily by

ascomatal color, color reactions in KOH and lactic acid

(Rossman et al., 1999). Species with red or orange-red

perithecia, which change color in KOH or lactic acid are

assignable to the Nectriaceae, whereas those with orange,

pale yellow or white perithecia, which lack a color reaction

in either reagent, are assignable to the Bionectriaceae.

Nectriaceous fungi have anamorphs in several

hyphomycetous form-genera (Seifert, 1985; Samuels

and Seifert, 1987) and in most cases the form-genera

do not overlap in the two families, with Acremonium as

the only exception which can be found in taxa of both

families, such as Cosmospora vilior (Nectriaceae) and

Hydropisphaera suffulta (Bionectriaceae). Common

Botanical Studies (2007) 48: 187-203.

*

Corresponding author: E-mail: yumingju@gate.sinica.edu.

tw; Tel: +886-2-27899590 ext 314.

anamorphs of Nectriaceae are in Acremonium Link,

Actinostilbe Petch, Antipodium Piroz., Chaetopsina

Rambelli, Cylindrocarpon Wollenw., Fusarium Link,

Tubercularia Tode, and Volutella Fr., whereas those of

Bionectriaceae are in Acremonium, Clonostachys Corda,

Dendrodochium Bonord., and Stilbella Lindau (Schroers,

2001). The recognition of these two families is also

supported by molecular phylogenetic studies (e.g., Rehner

and Samuels, 1994, 1995; Ogawa et al., 1997).

Ninteen nectriaceous fungi have previously been

reported in Taiwan. Sawada (1928, 1931, 1959a, b)

recorded Nectria citri Henn., N. diversispora Petch, N.

durantae Sawada, N. elephatopodis Sawada, and N.

pterospermi Sawada, and Gibberella fujikuroi (Sawada)

Wollenw. Nectria swieteniae-mahogani C.-C. Chen was

reported to grow on Swietenia mahogani (Chen, 1965).

Nectria flammea (Tul. & C. Tul.) Dingley was found on

scale insects on Paulownia (Chen, 1975). Calonectria

theae Loos and C. crotalariae (Loos) D. K. Bell &

Sobers

caused a root rot (Chang, 1992) on Cinnamomum

osmophloeum Kanehira and a black rot on Sassafras sp.

(Chang, 1994), respectively. Nectria galligena Bres., now

accepted as Neonectria galligena (Bres.) Rossman &

Samuels (Rossman et al., 1999), caused an apple canker

in Li-shan (Sun and Yu, 1975). Nectria haematococca

Berk. & Broome [¡Ý Haematonectria haematococca (Berk.

& Broome) Samuels & Rossman], with the anamorph

Fusarium solani (Mart.) Sacc., caused a collar and root rot

on passion fruit Passiflora edulis Sims (Lin and Chang,

1982). Chen and Hsieh (1996) recorded Thyronectria

pseudotrichia (Schwein.) Seeler a common encounter

mICROBIOlOgy

Nectriaceous fungi collected from forests in Taiwan

Jia-Rong GUU

1

, Yu-Ming JU

2,

*, and Huan-Ju HSIEH

1

1

Department of Plant Pathology and Microbiology, National Taiwan University, Taipei 10617, Taiwan

2

Institute of Plant and Microbial Biology, Academia Sinica, Nankang, Taipei 11529, Taiwan

(Received June 9, 2006; Accetped November 2, 2006)

ABSTRACT.

Nineteen fungi of the family Nectriaceae collected from forests in Taiwan are reported,

including Albonectria albida, A. rigidiuscula, Cosmospora cf. consors, C. diminuta, C. cf. glabra, C. joca, C.

triqua, C. vilior, Haematonectria haematococca, Lanatonectria flavolanata, L. flocculenta, Nectria

balsamea,

N. pseudotrichia, Neonectria coronata, Neo. discophora, Neo. jungneri, Neo. lucida, Neo. rugulosa, a nd

Ophionectria trichospora. Most of them were found on recently dead broad-leaved trees. Only A. rigidiuscula,

H. haematococca, and N. pseudotrichia have previously been reported in Taiwan; others represent new

records. Nectria albida is newly combined into Albonectria as A. albida on the basis of its ascomatal color

and Fusarium anamorph. A dichotomous key to these species is also given.

Keywords: Albonectria; Bionectriaceae; Cosmospora; Haematonectia; Hypocreales; Neonectria; Nectria;

Nectriaceae; Neonectria; Ophionectria; Systematics.

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Botanical Studies, Vol. 48, 2007

on dead branches. Wang (2000) described a new species

Nectria caricae C.-L. Wang from dead papaya stems.

Sun and Huang (1996) reported two Nectria species, N.

haematococca and N. rigidiuscula Berk. & Broome [¡Ý

Albonectria rigidiuscula (Berk. & Broome) Rossman &

Samuels] and six Gibberella species. Unlike most of the

specimens included in our study, which were collected

from recently dead, broad-leaved trees in forests, those

previously recorded species were mostly reported from

crops. This apparently accounts for the fact that the

majority of the species that we deal with herein have not

been recorded in Taiwan before.

In the present study, we report nineteen species of the

Nectriaceae found in Taiwan, including two Albonectria

species, seven Cosmospora species, one Haematonectria

species, two Lanatonectria species, one Nectria species,

five Neonectria species, and one Ophionectria species. In

addition, efforts were made to show teleomorph-anamorph

connections of these taxa via culturing. Anamorphs of

these nineteen nectriaceous fungi are distributed among

eight form-genera, with Cosmospora connected to

Acremonium, Chaetopsina and Volutella, Lanatonectria to

Actinostilbe, Ophionectria to Antipodium, Neonectria to

Cylindrocarpon, Haematonectria to Fusarium, and Nectria

to Tubercularia and Zythiostroma

Hohn. ex Falck.

mATERIAlS AND mETHODS

Perithecia were immersed in 10% KOH and lactophenol

to observe color changes. Perithecia becoming purple

or deeply red in 10% KOH but pale red or yellowish in

lactophenol were recorded as positive color changes.

Perithecial wall layers were described based on

longitudinal sections made by a freezing microtome

(Yamatokohki FX-801). Sections of perithecia were

approximately 15 £gm thick. Cultures were initiated from

single or multiple ascospores on Difco potato dextrose

agar (PDA). Resulting colonies were transferred to 9-cm

plastic Petri plates containing PDA, from which the culture

descriptions were made. Cultures were incubated under

12-hour fluorescent light at 20¢XC. The cited specimens

were deposited in HAST (Botanical Herbarium, Academia

Sinica, Taipei). Obtained cultures were deposited in the

BCRC (the Bioresource Collection and Research Center)

in Taiwan. Descriptions of teleomorphs, cultures, and

anamorphs were prepared in brief formats due to detailed

descriptions can readily be found in recent literature.

Terminology used in descriptions of teleomorphs and

anamorphs follows Kirk et al. (2001). The numbers of

ascospores, conidia, perithecia, asci, and conidiophores

that were measured to form the size ranges in the

descriptions are 20, 20, 5, 5, and 5, respectively.

TAXONOmy

Key to the nectriaceous fungi studied

1. Perithecia not changing color in 10% KOH or lactophenol ............................................................................................... 2

1. Perithecia red, becoming purple or deeply red in 10% KOH but pale red or yellowish in lactophenol ....... 4 (Nectriaceae)

2.

Ascospores mostly two-celled; anamorph often Acremonium-like or Clonostachys .......................................................

....................................................................................................................... Bionectriaceae (not further treated herein)

2. Ascospores phragmosporous; anamorph Fusarium ............................................................ 3 (Albonectria, Nectriaceae)

3.

Perithecia white or pale yellow, with coarsely concolorous warts on the surface; ascospores 3-septate, 22.5-27 ¡Ñ 6.5-8.5

£gm ..................................................................................................................................................................A. rigidiuscula

3. Perithecia light yellow, smooth; ascospores 3-septate, 32-39 ¡Ñ 7-8 £gm

.................................................................A. albida

4. Ascospores muriform ......................................................................................................................................5 (Nectria)

4. Ascospores didymosporous or phragmosporous ............................................................................................................ 6

5.

Perithecia red, covered with white scales; asci filled with ascoconidia; ascospores (18-)20-25.5(-30) ¡Ñ 4.5-)5.5-6(-6.5)

£gm ..................................................................................................................................................................... N. balsamea

5.

Perithecia red to dark red, covered with concolorous scales; asci not filled with ascoconidia; ascospores (15-)19-25(-30)

¡Ñ 9-11(-13) £gm ............................................................................................................................................N. pseudotrichia

6. Ascospores scolecosporous, 13- to 20-septate, 190-210 ¡Ñ 9-11 £gm ....................................... Ophionectria trichospora

6. Ascosporous didymosporous .......................................................................................................................................... 7

7. Perithecia covered with echinulate, golden-yellow hairs ......................................................................... 8 (Lanatonectria)

7. Perithecia lacking hairs on the surface

................................................................................................................................ 9

8. Ascospores 11-13 ¡Ñ 3-4 £gm; anamorph sporodochial .................................................................................L. flocculenta

8. Ascospores 15-20 ¡Ñ 5-5.5 £gm; anamorph synnematous ............................................................................. L. flavolanata

9. Perithecia greater than 250 £gm diam; anamorph Cylindrocarpon

............................................................... 10 (Neonectria)

9.

Perithecia less than 250 £gm diam, smooth or coarsely warted; anamorph Fusarium, Volutella, Chaetopsina,

Acremonium-like, often found on natural substrata .......................................................................................................... 14

GUU et al. ¡X Nectriaceae in Taiwan

189

Albonectria albida (Rossman) J.-R. Guu & Y.-M. Ju,

comb. nov.

Figures 1B, 2F-I

Basionym. Nectria albida Rossman, Mycol. Pap. 150: 79.

1983.

Anamorph. Fusarium sp.

Stromata well-developed, pulvinate, white or buff-

colored. Perithecia aggregated, superficial, light

yellow, not changing color in 10% KOH or lactophenol,

obpyriform, 200-260 ¡Ñ 160-180 £gm, smooth, not collapsed

when dry; perithecial wall 15-20 £gm thick, two-layered;

ostioles obtuse. Asci clavate, 55-87 ¡Ñ 14-16 £gm, lacking

an apical ring. Ascospores hyaline, fusiform, 3-septate,

32-39 ¡Ñ 7-8 £gm, smooth.

Cultures and anamorph. Colonies attaining 2.5 cm diam

in 14 days at 20¢XC, with scanty aerial hyphae, diffusing

luteous to light yellow pigments into the medium.

Conidiophores producing macroconidia only, aggregated

into sporodochia, irregularly branched, bearing a phialide

at each terminus, with the phialides cylindrical, 10-22 ¡Ñ

4-6 £gm. Macroconidia hyaline, fusiform, slightly curved,

with a pointed tip and foot cell, 3- to 7-septate, 72-90 ¡Ñ

6-7 £gm.

Specimens examined. Nantou County, Luku, Shitou, on

bark, with Haematonectria haematococca and Valetoniella

sp., 19 Apr 2003, Ju, Y.-M. 92041906 (cultured) &

92041907 (cultured). Taipei County, Hsintien, Shih-tzu-

tou-shan, on bark, 11 Dec 2004, Guu, J.-R. 93121106

(cultured).

Notes. The perithecia of Albonectria do not change

color in KOH or lactophenol and are therefore atypical for

the Nectriaceae. Nonetheless, the Fusarium anamorphs

and molecular phylogenies based on sequences of the

28s rDNA suggest the disposition of Albonectria in the

Nectriaceae rather than Bionectriaceae (Guadet et al.,

1989; O¡¦Donnell, 1993; Rehner and Samuels, 1995).

Rossman (1983) described the anamorph of A. albida and

assigned it to Fusarium. The anamorph that we obtained

from the Taiwan specimens fits well her description. We

feel justified to combine the epithet of Nectria albida

with Albonectria due to its possession of light yellow

perithecia and a Fusarium anamorph. Also see notes on A.

rigidiuscula.

Albonectria rigidiuscula (Berk. & Broome) Rossman &

Samuels in Rossman, Samuels, Rogerson & Lowen,

Stud. Mycol. 42: 105. 1999.

Figures 1D, 2A-E

Anamorph. Fusarium decemcellulare Brick, Jahresber.

Vereinigung Angew. Bot. 6: 277. 1908.

Stromata flattened to pulvinate, sometimes

inconspicuous. Perithecia solitary to aggregated in groups

of up to 30, white or pale yellow, not changing color in

10% KOH or lactophenol, globose to subglobose, 225-350

¡Ñ 180-325 £gm, uncollapsed when dry, ornamented

with large, concolorous, coarse warts up to 50 £gm

high; perithecial wall 25-80 £gm thick, three-layered;

ostioles slightly papillate, darker than perithecial body.

Asci clavate, 70-90 ¡Ñ 10-14 £gm, lacking an apical ring.

Ascospores hyaline, oblong-ellipsoidal, 3-septate, 22.5-27

¡Ñ 6.5-8.5 £gm, faintly striated.

10. Ostioles encircled with a crown ..............................................................................................................Neo. coronata

10. Ostioles not encircled with a crown ...........................................................................................................................11

11. Perithecia with apapillate ostioles, tending to be orange-red, covered with red warts .................................Neo. rugulosa

11. Perithecia with papillate ostioles, smooth, red to blood-red ........................................................................................... 12

12. Ostiloes slightly papillate .................................................................................................................... Neo. discophora

12. Ostioles coarsely papillate

......................................................................................................................................... 13

13.

Ascospores echinulate, 13-15 ¡Ñ 5-6 £gm

............................................................................................................ Neo. lucida

13. Ascospores striated, 20-21.5 ¡Ñ 8-9.5 £gm ...................................................................................................... Neo. jungneri

14. Perithecia roughened or scaly, cupulate when dry ...................................................... Haematonectria haematococca

14. Perithecia smooth, laterally pinched or not deformed when dry........................................................15 (Cosmospora)

15. Ascospores longer than 20 £gm ........................................................................................................................................ 16

15. Ascospores shorter than 20 £gm ....................................................................................................................................... 17

16. Ascospores (28-)34-48 ¡Ñ (6-)7-8(-9) £gm, smooth..................................................................................... C. cf. glabra

16. Ascospores 22-40 ¡Ñ 7.5-10.5 £gm, striate ....................................................................................................C. diminuta

17. Ascospores less than 3 £gm broad, 7.5-9(-9.5) ¡Ñ 2-2.5 £gm; anamorph Volutella sp. ..................................... C. cf. consors

17. Ascospores more than 3 £gm broad; anamorph Acremonium sp. ..................................................................................... 18

18. Ascospores warted ..................................................................................................................................................... 19

18. Ascospores granulated ..................................................................................................................................... C. triqua

19. Ascospores constricted at the septum, 5-8.5 ¡Ñ 3-4 £gm .......................................................................................... C. vilior

19. Ascospores not constricted at the septum, 9.5-13 ¡Ñ (4.5-)5-5.5(-6) £gm .................................................................. C. joca

190

Botanical Studies, Vol. 48, 2007

Figure 1. Macroscopical details of ascomata of selected nectriaceous fungi on their natural substrata. A, Nectria pseudotrichia; B,

Albonectria albida; C, Haematonectria haematococca; D, Albonectria rigidiuscula; E, Neonectria discophora; F, Neonectria lucida;

G, Neonectria rugulosa; H, Neonectria jungneri; I, Neonectria coronata; J, Lanatonectria flocculenta; K, Lanatonectria flavolanata;

L, Ophionectria trichospora; M, Nectria balsamea; N, Cosmospora cf. consors; O, Cosmospora cf. glabra; P, Cosmospora joca; Q,

Cosmospora vilior. Scale bars: A, D = 500 £gm; E, L, M = 400 £gm; C, H, K, T = 300 £gm; B, F, G, J, N, O, Q = 200 £gm; P = 100 £gm; I

= 100 £gm.

GUU et al. ¡X Nectriaceae in Taiwan

191

Cultures and anamorph. Colonies attaining 7 cm

diam in 14 days at 20¢XC, with short aerial hyphae,

diffusing a rosy pigment into the medium. Conidiophores

producing macroconidia and microconidia; those bearing

macroconidia densely aggregated into slimy, pale luteous

sporodochia, arising from the aerial hyphae and agar

surface, scarcely branched, producing a phialide at each

terminus, with the phialides cylindrical, up to 100 £gm long;

those bearing microconidia simple, producing verticillate

phialides, 25-37 ¡Ñ 4-5 £gm. Macroconidia slightly curved,

cylindrical to fusiform, 7-septate, hyaline, smooth, 55-120

¡Ñ 6-9 £gm. Microconidia formed in chains, ovoid, aseptate

or, occasionally, 1-septate, hyaline, smooth, 5-8 ¡Ñ 2-4 £gm.

Chlamydospores absent.

Specimens examined. Taipei County, Sunshai, Mann-

yue-yuan, on bark, 27 Sep 2000, Ju, Y.-M. and Hsieh,

H.-M. 89092703 (cultured). Taipei City, Nankang, Hu-

shih park, on bark, 6 Apr 2003, Ju, Y.-M. and Hsieh, H.-M.

92040603 (cultured). Taipei City, Nankang, Hu-shih

park, on bark, 31 Jul 2003, Guu, J.-R. 92073104. Taipei

City, Songshan, Hu-shan trail, on bark, 4 Aug 2003, Guu,

J.-R. 92080404. Taipei County, Hsintien, Jih-tan-shan,

on bark, with Bionectria ochroleuca, 9 Oct 2003, Guu,

J.-R. 92100905. Taipei County, Hsintien, Yinhodong, on

bark, mixed with Bionectria ochroleuca, Cosmospora sp.,

and Haematonectria haematococca, 16 Oct 2003, Guu,

J.-R. 92101603. Taipei County, Pingshi, Chung-yang-

chien-shan, on bark, 20 Nov 2003, Guu, J.-R. 92112003

(cultured). Taipei County, Sungi, Datunshi historical trail,

on bark, 6 Dec 2003, Guu, J.-R 92120603 (cultured).

Taipei County, Jingtung, Shu-lung-chien, on bark,

5 Jan

2004, Guu, J.-R 93010503 (cultured). Taipei County,

Hsintien, Shih-tzu-tou-shan, on bark, 2 May 2004, Guu,

J.-R. 93050203. Pingtung County, Chaochou, on bark of

Leucaena leucocephala, 27 Aug 2004, Shih, H.-H. and

Chao, T.-H. 93082702. Taipei County, Jingtung, Jingtung

historical trail, on bark, 4 Sep 2004, Kuo, H.-P. 93090406.

Taipei City, Muja, on bark, 25 Sep 2004, Shih, H.-H.

93092507. Taipei County, Pingshi, Chou-tou-shan, on

bark of Ficus septica, 31 Oct 2004, Guu, J.-R. 93103107.

Taipei County, Hsintien, Yinhodong, on bark, 25 Nov

2004, Shih, H.-H. 93112501 (cultured). Taipei County,

Hsintien, Shih-tzu-tou-shan, on bark, 11 Dec 2004, Guu,

J.-R. 93121103. Kaohsiung County, Maolinn, on bark,

mixed with Haematonectria haematococca, 14 Sep 2005,

Guu, J.-R. 94091405.

Notes. Although both four- and eight-spored asci have

been found in this species (Rossman, 1983), only four-

spored asci were observed among the Taiwan collections.

Some of the cultures that we obtained produced the

teleomorph. The teleomorph, cultures, and anamorph of

the Taiwan material match those described by Rossman

(1983) and Rossman et al. (1999). Albonectria rigidiuscula

differs from A. albida in having warted perithecia and

smaller, oblong-ellipsoidal ascospores ornamented with

faint striations.

Cosmospora cf. consors (Ellis & Everh.) Rossman &

Samuels in Rossman, Samuels, Rogerson & Lowen,

Stud. Mycol. 42: 119. 1999.

Figure 1N, 2Q

Anamorph. Volutella sp.

Stromata lacking. Perithecia mostly solitary, superficial,

rarely aggregated, red, changing color to dark red in 10%

KOH and yellow in lactophenol, obpyriform, 200-270 ¡Ñ

150-200 £gm, smooth; perithecial wall 15-20 £gm thick,

two-layered; ostioles papillate. Asci narrowly clavate,

45-60 ¡Ñ 5-8 £gm, with an apical ring. Ascospores hyaline,

fusiform, 1-septate, 7.5-9(-9.5) ¡Ñ 2-2.5 £gm, smooth.

Anamorph not obtained in culture but found on natural

substrate, sporodochial, produced among the perithecia.

Sporodochia stipitate, white to pale yellow, bearing

hemispherical conidial masses. Setae arising from the

sporodochial base, cylindrical, ca 240 £gm long, septate,

tapering toward the tip. Conidiophores aggregated,

branched, bearing 1, 3 or 4 cylindrical phialides. Conidia

hyaline, ellipsoidal, aseptate, (4.5-)5.5-6.5 ¡Ñ 1-1.5 £gm.

Specimens examined. Taipei City, Muja, campus of

National Chengchi Univ., on bark, mixed with Albonectria

rigidiuscula and Haematonectria haematococca, 2 5

Sep 2004, Shih, S.-H. 93092505. Taipei County, Sungi,

Datunshi historical trail, on bamboo culm, 10 Jul 2005,

Guu, J.-R. 94071001.

Notes. Our specimens have ascospores smaller than

those reported in Samuels (1977), i.e., 7.5-9(-9.5) ¡Ñ 2-2.5

£gm vs. (9-)10-11(-13) ¡Ñ (2.5-)3-4 £gm. While typical

C. consors was reported to be constantly associated

with decaying herbaceous debris (Samuels, 1977), our

specimens were collected from the bark of a woody

plant and bamboo culm. The anamorph of C. consors is

Volutella minima Hohn. (Samuels, 1977). Unfortunately,

we were unable to obtain cultures from our specimens.

However, the anamorph produced on the natural substrate

is in general as described by Samuels (1977) except for

the conidia being slightly narrower, i.e., 1-1.5 £gm vs.

1.5-2.5(-3) £gm. It is likely that our specimens represent an

undescribed taxon of Cosmospora, but we prefer to defer

the decision of making a new taxon until further culturable

material is available. The present fungus differs from the

other species of the genus found in Taiwan in

having a

purported Volutella anamorph.

Cosmospora diminuta (Berk.) Rossman & Samuels in

Rossman, Samuels, Rogerson & Lowen, Stud. Mycol.

42: 120. 1999.

Figure 2P

Anamorph. Unknown.

Stromata lacking. Perithecia solitary, superficial, red,

changing color to dark red in 10% KOH and yellow in

lactophenol, obpyriform, 150-190 ¡Ñ 160-210 £gm, smooth;

perithecial wall 15-20 £gm thick, two-layered; ostioles

obtuse. Asci clavate, 55-70 ¡Ñ 17-22 £gm, lacking an apical

ring. Ascospores hyaline, of two types; one type fusiform,

1-septate, 30-40 ¡Ñ 8.5-10.5 £gm, striate; the other type

oblong-ovoid, 3-septate, 22-28.5 ¡Ñ 7.5-9 £gm, granulated.

192

Botanical Studies, Vol. 48, 2007

Specimen examined. Taipei County, Hsintien, Shih-tzu-

tou-shan, on bark, 21 Mar 2004, Guu, J.-R. 93032101.

Notes. This fungus has previously been reported from

North America and Sri Lanka only (Samuels and Brayford,

1994). The Taiwan material is much as described by

Samuels and Brayford (1994) and Rossman et al. (1999).

Unfortunately, we were unable to obtain cultures from

our material. Rossman et al. (1999) observed a Volutella-

like hyphomycete associated with the type material of

Dialonectria gigaspora Cooke & Massee, which was

considered a synonym of C. diminuta by these authors,

and suspected that the hyphomycete is the anamorph of

Figure 2. A-E, Albonectria rigidiuscula. A, Median section of ascomata; B, Lateral wall; C, Macroconidium; D, Ascus; E, Ascospore.

F-I, Albonectria albida. F, Median section of ascoma; G, Lateral wall; H, Ascus; I, Ascospores. J-M, Cosmospora vilior. J, Ascus; K,

Median section of ascoma; L, Ascospores; M, Conidiophores. N-O, Cosmospora cf. glabra. N, Ascospores; O, Ascal tip with an apical

ring; P, Ascospores of Cosmospora diminuta; Q, Sporodochium of Cosmospora cf. consor. Scale bars: A = 100 £gm; B = 20 £gm; C, O

= 12 £gm; D, G = 15 £gm; E, J = 6 £gm; F = 30 £gm; H, I, L, P = 8 £gm; K = 50 £gm; M, N = 10 £gm; Q = 250 £gm.

GUU et al. ¡X Nectriaceae in Taiwan

193

C. diminuta. As with Samuels and Brayford (1994) and

Rossman et al. (1999), we also observed two types of

ascospores in the Taiwan specimen.

Cosmospora cf. glabra (Rossman) Rossman & Samuels

in Rossman, Samuels, Rogerson & Lowen, Stud.

Mycol. 42: 122. 1999.

Figures 1O, 2N, O

Anamorph. Unknown.

Stromata lacking. Perithecia solitary to aggregated,

superficial, red, not changing color in 10% KOH but

becoming yellow in lactophenol, obpyriform, 220 ¡Ñ 190

£gm, smooth; perithecial wall 10-14 £gm thick, one-layered;

ostioles obtuse. Asci clavate, 60-75 ¡Ñ 9-15 £gm, with an

apical ring. Ascopores hyaline, fusiform, 1- or 3-septate,

usually constricted at septa in 3-septate ascospores,

(28-)34.5-48 ¡Ñ (6-)7-8(-9) £gm, smooth.

Specimen examined. Taipei County, Hsintien,

Yinhodong, on bark, 16 Oct 2003, Guu, J.-R. 92101604,

mixed with A. rigidiuscula and H. haematococca.

Notes. Rossman (1983) described C. glabra possessing

3-septate ascospores in most of her cited specimens. Three

of her specimens, however, were described as having

mainly 1-septate but also some 3-septate ascospores.

Both 1-septate and 3-septate ascospores can be found in

the Taiwan specimen. Although our specimen in general

agrees with the protologue of C. glabra in Rossman

(1983), it deviates in having an ascal apical ring. Our

specimen is also similar to C. kurdica (Petr.) Rossman &

Samuels, which has a distinct ascal apical ring, but differs

in having larger ascospores. Therefore, we are uncertain in

respect to the conspecificity of the Taiwan specimen and

C. glabra. The former probably represents another taxon.

It is rather unfortunate that we were unable to germinate

ascospores from the Taiwan specimen.

Cosmospora joca (Samuels) Rossman & Samuels in

Rossman, Samuels, Rogerson & Lowen, Stud. Mycol.

42: 122. 1999.

Figure 1P

Anamorph. Acremonium sp.

Stromata inconspicuous. Perithecia solitary, superficial,

red, changing color to dark red in 10% KOH and yellow in

lactophenol, obpyriform, 200-235 ¡Ñ 175-200 £gm, smooth,

laterally collapsed when dry; perithecial wall 15-20 £gm

thick, one-layered; ostioles obtuse. Asci cylindrical, 55-70

¡Ñ 6-8 £gm, lacking an apical ring. Ascospores yellowish,

broadly ellipsoidal, 1-septate, not constricted at septum,

9.5-13 ¡Ñ (4.5-)5-5.5(-6) £gm, warted.

Cultures and anamorph. Colonies attaining 1 cm diam

in 21 days at 20¢XC, lacking aerial hyphae, usually slimy,

salmon-colored. Conidiophores scarcely produced, usually

unbranched, unseptated; phialides cylindrical, 10-25 ¡Ñ 1-3

£gm, tapering to 1 £gm near apex. Conidia hyaline, oblong

ellipsoidal, aseptate, 3-5 ¡Ñ 1-2.5 £gm, smooth.

Specimens examined. Pingtung County, Hengchun,

Kenting, on Biscogniauxia stromata, 12 Apr 2000, Ju,

Y.-M. 89041201 (cultured). Taipei County, Hsintien, Shih-

tzu-tou-shan, on depauperate Biscogniauxia stromata, 11

Dec 2004, Guu, J.-R. 93121108 (cultured).

Notes. This fungicolous species is not commonly

encountered and has only been reported once from Brazil

(Samuels, 1991). It greatly resembles C. vilior but differs

in having broader ascospores and producing a salmon-

colored pigment in culture. The Acremonium anamorph

produced by the Taiwan material much agrees with that

reported by Samuels (1991).

Cosmospora triqua (Samuels) Rossman & Samuels in

Rossman, Samuels, Rogerson & Lowen, Stud. Mycol.

42: 125. 1999.

Anamorph. Acremonium sp.

Stromata absent. Perithecia solitary, superficial, red,

changing color to dark red in 10% KOH and yellow

in lactophenol, obpyriform, 180-210 ¡Ñ 160-190 £gm,

smooth, laterally collapsed when dry; ostioles obtuse.

Asci cylindrical, 50-65 ¡Ñ 6-8 £gm, with an apical ring.

Ascospores yellowish, broadly ellipsoidal, 1-septate, not

constricted at septum, 8-11 ¡Ñ 4-5 £gm, warted.

Cultures and anamorph. Colonies attaining 1 cm diam

in 20 days at 20¢XC, lacking aerial hyphae, slimy, salmon-

colored around the inoculation points. Conidiophores

scarcely produced, mostly unbranched, unseptated;

phialides cylindrical, 12-25 ¡Ñ 1.5-2 £gm, tapering to 1 £gm

near apex. Conidia hyaline, oblong ellipsoidal, aseptate,

2.5-4 ¡Ñ 1-2 £gm, smooth.

Specimen examined. Pingtung County, Hengchun,

Kenting, on Eutypella stromata, 29 Jul 2001, Ju, Y.-M. &

Hsieh, H.-M. 90072904 (cultured).

Notes. This is another fungicolous Cosmospora species

that is uncommonly encountered. It has only been reported

from French Guiana previously (Samuels, 1991). The

Acremonium anamorph that we obtained is consistent

with that reported by Samuels (1991). Cosmospora triqua

greatly resembles C. joca but differs in having an ascal

apical ring and granulated ascospores.

Unlike the colonies

of C. joca, which diffused a salmon-colored pigment into

the medium, those of the present fungus form a salmon-

colored pigment that remain restricted to the inoculation

sites.

Cosmospora vilior (Starb.) Rossman & Samuels in

Rossman, Samuels, Rogerson & Lowen, Stud. Mycol.

42: 126. 1999.

Figures 1Q, 2J-M

Anamorph. Acremonium berkeleyanum (P. Karst.) W.

Gams, Netherlands J. Pl. Pathol. 88: 76. 1982.

Stromata inconspicuous. Perithecia solitary, rarely

aggregated, superficial, red, changing color to dark red

in 10% KOH and yellow in lactophenol, obpyriform,

212-250 ¡Ñ 187-215 £gm, smooth, laterally collapsed when

dry; perithecial wall 20-40 £gm thick, one-layered; ostioles

obtuse. Asci cylindrical, 51-65 ¡Ñ 5-7 £gm, with an apical

ring. Ascospores yellowish, broadly ellipsoidal, 1-septate,

constricted at septum, 5-8.5 ¡Ñ 3-4 £gm, warted.

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Botanical Studies, Vol. 48, 2007

Cultures and anamorph. Colonies attaining 1-1.8 cm

diam in 7 days at 20¢XC, with abundant aerial hyphae,

usually slimy, yellow-green; aerial hyphae often

aggregated into acute strands. Conidiophores arising

from aerial hyphae and agar surface, usually unbranched,

aseptate; phialides cylindrical, 40-60 ¡Ñ 2-4 £gm, tapering

to 1 £gm near apex. Conidia hyaline, oblong ellipsoidal,

aseptate, 3-5 ¡Ñ 1-2.5 £gm, smooth.

Specimens examined. Tainan County, Hsinhwa, on

Kretzshmaria sp., 15 May 2000, Ju, Y.-M & Hsieh, H.-M.

89051501 (cultured). Taipei County, Wulai, Hsinsian

elementary school, on Rosellinia, 29 Jun 2000, Ju, Y.-M.

89062901 (cultured). Ilan County, Fu-shan, on Rosellinia,

27 Oct 2001, Ju, Y.-M & Hsieh, H.-M. 90102708. Pingtung

County, Hengchun, Kenting, on decaying stromata of a

fungus, 26 Nov 2000, Ju, Y.-M & Hsieh, H.-M. 89112603.

Pingtung County, Hengchun, Kenting, on decaying

Hypoxylon stromata, 23 Sep 2002, Ju, Y.-M & Hsieh,

H.-M. 91092332 (cultured). Miaoli County, Taiann, on

decayed twigs, 8 Jul 2005, Ju, Y.-M & Hsieh, H.-M.

94070809 (cultured). Taipei City, Tienmu historical trail,

on bark, 25 Aug 2005, Ju, Y.-M & Hsieh, H.-M. 94082503

(cultured). Taipei County, Nankang-shan, on stromata

of Krezschmaria pavimentosa, 1 Nov 2005, Guu, J.-R

94110102(cultured). Hwalien County, Showlinn, Sacadung

trail, on stromata of Kretzschmaria pavimentosa, 10 Dec

2005, Guu, J.-R. 94121003 (cultured).

Notes. This fungus is commonly encountered in tropical

and subtropical areas. It has been found to grow on the

stromata of various xylariaceous fungi in Taiwan. Cultures

of C. vilior are peculiar in abundantly diffusing a yellow-

green pigment into the medium. The anamorph produced

by the Taiwan material is much as described in Samuels et

al. (1990).

Haematonectria haematococca (Berk. & Broome)

Samuels & Nirenberg in Rossman, Samuels,

Rogerson & Lowen, Stud. Mycol. 42: 135. 1999.

Fiugres 1C, 5G-K

Anamorph. Fusarium solani (Martius) Saccardo, Michelia

2: 296. 1881.

Stromata lacking. Perithecia solitary to aggregated

in groups of 20, superficial, red, becoming dark red in

10% KOH and yellow in lactophenol, subglobose to

broadly obpyriform, 225-270 ¡Ñ 230-250 £gm, surface

with red to yellowish warts up to 70 £gm high, collapsed

when dry; perithecial wall 60-95 £gm thick, two-layered;

ostioles acute, paler than perithecial body. Asci cylindrical

to clavate, 55-90 ¡Ñ 8-11 £gm, lacking an apical ring.

Ascospores yellowish, oblong-ellipsoidal, 1-septate, 10-14

¡Ñ 4-6.5 £gm, striate.

Cultures and anamorph. Colonies averaging 7 cm

diam in 7 days at 20¢XC, floccose, white, sometimes

becoming pale brown in the center. Conidiophores

producing macroconidia and microconidia; those bearing

macroconidia aggregated into cream-colored sporodochia,

cylindrical, 9-12 ¡Ñ 5-7 £gm, irregularly branched, bearing

1-3 phialides at each terminus, with the phialides

cylindrical, 11-23 ¡Ñ 3-5 £gm; those bearing microconidia

usually unbranched, with the phialides cylindrical, 22-31

¡Ñ 3-4 £gm. Macroconidia hyaline, cylindrical to fusoid,

with wedge-shaped to obtuse foot cells, 3- to 5-septate,

32-77 ¡Ñ 4-6 £gm, smooth. Microconidia hyaline, allantoid

to cylindrical, (6-)9-13 ¡Ñ 2-4 £gm, smooth, produced singly

or in small, slimy droplets. Chlamydospores globose,

hyaline, smooth to roughened, 7-10 £gm, produced singly

or in chains.

Specimens examined. Kaohsiung County, Maolinn, on

bark, 14 Sep 2005, Guu, J.-R. 94091406 (cultured). Taipei

County, Wanlee, Ruei-chuan historical trail, on bark, 23 Jul

2005, Guu, J.-R. and Kuo, H.-P. 94072307. Taipei County,

Pingshi, Chou-tou-shan, on bark, Guu, J.-R. 94043010

(cultured). Ilan County, Fu-shan botanical garden, on bark,

24 Jan 2005, Guu, J.-R. 94012410. Taipei County, Sungi,

Datunshi historical trail, on bark, 22 Jan 2005, Guu, J.-R.

94012205. Taipei County, Wanlee, Linnshih historical

trail, on bark, 15 Jan 2005, Guu, J.-R. 94011504 (cultured).

Ilan County, Guei-shan-dao, on bark, 9 Jun 2003, Ju, Y.-M.

92060901 (cultured); Nantuo County, Tsuifong, on bark, 6

Apr 2003, Ju, Y.-M. 92040605 (cultured). Taipei County,

Hsintien, Shih-tzu-tuo-shan, on bark, 11 Dec 2004, Guu,

J.-R. 93121104. Taipei City, Shilinn, Shengren waterfall,

on bark, 10 Oct 2004, Guu, J.-R. 93101003. Taipei County,

Sijih, Hsin-shan-mon-hu, on bark, 14 Nov 2004, Guu,

J.-R. 93111402. Taipei County, Pingshi, Chou-tuo-shan,

on bark, 31 Oct 2004, Guu, J.-R. 93103104. Taipei City,

Muja, campus of National Chenchi Univ., on bark, 25 Sep

2004, Shih, H.-H. 93092506. Taipei County, Hsintien, Ta-

tung-shan, on bark, 19 Sep 2004, Kuo, H.-P. 93091906.

Ilan County, Tatung, Chi-lan-shan, on bark, 7 Sep 2004,

Ju, Y.-M. 93090706; Hwalien County, Jaofong farm, on

bark, 22 Jul 2004, Shih, H.-H. 93072201 (cultured). Taipei

County, Jingtung, Jingtung historical trail, on bark, 21

Dec 2003, Guu, J.-R. 92122103 (cultured). Taipei County,

Pingshi, Chung-yang-chien-shan, on bark, 4 Dec 2003,

Guu, J.-R. 92120403. Taipei County, Pingshi, Chung-

yang-chien-shan, on bark, 20 Nov 2003, Guu, J.-R.

92112007 (cultured). Taipei County, Hsintien, Yinhodong,

on bark, 16 Oct 2003, Guu, J.-R. 92101606. Taipei City,

Shihlinn, Ping-ding-gu-jyun, on bark, 9 Aug 2003, Guu,

J.-R. 92080906.

Notes. The teleomorph of this species is frequently

encountered on decaying branches and fruits. As with

Samuels (1976) and Samuels et al. (1990), the teleomorph

was also produced in cultures obtained from the Taiwan

collections. The anamorph that we obtained from the

Taiwan material fits well the description

in Booth (1960).

It is interesting to note that perithecia of Ophionectria

trichospora and H. haematococca were found to be mixed

on the same substrata in several occasions.

Lanatonectria flavolanata (Berk. & Broome) Samules &

Rossman in Rossman, Samuels, Rogerson & Lowen,

Stud. Mycol. 42: 139. 1999.

Figures 1K, 3H, I

GUU et al. ¡X Nectriaceae in Taiwan

195

Anamorph: Actinostilbe sp.

Stromata usually erumpent from the bark, sometimes

with conidiomata. Perithecia sometimes accumulating at

base of synnemata or formed without anamorph, mostly

aggregated, in groups of 5-15, rarely solitary, superficial,

red, changing color to dark red in 10% KOH and yellow in

lactophenol, subglobose to broadly obpyriform, 300-450

¡Ñ 200-320 £gm, uncollapsed when dry, with surface

slightly roughened with perithecial hairs; perithecial wall

25-35 £gm thick, two-layered; perithecial hairs septate,

unbranched, flexuous, verrucose, golden yellow, hooked,

40-60 ¡Ñ 4-6 £gm, usually collapsed in age; ostioles slightly

papillate, paler than perithecial body. Asci clavate, 45-60 ¡Ñ

7-8 £gm, with an apical ring. Ascospores hyaline, oblong-

ellipsoidal, 1-septate, 15-20 ¡Ñ 5-5.5 £gm, striated.

Cultures and anamorph. Colonies 2-3 cm in 7 days

at 20¢XC, with abundant aerial hyphae; reverse yellowish

brown. Synnemata abundant, 0.5-5 mm long ¡Ñ 0.1-0.3

mm broad, white, furfuraceous with short, golden, septate,

3-4-£gm-wide hairs along the length, bearing a terminal,

hemispherical conidial mass; hairs often covered with

droplets of golden exudates. Conidiophores aggregated

at top of synnemata, anastomosing laterally, branched,

bearing 2-3 phialides at each terminus; phialides smooth or

finely roughened, cylindrical, 20-35 ¡Ñ 1.5-3 £gm. Conidia

hyaline, oblong-ellipsoidal, 1-septate, 10-14 ¡Ñ 4-5 £gm,

smooth, with a flat base.

Specimens examined. Nantou County, Luku, Shitou,

o n Brugmansia suaveolens (Willd.) Bercht. & Presl, 19

Apr 2003, Ju, Y.-M. & Hsieh, H.-M. 92041910 (cultured).

Taipei County, Jingtung, Jingtung historical trail, 21 Dec

2003, Guu, J.-R. 92122105

(cultured). Taipei County,

Jingtung, Jingtung historical trail, 18 Mar 2004, Guu, J.-R.

93031804.

Notes. Lanatonectria is mainly characterized by the

golden-yellow hairs covering the perithecia. Our material

o f L. flavolanata produced an Actinostilbe anamorph in

culture, which is much as that described by Rossman

et al. (1999). Lanatonectria flavolanata differs from L.

flocculenta in having larger ascospores and a synnematous

anamorph. The anamorph of L. flocculenta is sporodochial.

Lanatonectria flocculenta (Henn. & E. Nyman) Samuels

& Rossman in Rossman, Samuels, Rogerson & Lowen,

Stud. Mycol. 42: 138. 1999.

Figures 1J, 3A-G

Anamorph. Actinostilbe macalpinei (Agnihothr. & G.C.S.

Barua) Seifert & Samuels, Stud. Mycol. 42: 138. 1999.

Stromata erumpent from bark, usually associated with

conidiomata, inconspicuous. Perithecia solitary or in

groups of 2-20, superficial, red, changing color to dark

red in 10% KOH and yellow in lactophenol, globose to

subglobose, 180-300 ¡Ñ 200-295 £gm, uncollapsed when

dry, slightly roughened with perithecial hairs on the

surface; perithecial wall 25-30 £gm thick, two-layered;

perithecial hairs straight or hooked, septate, unbranched,

golden yellow, conspicuously verrucose, 30-70 ¡Ñ 4-6 £gm;

ostiols usually papillate, paler than perithecial body. Asci

clavate, 35-70 ¡Ñ 6-10 £gm, with an apical ring. Ascospores

hyaline, oblong-ellipsoidal, 1-septate, 11-13 ¡Ñ 3-4 £gm,

striated.

Cultures and anamorph. Colonies attaining 5 cm diam

in 7 days at 20¢XC, with abundant floccose aerial hyphae;

reverse yellow. Conidiophores aggregated into well-

developed, solitary or gregarious sporodochia, irregularly

branched, usually anastomosing laterally. Conidia hyaline,

yellow in masses, oblong ellipsoidal, 1-septate, 6-8 ¡Ñ 2-3.5

£gm, smooth.

Specimens examined. Tainan County, Hsinhwa, on

bark, 14 May 2000, Ju, Y.-M & Hsieh, H.-M. 89051406

(cultured). Taipei City, Nankang, campus of Academia

Sinica, 8 Jun 2001, Ju, Y.-M. 90060801. Nantou County,

Luku, Shitou, on Brugmansia suaveolens (Willd.)

Bercht. & Presl, 19 Apr 2003, Ju, Y.-M. & Hsieh, H.-M.

92041905 (cultured). Taipei City, Nankang, Tu-ku-

yue, on bark, 10 Aug 2003, Guu, J.-R. 92081001. Taipei

County, Shihding, Huang-di-dian north peak trail, on

bark, 27 Nov 2003, Guu, J.-R. 92112705 (cultured). Taipei

City, Muja, Jihnan temple, on bark, 7 Dec 2003, Guu,

J.-R. 92120701 (cultured). Taipei City, Muja, Chanshan

temple, on bark, 11 Apr 2004, Guu, J.-R. 93041104.

Hwalien County, Jaofong farm, on bark of Zelkova serrata

(Thunb.) Makino, 22 Jul 2004, Shih, S.-H. and Chao,

T.-H. 93072202

(cultured). Taipei City, Muja, Changshan

temple, on bark, 25 Sep 2004, Shih, S.-H. 93092511

(cultured). Taipei County, Wulai, Chia-jiou-liao trail, on

bark, 21 Aug 2005, Guu, J.-R. 94082102 (cultured). Tainan

County, Meilinn, on Areca catechu L., 29 Aug 2005, Ju,

Y.-M. & Hsieh, H.-M. 94082904 (cultured).

Notes. This species is one of the most commonly

encountered nectriaceous fungi in pantropical regions

(Rossman et al., 1999; Samuels et al., 1990; Samuels

and Brayford, 1994). The other Lanatonectria species

found in Taiwan has larger ascospores and a synnematous

anamorph. The cultures from specimen 94082904

produced the teleomorph on PDA. Like Rossman et al.

(1999), we also observed an Actinostilbe macalpinei

anamorph produced in the cultures initiated from our

collections of L. flocculenta.

Nectria balsamea Cooke & Peck in Cooke, Grevillea 12:

81. 1884.

Figures 1M, 5E, F

Anamorph. Zythiostroma sp.

Stromata inconspicuous. Perithecia solitary, superficial,

red, changing color to dark red in 10% KOH and yellow

in lactophenol, subglobose, 250-380 £gm, overlain with

white scales, uncollapsed when dry; ostioles papillate.

Asci clavate, 72-89 ¡Ñ 9-13 £gm, lacking an apical ring.

Ascospores hyaline, broadly fusiform, muriform,

(18-)20-25.5(-30) ¡Ñ (4.5-)5.5-6(-6.5) £gm, smooth.

Cultures and anamorph. Colonies attaining 3 cm diam

in 10 days at 20¢XC, submerged, with scarce aerial hyphae,

slimy at the inoculation points; reverse light yellow first

196

Botanical Studies, Vol. 48, 2007

but becoming brown after 14 days. Conidiomata dark

red, pulvinate, multiloculate. Conidiophores cylindrical,

5-10 ¡Ñ 1-2 £gm. Conidia hyaline, allantoid, 4-6 ¡Ñ 1-2 £gm,

smooth, usually aggregated into orange-red, slimy masses

at ostioles.

Specimen examined. Taichung County, Pilushi, on twig

of Pinus taiwanensis Hayata cone, 24 May 2005, Guu,

J.-R. 94052402 (cultured).

Notes. Nectria balsamea features

muriform ascospores

that produce ascoconidia within asci. It is the only Nectria

species found to be associated with a conifer in Taiwan.

Our specimen agrees well with the descriptions and

illustrations given by Seeler (1940), Booth (1959), and

Rossman (1985) [as Thyronectria]. The anamorph that

we obtained from the Taiwan material closely resembles

that described and illustrated in Booth (1959). Rossman

et al. (1999) assigned the anamorph to the form-genus

Zythiostroma.

Nectria pseudotrichia (Schwein.) Berk. & M. A.

Curtis, J. Acad. Nat. Sci. Philadelphia 2, 2: 289. 1853.

Figures 1A, 5A-D

Anamorph. Tubercularia lateritia (Berk.) Seifert, Stud.

Mycol. 27: 119. 1985.

Stromata inconspicuous. Perithecia solitary or

aggregated, superficial, red, changing color to dark red

in 10% KOH and yellow in lactophenol, globose or

subglobose, usually associated with synnemata, 250-400

£gm, scaly, cupulate when dry; perithecial wall 50-85 £gm,

three-layered; ostioles slightly papillate. Asci clavate,

60-90 ¡Ñ 12-22 £gm, lacking an apical ring. Ascospores

hyaline, light yellow in age, broadly ellipsoidal, muriform,

19-24 ¡Ñ 9-12 £gm, smooth.

Cultures and anamorph. Colonies attaining 9 cm in 5

days at 20¢XC with scanty aerial hyphae, a pink pigment

diffusing into the medium. Conidiophores mononematous,

monoverticillate or biverticillate, bearing cylindrical

phialides; stipes warted, cylindrical, straight, curved

or twisted. Conidia ellipsoidal, hyaline, 3-7 ¡Ñ 2-3 £gm.

Synnemata produced in nature but lacking in culture,

solitary to aggregated, erected, unbranched, 500-5000 ¡Ñ

50-250 £gm, red-brown, grading into light cream-yellow

towards the top, bearing a spherical conidial mass.

Specimens examined. Taipei County, Wulai, Chia-jiou-

liao trail, on bark, 21 Aug 2005, Guu, J.-R. 94082103

(cultured). Taipei County, Wanlee, Ruei-chuan historical

trail, on bark, 23 Jul 2005, Guu, J.-R. and Kuo, H.-P.

94072304 (cultured). Taipei City, campus of National

Taiwan Univ., on bark, 17 Jul 2005, Guu, J.-R. 94071701

(cultured). Kaohsiung County, Lioguei, Shanping, on

bark, 10 Mar 2005, Guu, J.-R. 94031010 (cultured). Taipei

County, Wanlee, Linnshih historical trail, on bark, 15 Jan

2005, Guu, J.-R. 94011505 (cultured). Taipei County,

Hsintien, Shih-tzu-tuo-shan, on bark, 11 Dec 2004, Guu,

J.-R. 93121109; Taipei County, Hsintien, Yinhodong,

on bark, 25 Nov 2004, Shih, H.-H. 93112506 (cultured).

Pingtung County, Hengchun, Kenting, on bark, 28 Aug

2004, Ju, Y.-M and Hsieh, H.-M. 93082818. Pingtung

County, Meinong, on bark of Swietenia macrophylla King,

27 Aug 2004, Shih, H.-H. 93082701. Taipei City, campus

of National Taiwan Univ., on bark, 1 Aug 2004, Guu, J.-R.

93080101. Taipei City, Wolong street, on bark, 8 May

2004, Shih, H.-H. 93050801; Taipei County, Hsintien,

Shih-tzu-tuo-shan, on bark, 2 May 2004, Guu, J.-R.

93050201 (cultured). Tainan County, Tungshan, on bark of

Mangifera indica, 4 Apr 2004, Guu, J.-R. and Chang, K.-T.

93040403. Taipei County, Shihding, Huang-di-dian north

peak trail, on bark, 27 Nov 2003, Guu, J.-R. 92112708

(cultured). Taipei County, Shihding, Huang-di-dian, on

bark, 13 Nov 2003, Guu, J.-R. 92111301 (cultured). Taipei

County, Mudan, Diaoshan historical trail, on bark, 2 Nov

2003, Guu, J.-R. 92110201 (cultured). Taipei County,

Hsintien, Yinhodong, on bark, 16 Oct 2003, Guu, J.-R.

92101605. Taipei County, Hsintien, Jih-tan-shan, on bark,

9 Oct 2003, Guu, J.-R. 92100904. Taipei City, Shihlinn,

Ping-ding-gu-jyun, on bark, 14 Sep 2003, Guu, J.-R.

92091404. Taipei City, Songshan, Hu-shan trail, on bark,

10 Sep 2003, Guu, J.-R. 92091003. Ilan County, Fu-shan,

on fruit of persimmon, 12 Nov 2002, Ju, Y.-M. 91111221;

Ilan County, Fu-shan, on bark, 19 Aug 2001, Ju, Y.-M.

90081911. Taipei County, Sunshai, Mann-yue-yuan, on

bark, 27 Sep 2000, Ju, Y.-M. 89092709. Pingtung County,

Hengchun, Kenting, on bark, 16 Sep 2000, Ju, Y.-M and

Hsieh, H.-M. 89091638. Taipei City, Nankang-shan, on

bark, 22 Mar 2000, Ju, Y.-M and Hsieh, H.-M. 89032211.

Notes. This is a frequently encountered species in

the tropics and subtropics (Seifert, 1985; Samuels et al.,

1990). It features muriform ascospores and a synnematous

anamorph, with which the perithecia are constantly

associated. Chen and Hsieh (1996, as Thyronectria) has

previously reported this speices in Taiwan. Also see Seifert

(1985) for a description of the anamorph.

Neonectria coronata (Penz. & Sacc.) Mantiri & Samuels,

Canad. J. Bot. 79: 339. 2001.

Figures 1I, 4F-I

Anamorph. Cylindrocarpon coronatum Brayford &

Samuels, Mycologia 85: 620. 1993.

Stromata inconspicuous. Perithecia solitary to

aggregated, superficial, red, changing color to dark red

in 10% KOH and yellow in lactophenol, subglobose or

broadly obpyriform, 200-450 ¡Ñ 150-300 £gm, slightly

roughened, uncollapsed when dry; perithecial wall 15-30

£gm thick, one-layered; ostioles papillate, located in the

center of a black disc 150-200 £gm diam,

encircled with a

conspicuous crown of saccate cells. Asci clavate, 60-75

¡Ñ 10-15 £gm, lacking an apical ring. Ascospores hyaline,

becoming yellowish, broadly fusiform, 1-septate, 19-23 ¡Ñ

6.5-8 £gm, with discontinuous striations.

Cultures and anamorph. Colonies attaining 2-5 cm

diam in 10 days at 20¢XC, floccose, light brown, with scanty

aerial hyphae. Conidiophores producing macroconidia

and microconidia; those bearing macroconidia aggregated

into slimy, scattered sporodochia, branched, producing a

GUU et al. ¡X Nectriaceae in Taiwan

197

phialide at each terminus, with the phialides cylindrical,

10-20 ¡Ñ 3-5 £gm, with periclinal thickening at the apex;

those bearing microconidia cylindrical, unbranched, 12-26

¡Ñ 2-4 £gm. Macroconidia curved, fusiform, mostly 5- to

7-septate, 60-81 ¡Ñ 6-9 £gm, with blunt rounded apical cell,

less frequently, 1-septate, 30-40 ¡Ñ 6-7 £gm, or 3-septate,

40-52 ¡Ñ 6-7 £gm. Microconidia ellipsoidal, hyaline, 6-8 ¡Ñ

2-3.5 £gm, smooth.

Specimens examined. Taipei County, Hsintien, Jhih-

tan-shan, on bark, 9 Oct 2003, Guu, J.-R. 92100902

(cultured). Taipei County, Pingshi, Chung-yang-chien-

shan, on bark, 20 Nov 2003, Guu, J.-R. 92112005. Taipei

County, Jingtung, Jingtung historical trail, on bark, 21

Dec 2003, Guu, J.-R. 92122102 (cultured). Taipei County,

Jingtung, Jingtung historical trail, on bark, 18 Mar 2004,

Guu, J.-R. 93031807. Taipei County, Hsintien, Shih-tzu-

tou-shan, on bark, 21 Mar 2004, Guu, J.-R. 93032105. Ilan

County, Fu-shan, Hapenshi, on bark, 21 Aug 2004, Guu,

J.-R. 93082102 (cultured). Taipei County, Hsintien, Shih-

tzu-tou-shan, on bark, 11 Dec 2004, Guu, J.-R. 93121102.

Taipei County, Wanlee, Linnshih historical trail, on bark,

15 Jan 2005, Guu, J.-R. 94011502 (cultured). Taipei

County, Pingshi, Chou-tou-shan, on bark, 30 Apr 2005,

Guu, J.-R. 94043006 (cultured).

Notes. Neonectria coronata is commonly found on the

bark of recently dead trees. It is easily recognized by the

conspicuous crown around the ostiolar disc. It should be

noted that perithecia were formed on PDA from cultures

obtained from specimen 94043006. The anamorph that

we obtained from our collections is much as described by

Brayford and Samuels (1993).

Neonectria discophora (Mont.) Mantiri & Samuels,

Canad. J. Bot. 79: 339. 2001. Figures 1E, 4A-E

Anamorph. Cylindrocarpon ianthothele Wollenweber var.

majus Wollenweber, Z. Parasitenk. (Berlin) 1: 161.

1928.

Stromata inconspicuous. Perithecia solitary to

aggregated, superficial or partially immersed in

substratum or basal stroma, red, changing color to dark

red in 10% KOH and yellow in lactophenol, smooth and

shining, globose to subglobose, 293-355 ¡Ñ 390-430 £gm,

uncollapsed when dry; perithecial wall 39-45 £gm thick,

two-layered; ostioles blackened in age, slightly papillate.

Asci cylindrical, 60-72 ¡Ñ 6-9 £gm, with an apical ring.

Ascospores yellowish, broadly fusiform, 1-septate, 14-16

¡Ñ 5.5-6.5 £gm, echinulate.

Cultures and anamorph. Colonies attaining 2 cm

diam in 7 days at 20¢XC, aerial hyphae scanty, mostly flat,

arising from the inoculum; hyphae often filled with brown

inclusions, diffusing a violet pigment into the medium.

Conidiophores producing macroconidia only, aggregated

into slimy sporodochia, cylindrical, smooth, usually

unbranched, each producing a phialide, with the phialides

cylindrical, 10-15 ¡Ñ 2-4 £gm. Macroconidia hyaline,

oblong-ellipsoidal, slightly curved, with rounded ends, 3-

to 5-septate, 32-80 ¡Ñ 6-8 £gm, smooth.

Specimens examined. Ilan County, Fu-Shan, on bark,

23 Jun 2001, Ju, Y.-M. and Hsieh, H.-M. 90062304.

Taipei County, Jingtung, Jingtung historical trail, on bark,

21 Dec 2003, Guu, J.-R. 92122107 (cultured). Taipei

County, Jingtung, Jingtung historical trail, on bark, 18

Mar 2004, Guu, J.-R. 93031802. Ilan County, Fu-Shan,

on bark, 24 Jan 2005, Guu, J.-R. 94012407 (cultured).

Kaohsiung County, Liou-guei, Shan-ping, on bark, 10

Mar 2005, Guu, J.-R. 94031007 (cultured). Taipei County,

Pingshi, Chou-tou-shan, on bark, 30 Apr 2005, Guu, J.-R.

94043003 (cultured). Chiayi County, Shijuo, on bark, with

Neonectria lucida, 20 Nov 2005, Guu, J.-R. 94112003.

Notes. This fungus and N. lucida are the most

commonly encountered species in the N. discophora group

(Samuels et al., 1990). These two species are mainly

separated by colony color. The colonies of N. discophora

diffuse a violet pigment into the culture medium, while

those of N. lucida yield a tan-colored pigment restricted to

the surrounding area of inoculation points. The anamorph

that we obtained from the Taiwan material fits well

Cylindrocarpon ianthothele (Brayford et al., 2004).

Neonectria jungneri (Henn.) Samuels & Brayford,

Mycologia 96: 580. 2004.

Figures 1H, 4O-R

Anamorph. Cylindrocarpon victoriae Wollenw., Zentbl.

Bakt. ParasitKde, Abt. II 1: 161. 1928.

Stromata absent. Perithecia aggregated, superficial,

red, changing color to dark red in 10% KOH and yellow

in lactophenol, obpyriform, 350-400 ¡Ñ 220-330 £gm,

roughened, uncollapsed when dry; perithecial wall 20-35

£gm thick, two-layered; ostioles coarsely papillate, red

or gray, slightly constricted below. Asci clavate, 75-90

¡Ñ 15-22 £gm, lacking an apical ring. Ascospores hyaline,

broadly fusiform, 1-septate, 20-21.5 ¡Ñ (7.5-)8-9.5 £gm,

striated.

Cultures and anamorph. Colonies attaining 4 cm diam

in 7 days at 20¢XC, velutinous, white to light yellow first,

becoming brown from center outwards. Conidiophores

producing macroconidia only, aggregated into slimy

sporodochia, irregularly branched, producing 1 or 2

phialides at each terminus, with the phialides cylindrical,

15-22 ¡Ñ 3-5 £gm. Macroconidia oblong-ellipsoidal, hyaline,

5- to 7-septate; 5-septate conidia 93-100 ¡Ñ 10-10.5 £gm;

6-septate conidia 113-115.5 ¡Ñ 10 £gm; 7-septate conidia

106-121.5 ¡Ñ 10-10.5 £gm.

Specimens examined. Kaohsiung County, Meinong,

on bark of Swietenia macrophylla King, 16 Sep 2005,

Guu, J.-R. 94091601 (cultured). Pingtung County,

Machia, Liang-shan waterfall, on bark, 14 Sep 2005,

Guu, J.-R. 94091407 (cultured). Nantou County, Yuchee,

Lienhwachee, on bark of Castanopsis carlesii (Hesml.)

Hayata

var. sessilis Nakai, 19 Jul 2001, Ju, Y.-M. 90071905

(cultured). Pingtung County, Hengchun, Kenting, on bark

of Castanopsis carlesii var. sessilis, 26 Nov 2000, Ju, Y.-M

and Hsieh, H.-M. 89112623.

198

Botanical Studies, Vol. 48, 2007

Notes. This species resembles N. coronata but differs

from the latter in lacking an ostiolar disc, in the ostiole

not being surrounded with a crown, and in the ascospores

ornamented with discontinuous striations. Also see

Samuels and Brayford (1994) for a description of the

anamorph.

Neonectria lucida (Hohn.) Samuels & Brayford,

Mycologia. 96: 590. 2004. Figure 1F

Anamorph. Cylindrocarpon lucidum Booth, Mycol. Pap.

104: 21. 1966.

Stromata absent. Perithecia solitary or aggregated,

superficial, red, changing color to dark red in 10% KOH

and yellow in lactophenol, obpyriform, 400-460 £gm diam,

slightly roughened, uncollapsed when dry; perithecial wall

20-35 £gm thick, two-layered; ostioles coarsely papillate,

80-110 £gm diam, 12-30 £gm high, slightly darker than

the perithecial bodies. Asci cylindrical, 58-70 ¡Ñ 7-9 £gm,

with an apical ring. Ascospores hyaline, broadly fusiform,

1-septate, (12.5-)13-15 ¡Ñ (4.5-)5-6 £gm, echinulate.

Cultures and anamorph. Colonies attaining 3 cm diam

in 7 days at 20¢XC, tan-colored close to the inoculation

points, becoming white towards the edges. Conidiophores

producing macroconidia only, aggregated into slimy

sporodochia, irregularly branched, producing 1-2 phialides

at each terminus, with the phialides cylindrical, 15-22

¡Ñ 3-5 £gm. Macroconidia oblong-ellipsoidal, sometimes

curved, 3- to 5-septate; 3-septate conidia 40-82 ¡Ñ 5-7 £gm;

4-septate conidia 50-60 ¡Ñ 5-8 £gm; 5-septate conidia 60-85

¡Ñ 6-8 £gm.

Specimens examined. Nantou County, Tsuifong, on

bark, 23 Sep 2002, Ju, Y.-M and Hsieh, H.-M. 91092333.

Taipei County, Jingtung, Jingtung historical trail, 27 Nov

2003, Guu, J.-R. 92112704 (cultured). Taipei County,

Pingshi, Chou-tou-shan, on bark, 30 Apr 2005, Guu, J.-R.

94043002 (cultured). Kaohsiung County, Meinong, on

bark of Swietenia macrophylla King, 16 Sep 2005, Guu,

J.-R. 94091602 (cultured). Chiayi County, Shih-jhuo, on

bark, with Neonectria discophora, 20 Nov 2005, Guu,

J.-R. 94112004.

Figure 3. A-G, Lanatonectria flocculenta. A, Median section of ascoma; B, Lateral wall; C, Ascus; D, Verrucose perithecial hairs; E,

Ascal tip with an apical ring; F, Sporodochium; G, Conidia; H-I, Lanatonectria flavolanata; H, Striated ascospores; I, Synnema bearing

a conidial mass. Scale bars: A = 50 £gm; B, C = 10 £gm; D, G, H = 8 £gm; E = 3 £gm; F = 50 £gm; I = 500 £gm.

GUU et al. ¡X Nectriaceae in Taiwan

199

Figure 4. A-E, Neonectria disccophora. A, Median section of ascoma; B, Ascus; C, Ascospores; D, Ascal tip with an apical ring;

E, Macroconidia; F-I, Neonectria coronata. F, Median section of ascoma; G, Macroconidia and microconidia; H, Ascospores with

discontinuous striations; I, Lateral wall. J-N, Neonectria rugulosa. J, Median section of ascoma; K, Lateral wall; L, Ascus; M,

Ascospores with continuous striations; N, Macroconidia and microconidia. O-R, Neonectria jungneri. O, Macroconidia; P, Coarsely

papillate ostiole (arrow); Q, Median section of ascoma; R, Lateral wall. Scale bars: A, P = 50 £gm; B, E, G, K, M, O = 10 £gm; C, D = 5

£gm; F, R = 20 £gm; H = 8 £gm; I = 15 £gm; J, N, Q = 30 £gm; L = 12 £gm.

200

Botanical Studies, Vol. 48, 2007

Notes. Cultures from specimens 92112704 and

94091602 produced the teleomorph on PDA. Also see

notes on Neonectria discophora. The anamorph obtained

from the Taiwan material is much as described in Booth

(1966) and Brayford et al. (2004).

Neonectria rugulosa (Pat. & Gaillard) Mantiri

& Samuels, Canad. J. Bot. 79: 339. 2001.

Figures 1G, 4J-N

Anamorph. Cylindrocarpon rugulosum Brayford &

Samuels, in Samuels & Brayford, Sydowia 46: 148.

1994.

Figure 5. A-D, Nectria pseudotrichia. A, Median section of ascoma; B, Lateral wall; C, Ascus; D, Ascospore. E-F, Nectria balsamea. E,

Ascospores forming ascoconidia in ascus; F, ascospores germinating into ascoconidia. G-K, Haematonectria haematococca. G, Median

section of ascoma; H, Ascus; I, Ascospores; J, Conidiophores and macroconidia; K, Macroconidium; L-P, Ophionectria trichospora; L,

Median section of ascoma; M, Lateral wall; N, Ascus; O, Ascospore with spiral striations; P, Conidia. Scale bars: A = 60 £gm; B, J, M, N,

P = 20 £gm; G = 100 £gm; L = 50 £gm; C, F, H, I, K, O = 3 £gm; D, E = 8 £gm.

GUU et al. ¡X Nectriaceae in Taiwan

201

Stromata absent. Perithecia solitary to aggregated, red

or orange-red, changing color to dark red in 10% KOH and

yellow in lactophenol, globose to subglobose, 300-450 £gm,

roughened by red warts, uncollapsed when dry; perithecial

wall 15-23 £gm thick, two-layered; ostioles apapillate.

Asci clavate, 55-79 ¡Ñ 9-13 £gm, lacking an apical ring.

Ascospores hyaline, broadly fusiform, 1-septate, 22-25 ¡Ñ

7.5-9.5 £gm, striate.

Cultures and anamorph. Colonies attaining 7 cm in 14

days at 20¢XC, velutinous, white, or light brown, diffusing

a light brown pigment into medium. Conidiophores

producing macroconidia and microconidia; those bearing

macroconidia aggregated into slimy sporodochia, rarely

branched, producing 3-5 phialides at each terminus, with

the phialides cylindrical, 10-20 ¡Ñ 2-3 £gm; those bearing

microconidia cylindrical, unbranched, 30-90 ¡Ñ 2-3 £gm.

Macroconidia hyaline, oblong-ellipsoidal, sometimes

curved, 4- to 7-septate, 53-82 ¡Ñ 4.5-6.5 £gm. Microconidia

hyaline, ellipsoidal or oblong-ellipsoidal, aseptate, 7-13 ¡Ñ

2.5-3.5 £gm.

Specimens examined. Ilan County, Fu-shan, on bark

o f Castanopsis carlesii var. sessilis, 23 Jun 2001, Ju,

Y.-M 90062306 (cultured). Nantou County, Yuchee,

Lienhwachee, on bark of Castanopsis carlesii var. sessilis,

19 Apr 2001, Ju, Y.-M. 90041902 (cultured). Nantou

County, Yuchee, Lienhwachee, on bark of Castanopsis

carlesii var. sessilis, 19 Jul 2001, Ju, Y.-M. 90071903

(cultured). Taipei County, Yong-ming-shan, Lu-jiao-

keng shi, on bark, 10 Jul, Guu, J.-R. 93071001 (cultured).

Kaohsiung County, Liouguei, Shanping, on bark, 10 Mar

2005, Guu, J.-R. 94031008. Tainan County, Meilinn, on

Areca catechu L., 29 Aug 2005, Ju, Y.-M. & Hsieh, H.-M.

94082904 (cultured). Pingtung County, Machia, Liang-

shan waterfall, on bark, 14 Sep 2005, Guu, J.-R. 94091408

(cultured).

Notes. Among the Neonectria species included in the

present study, only this species and N. discophora have

globose perithecia. They differ in the former having

apapillate ostioles, striate ascospores, and producing

both macroconidia and microconidia in culture. Samuels

and Brayford (1994) reported the production of the

teleomorph in culture, but no perithecia formed in cultures

that we obtained from ascospores of our collections. The

anamorph obtained from the Taiwan material agrees well

with the description in Samuels & Brayford (1994).

Ophionectria trichospora (Berk. & Broome) Sacc.,

Michelia 1: 323. 1878.

Figures 1L, 5L-P

Anamorph. Antipodium spectabile Piroz., Canad. J. Bot.

52: 1144. 1974.

Stromata absent. Perithecia solitary to aggregated,

red, changing color to dark red in 10% KOH, yellow in

lactophenol, ovoidal, 400-600 ¡Ñ 250-300 £gm, roughened

by concolorous warts, sometimes collapsed when dry;

perithecial wall 30-90 £gm thick, two-layered; ostioles

inconspicuous. Asci clavate, 180-220 ¡Ñ 25-30 £gm, lacking

an apical ring. Ascospores hyaline, vermiform, 13- to

20-septate, 190-210 ¡Ñ 9-11 £gm, inconspicuously striate.

Cultures and anamorph. Colonies attaining 2 cm diam

in 14 days at 20¢XC, white, flat, scanty, yielding a brown

pigment at the inoculation point, slowly diffusing to the

colony margins with age. Conidiophores arising from the

colony surface, cylindrical, septate, smooth, 100-200 ¡Ñ

10-15 £gm, hyaline, unbranched, each producing a phialide;

phialides cylindrical, 35-50 £gm, with conspicuous

periclinal thickening at tip, collarette, unflared. Conidia

hyaline, fusiform, 3- to 5-septate, 110-140 ¡Ñ 25-29 £gm,

smooth to slight roughened.

Specimens examined. Taipei County, Shihding, Huang-

di-dian north peak trail, on bark of Ficus fistulosa Reinw.

ex Bl., 27 Nov 2003, Guu, J.-R. 92112702. Taipei County,

Jingtung historical trail, on bark, 21 Dec 2003, Guu,

J.-R. 92122101 (cultured). Nantou County, Luku, Shitou,

on stem of Brugmansia suaveolens (Willd.) Bercht. &

Presl, 19 Apr 2003, Ju, Y.-M. 92041904 (cultured). Taipei

County, Hsintien, Ta-tung-shan, on bark of Syzygium

samarangense (Bl.) Merr. & Perry, 9 Oct 2004, Guu, J.-R.

93100904.

Notes. This species is easily recognized by its ovoidal

perithecia, cylindrical ascospores, and the Antipodium

anamorph. The anamorph that we obtained is typical for

A. spectabile as described in Rossman (1983). Perithecia

of this fungus are frequently mixed with those of H .

haematococca on the same substrata. The teleomorph

was produced on PDA in those cultures initiated from

specimen 92122101.

Acknowledgments. This study was supported by National

Science Council of R.O.C. Grant NSC 94-2621-B-001-001

to Y.-M. J. We highly appreciate two anonymous reviewers

for critically reviewing the manuscript and providing

invaluable suggestions.

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