Revision of Hendersonula

Myca/. Res. 93 (4): 466-488 (1989) Printed in Great Britain

Revision of Hendersonula

B. C. SUTTON AND B. J. DYKO"

CAB Intemational Mycological Institute, Ferry Lane, Kew, Richmond, Surrey TW9 3AF

Revision of Hendersonula. Mycological Research 93 (4): 466-488 (1989).

The history of Hendersonula is briefly reviewed and three species are accepted. Fifteen are of uncertain or dubious identity, one isreferred each to Fusicoccum, Sphaeropsis, Dothidella and Hendersoniopsis, and two to Seimatosporium. New combinations proposedinclude Hendersonula symploci (Berk. & Broome) comb.nov., Dichomera rhamnicola (Cooke) comb.nov., Dichomera conglobata (Sacc.)comb.nov., Torosporiopsis macrosperma (Cavara) comb.nov. and Stilbospora meridionalis (D. Sacc.) comb.nov. Dichomera neorhamni

nom.nov. is proposed for Camarosporium rhamni and the new generic name Nattrassia, type species N. mangiferae (H. & P. Syd.)comb.nov. introduced for the fungus formerly known as HendersonlAla toruloidea. The synanamorph of N. mangiferae is namedScytalidium dimidiatum (Penz.) comb.nov.

Key words: Nattrassia, Hendersonula, Systematics, Mycoparasites.

466

The generic name Hendersonula was introduced by Spegazzini(1880). At least 32 species have been described in the genusso far and these are augmented by the occasional use of thegeneric name in unpublished herbarium annotations. In termsof distribution, frequency and economics the most importantspecies is H. toruloidea Nattrass. This is a polymorphic fungus,readily cultured in its characteristic Scytalidium arthric stateyet also sometimes producing pycnidial conidiomata undersuitable conditions. It is frequently implicated in plant andhuman diseases and has a wide geographical distributionincluding Africa, Asia, North, Central and South America,Australasia and Europe (Punithalingam & Waterston, 1970 b;herb. IMI unpublished records). The nature of H. toruloidea invivo and in vitro is at variance with the type species, H.australis, which was originally described as a stromaticcoelomycete on Solanum leaves and subsequently found byPetrak (1951) to be a mycoparasite on Dothidella australisSpeg. (syn. Placostroma australe (Speg.) Theissen & Syd.). Sinceonly one of the remaining Hendersonula species was describedas a mycoparasite and Petrak (1949) considered the genus tobe heterogeneous, a more detailed revision is desirable.

Hendersonula was described by Spegazzini (1880) with asingle species, H. australis Speg. on living leaves of Solanumboerhaviifolium from Argentina. The generic diagnosis gaveconidiomata as foliicolous, stromatic, immersed, circular toirregular, inside white, black with little or barely markedpunctiform ostioles, and spores produced in locules, ellipsoid,

• National Science Foundation Post-Doctoral Research Fellow, eM!,1977-8.

multiseptate and coloured. The relationship of the genus toknown members of the group was not discussed. The specieswas provided with a slightly more detailed description, thelocules referred to as dothideaceous and crowded togetherand minute (80-90 !-lm) and conidia as elliptical or fusoid, bothends more or less rounded, 2-3 septate, constricted at thesepta, dilutely olivaceous, 14-20 x 4-5 !-lm. Conidia spreadingfrom the stromata were referred to as powdery olivaceous.

Saccardo (1884) accepted the genus in the Sphaeropsideae,Sphaerioideae, Phragmosporae, thought it to be a HendersoniaSacCo with dothideaceous stromata, and added a secondspecies, Hendersonula morbosa SacCo This was interpreted as thepycnidial anamorph of Plowrightia morbosa Schwein. (syn.Dibotryon morbosum (Schwein.) Theissen). The classic Europeanfloras by Allescher (1903), Diedicke (1914) and Migula (1921)

accepted the interpretation by Saccardo (1884), by which timethe number of species assigned to the genus had risen to ten.Stevens (1913, 1919) included H. morbosa in his account offungi involved in plant disease. In the following 60 years thetotal species included in Hendersonula rose to 32 and there hasbeen a parallel re-evaluation of the genus and its status,notably in a series of papers by Petrak. Clements & Shear(1931) again placed Hendersonula in the Phomales, Phomaceae(Sphaerioidaceae), Phaeophragmiae, a position scarely advanced from that of Saccardo (1884). Hahnel (1923)incorporated the genus in his radical reclassification ofDeuteromycetes but proferred no revision.

Petrak & Sydow (1935) redescribed H. australis inconsiderable detail from the type collection and concludedthat the genus is like Camarosporium Schulz. with a strongly

B. C. Sutton and B. ]. Dyko

developed stroma but lacking longitudinal septa in the conidiaor as a Oichomera Cooke with phragmospores. It wasmaintained as distinct yet heterogeneous. In describingHendersonula astragalina Petrak (1949) gave a resume ofseveral species which had been described in the genus. Hesuggested that (1) H. nlorbosa. interpreted as the anamorph ofOibotryon morbosum, should not be included with H. australis;(2) H. phyllachoroides SacCo could not be accurately placed fromthe original description; (3) H. macrosperma Cavara wasgenerically distinct from Hendersonula; (4) H. cerberae Tassi isnot typical of the genus; (5) H. bofryosphaerioides Bres. is anabnormal and poorly developed form of Dichomera saubinetii(Mont.) Cooke, as is H. conglobata (Sacc.) v. Hahnel anabnormal and poorly developed form of Dichomera; (6) H. piniDied. is referred to Macrophoma pinea (Desm.) Petrak &Sydow; (7) H. leptosphaerioides Fragoso is distinct from thetype species of the genus and H. mori Sacco & Vogl. is similar;(8) H. [rucficola Syd. has large conidia formed in pycnidia andcannot be retained in Hendersonula. Later, Petrak (1951) foundthat H. australis, rather than being a foliicolous pathogen, wasin fact a mycoparasite, the conidiomata immersed in stromatictissue of Oothidella australis. Because he thought the originalaccount of the species as having dothideoid stromata waserroneous he referred the genus to Hendersonia Berk. Since thebinomial H. australis was already preoccupied by Hendersoniaaustralis Tassi he introduced the nomen novum, H. dothidellaePetrak. Since according to his interpretation Hendersonula wasunavailable for stromatic ostiolate coelomycetes with septate,olivaceous conidia he removed H. astragalina to a newlydescribed genus, Hendersonulina Petrak. Sutton (1980) brieflyreviewed the history of Hendersonula, illustrated the typespecies, and concluded that it probably contains only twOspecies and that H. toruloidea should be accommodated in aseparate genus.

HendersonuIa Spegazzini. An. Soc. Cienc. Arg. 10: 36(1880).

Mycoparasitic, fruiting within ascomata. Mycelium immersed,branched, septate, hyaline to pale brown. Conidiomataeustromatic and multilocular or pycnidial and unilocular,immersed in or amongst host fungal tissue, brown; dehiscenceostiolar or by breakdown of overlying wall. Conidiophoresabsent. Conidiogenous cells discrete, determinate or indeterminate, hyaline, smooth, ampulliform, doliiform or cylindrical.proliferating enteroblastically with conidia seceding at thesame level or at successively higher levels; periclinal thickeningdistinct or not, collareUe absent Conidia holoblastic, palebrown, smooth or verruculose, with prominent transverseeusepta, cylindrical to fusiform, eguttulate, occasionally witha mucilaginous apical appendage; secession schizolytic.

Type species: H. australis Speg.

Key to accepted Hendersonula species

467

Hendersonula australis Spegazzini, Ann. Soc. Cien. Argent.10: 36 (1880). (Fig. 1)

Hendersonia dothidellae Petrak, Sydowia 5: 421 (1951).

Fruiting below the stroma of Dothidella australis Speg., whichin tum is fruiting on Solanum boerhavii[olium. Myceliumimmersed, branched, septate, hyaline. Conidiomata eustromatic,subcuticular, becoming epidermal. irregular in shape butgenerally circular in outline, 2-2'5 mm diam, becomingerurnpent especially at the periphery, irregularly pulvinate,carbonaceous, blackish brown, irregularly multilocular; wallsof 5-9 cell layers, up to 50 101m thick, composed of dark brown,thick-walled textUTa angularis, inner wall hyaline to pale brown,of thin-walled textura angularis. Ostiole absent; opening byirregular breakdown of the upper wall. Conidiophores absent.Conidiogenous cells cylindrical. percurrently proliferating, indeterminate, discrete, hyaline, smooth, 8-13 x 4-4'5 101m. Conidiaholoblastic, cylindrical to fusiform, pale brown, verruculose,thin-walled, 1-3 euseptate, septa thick and prominent, apexobtuse, base truncate, eguHulate, 8-20 x 4-5 \-1m.

The conidiomata of H. australis were found below thestromata of Dothidella australis; the stromata of the two fungiare quite distinct and can readily be distinguished. Neither ascinor spores of D. australis remained in the material which wasexamined. Often if a pycnidial anamorph occurs with anascogenous teleomorph, they do so in the same stromatictissue side by side in separate locules. Muller & V. Arx (1962)in a treatment of the didymosporous pyrenomycetes did notreport any anamorphic state for D. australis. Either H. australisis a secondary invader of an old fungal stroma of aDothideaceous fungus or it is a mycoparasite on D. australis.No colledions other than the holotype have been examinedso resolution of this problem must await additional collectionsand data.

Originally described as a stromatic coelomycete on leavesof Solanum by Spegazzini (1880). Petrak (1951) found theconidiomata of this fungus to be immersed in the stromatictissue of Dothidella australis Speg. (Placostroma australe ($peg.)Theissen & Syd.) and placed Hendersonula in synonymy withHendersonia Berk. because he regarded the original account ofthe fungus by Spegazzini to be erroneous. As the transfer toHendersonia would have created a later homonym with H.australis Tassi (1899) he proposed the new name Hendersoniadothidellae. Hendersonia Berk., however, is a nomen rejiciendumfor Stagonospora (Sacc.) Sacco and has holoblastic, hyalineseptate conidia formed in pycnidial conidiomata. Hendersonulaaustralis is taxonomically quite distind.

Specimens examined: On Dothidella australis (Placostroma australe(Speg.) Theiss &; Syd.), on leaves of Solanum boerhaviifolium Sndtn.,Boca del Rachuelo, Buenos Aires, Argentina, 10 July 1880. C.Spegazzini, Dec. Myc. Argent. no. 42, holotype, LPS.

H. australis2

1. Conidiomata eustromatic; conidia 1-3 septate, cylindrical, 8-20 x 4-5 \.lm, verruculose1. Conidiomata pycnidial; conidia 0-3 septate, fusifonn .

2. Conidiogenous cells 'phialidic'; conidia 16-23 x 4-4'5 \.lm. smooth, sometimes with a mucilaginous apical appendage.. . . H. monochaetiellae

2. COnidiogenous cells'annellidic'; conidia 17-22 x 4-5 \.lID, verruculose, with no appendage . H. symploci

31 MYC 93


Fig. 1. Hendersonula australis. A. Conidia; B. conidiogenous cells and developing conidia; C. vertical section of a conidioma belowstroma of Dothidella australis. Reproduced by pennission of CAB International from The Coelomycetes: 348 (1980).

468

A

B

lOjlm

Hendersonula monochaetieIlae Tommerup & Langdon,Trans. Br. mycol. Soc. 65: 478 (1970). (Fig. 2)

Fruiting amongst stromatic tissue of Hypnotheca graminisTommerup and its anamorph Monochaetielliopsis themedae(Kamasw. & Sundar.) Sutton & di Cosmo. Mycelium immersed,branched, septate, hyaline to dark brown. Conidiomatapycnidial, unilocular, 130 I.lm diam x 60 I.lm deep; walls 2-4cell layers. 3-5 I.lm thick composed of a pale brown outerlayer somewhat darker around the ostiole and an inner hyalinelayer. Ostiole central to each pycnidium. circular. Conidiophoresabsent. Conidiogenous cells ampulliform, doliiform, more rarelylageniform, not percurrently proliferating, hyaline, smooth,discrete, determinate, arising from the cells lining the wall,5'5-8 x 4-7 \-lm. Conidia holoblastic, fusiform, pale brown,smooth. 0-3 euseptate. apex conic to long conic, sometimeswith an apical cap-like mucilaginous appendage, base truncate,16-23 x 4-4'5 \-lID.

Tommerup & Langdon (1970) described H. monochaetiellaeas a mycoparasite of Hypnotheca graminis and its anamorphfrom leaves of Heteropogon contortus and Themeda australis.Hyphae of H. monochaetiellae ramify between cells of H.graminis and conidiomata are formed at the sites of infectionof H. graminis. Penetration of infected leaves by H.monochaetiellae is through splits in the epidermis whereascocarps or acervular conidiomata of H. graminis are found.

The same reservations concerning Hendersonula as anappropriate genus for H. symploci (p. 468) apply to this speciesbecause conidiomata are again pycnidial and ostiolate.

Hendersonula monochaetiellae is distinct from H. australis andH. symploci on account of the conidial shape which is much

more fusiform and the fact that occasionally conidia bear anapical mucilaginous appendage. Another difference is thatconidia in H. monochaetiellae are smooth rather than verruculoseas in the other two species.

Conidial morphology invites close comparison withAscochyta paspali (H. Sydow) Punith. (Buchanan, 1987). Theshape is very similar but conidia are somewhat broader andthere is no reported apical appendage in this species. A. paspaliis a well-known pathogen of Paspalum (Buchanan, 1984 a, b)and there are no reports in the literature concerning itsassociation with or hyperparasitism of other fungi likeHypnotheca or Monochaetiellopsis. However, H. monochaetiellae grew and sporulated readily on common laboratorymedia which indicates the species is not obligately mycoparasitic. It would be interesting to conduct cross inoculationstudies with H. monochaetiellae on Paspalum species todetermine if it is capable of inducing a similar diseasecondition as does A. paspali.

Specimen examined: On Monochaetiellopsis themedae on Heteropogoncontortus, Esk. Queensland, Australia. 16 March 1967, R. F. Langdon,BRIV 2413. IMI 223866. holotype.

Hendersonula symploci (Berk. & Broome) Sutton & Dyko,comb.nov. (Fig. 3)

Hendersonia symploci Berk. & Broome, ]. Linn. Soc. 14: 88(1873).

Hendersonula yakushimensis Kobayashi, Mem. Nat. Mus.Tokyo 9: 91 (1976).

Fruiting in stroma of Dermatodothis sp. (Trematosphaeriayakushimensis Kobayashi, 1976) and Dermatodothis zeylanicaH. Sydow (1923) growing on 5ymplocos spp. Mycelium

B. C. Sutton and B. J. Dyko

Fig. 2. Hendersonula monocMetiellae. A. Conidia; B, conidiogenous cells and develuping conidia; C vertical section of a conidiomaamongst stromatic tissue of Hypnotheca graminis.

469

A

B 1OJ.Lffi

immersed, branched, septate, hyaline to pale brown. Conidiomata pycnidial. in locule of ascomycetous host, unilocular,52-100 lJ.m diam; walls 3-7 cell layers, 9-20 lJ.m, thick,composed pale brown, irregularly thickened thin-walled textumangularis. Ostiole central to each locule, circular. Conidiophoresabsent. Conidiogenous cells lageniform, with 1-2 percurrentproliferations, hyaline, smooth, discrete, indeterminate, arising

from cells lining locule, 5-12 x 2-4 lJ.m. Conidia holoblastic,fusiform, medium brown, verruculose, 3 euseptate, apexobtuse to conic, base truncate, 17-22 x 4-5 lJ.m.

This taxon does not fit entirely satisfactorily in Hendersonulabecause the conidiomata are pycnidial and ostiolate. In thetype species the conidiomata are eustromatic with no definite

31-1

Revision of Hendersonula 470

Fig 3•. Hendersonula symploci A, Conidia; B, conidiogenous cells and developing conidia; C vertical section of conidioma in stroma ofDermatodothis zeylanica.

A

10 J.lITl

B

method of dehiscence. However, the conidium shape, colour,septation and ontogeny are very similar or the same. It ispossible the differences in conidiomatal structure are the resultof a difference in host parasitism. In the absence of additionalinformation, it seems best to maintain Hendersonia symploci inHendersonula for the moment.

Three fungi are represented on the type specimen,Dermatodothis zeylanica H. Sydow, Hendersonula symplociwhich parasitizes it and Scolecosporiella varians (Sacc.) Dyko &

Sutton. Petch (1924) identified two species on the typematerial in his possession, an ascomycete identified as D.zeylanica and a coelomycete H. symploci the diagnosis of whichwas based on a combination of the two. Berkeley's specimenno. 455 (herb. K), however, speCifically notes in pencil 'thesmall spots x 0'0012 long' for H. symploci. These measurements refer to the parasitized Dermatodothis, although Berkeleywas unaware of the fungicolous nature of H. symploci. Sydow

(1923) noted a conidial state in his description of D. zeylanicabased on Petch specimens nos. 6400, 6005 and 6605 (herb. K)but these conidia are much shorter than those reported byBerkeley & Broome (1873). These conidia were also observedin this study but there is insufficient data to describe thestate.

Kobayashi (1976) described Trematosphaeria yakushimensisKobay. with an associated coelomycete state, Hendersonulayakushimensis Kobay., on living leaves of Symplocos theophrastifolia. The author describes teleomorph and anamorphstates occurring together in the ascostroma in separate locules.Examination of the type material shows a pycnidial funguswithin the locular cavity of the stroma of an ascomycete. Theascomycete is referable to Dermatodothis but as no asci orascospores remain in the material received for study, it is notpossible to give a specific epithet for it. This Dermatodothis,however, was not conspecific with D. zeylanica. The pycnidial


fungus is mycoparasitic on the Dermatodothis and is referrableto H. symploci.

Specimens examined: On Dermatodothis sp. on leaves of Symplocos

/heophras/ijolia, Yaku Island, Kayoshima PreL Japan, 1 July 1975,T. Kobayashi, no. 4450, Lab., TFM; on Dermatodo/his uyIanica on S.ob/usa, Nuwara Eliya, Sri Lanka, 7000 ft, no. 592, Hendersonia

symploci, holotype, ex herb. Berk., K; on D. uylanica on S. obtusa,

Central Province, Sri Lanka, 5000 ft, no. 455, ex Herb. Berk., K; onD. uylanica on S. obtusa, Hakgala, Sri Lanka, 3 March 1922, T. Petch,no. 404, Dermatodothis uyIanica, holotype, K; on D. uyIanica on S.latiflora, Hakgala, Sri Lanka, April 1919, T. Petch, no. 6005, K; on D.uylanica on S. elegans, Hakgala, Sri Lanka, April 1917, T. Petch, no.5232, K; on D. uylanica on S. ob/usa, Hakgala, Sri Lanka, Feb. 1923,T. Petch, no. 6605 'pycnidial', K.

DOUBTFUL AND REJECTED SPECIES

Hendersonula aristidae

Dothidella aristidae (Schwein.) Ell. & Everh., North AmericanPyrenomycetes: 607 (1892).

Sphaeria aristidae Schwein., Syn. Fung. Amer. bor.: 1471(1831).

Phyllachora aristidae (Schwein.) Sacc., Syll. fungo 2: 623(1883).

Hendersonula aristidae (Schwein.) Ell. in Tracy & Earle, Miss.Agr. Mech. ColI. Exp. St. Bull. 34: 110 (1895).

Euryachora? aristidae (Schwein.) Theiss. & Syd., Annis mycol.13: 367 (1915).

This was interpreted as a stromatic coelomycete by Ellis inTracy & Earle (1895) who described conidia as oblong,12-14 x 3'5-14 \.lm, at first hyaline and I-septate, becomingbrown and 3-septate. Sprague (1950) described the disease ofsalt-grass as Char Spot noting that two fungi are involved, D.aristidae which appears to be virtually saprophytic andSeptogloeum oxysporum Bomm., Rouss & Sacco (which is a latername for Cheilaria agrostis Lib.) which is actively parasitic.

Hendersonula astragalina

Hendersonulina astragalina (Petrak) Petrak, Sydowia 5: 421(1951).

Hendersonula astragalina Petrak, Sydowia 3: 322 (1949).

Petrak (195 I) described this species from Astragalus sp.collected in Iran and cited specimen no. 7165. The fungus wassaid to produce erumpent more or less aggregated, multilocular, ostiolate stromata 1-3 mm diam, composed of brownpseudoparenchyma. Pycnidia were described as denselybotryose-congested. Conidia were reported as numerous,whei'l young more or less globose to Widely ovoid, continuousor with a median septum, later oblong to cylindrical thendavateto pyriform, straight, 2-3-septate and 8-16 x 4-6 \lmlong. .

The holotype from Petrak's herbarium (W) has beenexamined but no fungus corresponding with the account ofmature conidia has been found on it. All the material seems torepresent the immature form with conidia 5 x 1'5 \lm, verypale brown and occasionally with a single median septum.There is some difficulty in interpreting this observation.

471

Possibly this fungus is indeed the immature form of H.astragalina and later conidia formed in similar, thick-walledsclerenchymatous pycnidia are of the dimensions quoted byPetrak. It is also possible that there are two separate unrelatedspecies. Whatever the explanation, the mature conidia aremost unlikely to belong to a Hendersonula species sensustricto.

Petrak (1951) proposed the generic name Hendersonulina toaccommodate H. astragalina when he reduced Hendersonula tosynonymy with Hendersonia Sacc. Unfortunately, Hendersonulina is a later homonym of Hendersonulina Tassi (1902) andwould have to be replaced if material becomes available toresolve the taxonomic problem posed by Petrak's type andthe genus is maintained as distinct. Hendersonula leptosphaerioides G. Frag. (see p. 476) was also described fromAstragalus but the considerable difference in conidial sizeindicates that they are separate species.

Specimen examined: On Astragalus sp., ProVo Mazanderan, distr.Kudjur, Ulodi Mtn 3200 m, 9 Aug. 1948, K. H. & F. Rechinger, No.7165, ex herb. W.

Hendersonula botryosphaerioides

Dichomera rhamnicola (Cooke) Sutton & Dyko, comb.nov. (Fig. 4)

Hendersonia rhamnicola Cooke, Nuovo Giorn. Bot. Ital. 10: 2 I(1878).

Hendersonula botryosphaerioides Bresad., Verhandl. Zool. Bot.Gesell. Wien: 618 (1905).

Camarosporium frangulae Petrak, Annis mycol. 22: 32 (1924).

Fruiting on dead twigs of Rhamnus frangula. Myceliumimmersed, brown, branched, septate. Conidiomata eustromatic,immersed, subperidermal, globose to irregular in outline, darkbrown to black, up to 1'5 mm diam; wall composed of 8-20cell layers, 80-150~ thick, outer layers of dark brown, thickwalled textura angularis, inner layers of thin-walled, hyalinetextura angularis. Ostioles single to multiple, circular, papillate.Conidiophores absent. Conidiogenous cells doliiform to cylindrical, percurrently proliferating with up to 3 annellations,discrete, indeterminate, smooth, hyaline, arising from innercells of locule, 5'5-16 x 2-4'5 !Jm. Conidia holoblastic, ovoidto elliptical, brown, muriform with 3-6 transverse and 1-2longitudinal or oblique eusepta, sometimes slightly constrictedat septa, smooth, base truncate, apex obtuse,13-22'5 x 6'5-8'5 \lm.

The type material of H. botryosphaerioides is ample and ingood condition but contains only conidia in the earlier stagesof development; they are yellowish in colour and only show1-3 longitudinal septa. Comparison with the type specimensof C. frangulae and H. rhamnicola. which contain mature andimmature conidia show them to be conspecific with H.botryosphaerioides.

Petrak (1949) considered H. botryosphaerioides to be a poorand abnormal specimen of Dichomera saubinetii (Mont.)Cooke; however, study of the type specimen of Hendersoniasaubinetii Mont. shows them not to be conspecific. A modemdescription of D. saubinetii is given by Sutton (1980). Grove


Fig. 4. Dichomera rhamnicola. A, Conidia; B, conidiogenous cells and developing conidia.

B

10 JlIl1

A

(1937) considered H. ramnicola to be a later synonym ofHendersonia mammillana (Fr.) Currey based on Sphaeriamammillana Fr. Study of the relevant material in the Friesherbarium (UPS) shows 5. mammillana to be an ascomycete,Clypeosphaeria mammillana (Fr.) Lambotte. Grove also suggested that 5taurosphaeria rhamni Westend. (Dichomera rhamni(Westend.) Sacc.) was a synonym of H. rhamnicola. Westendrop's (1859) description and his drawings strongly suggestthat the fungus he described as 5. rhamni is Dichomerasaubinefii rather than D. rhamnicola. Dichomera saubinefii can bedistinguished from other Dichomera species occurring onRhamnus [rangula by the spherical to subglobose, brownconidia which are often triradiately to cruciately septate and10-12 ~m diam.

It also seemed possible that H. bofryosphaerioides might besynonymous with Camarosporium rhamni Allesch. as theyoccur on the same host and have an apparently similar conidialsize. The holotype of C. rhamni shows the two taxa to be quitedistinct, the most notable differences being the lineararrangement of the conidiomata, the greater number ofsecondary transverse and oblique septa, and the greater widthof the conidia found in C. rhamni.

Several genera of coelomycetes produce muriform conidia.Camarosporium Schulz. differs from Dichomera in conidiomatabeing pycnidial rather than eustromatic. Dichomera hassometimes been interpreted as having 'phialidic' (sensu Sutton,1980) conidial ontogeny (Morgan-Jones, Nag Raj & Kendrick,1972) whereas Camarosporium has been interpreted asholoblastic. In practice, however, pycnidial conidiomatalforms intergrade into stromatic forms and conidiogenesis inboth genera seems to be holoblastic with the facility topercurrently proliferate with conidia formed at successivelyhigher levels. Camarographium Bubak also produces conidiomata which are difficult to interpret, appearing as separatepycnidia or as pycnidia arranged in a linear stroma. Theconidia, however, are produced from 'phialides' and aredistoseptate with a thick peridinal wall. 5figmella Lev. producespycnidial conidiomata and holoblastic conidia but the

conidiomatal wall is thinner than is normally seen inCamarosporium. The conidiogenous cells of Sfigmella tend tobe cylindrical but that character overlaps with the presentdiversity found in Camarosporium, and the conidia of 5tigmellaare very thick-walled with reduced cell contents but stilleuseptate.

Camarosporellum Tassi differs from 5figmella and Camarosporium only is that its conidia are distoseptate. Dichomeracontains 40 taxa and requires revision. Camarographium.5figmella and Camarosporellum may require reconsiderationbut this and the revision of Dichomera can only be done in thebroader context of an extensive revision of Camarosporiumwhich now contains over 400 taxa.

The most common teleomorphs for Camarosporium andDichomera species are referable to Cucurbifaria S. F. Gray. Thisstromatic ascomycete produces muriform ascospores of similarsize and shape to the conidia. As the conidial and ascomatalstates are occasionally encountered in the same stroma, somecare must be used in observation. Additional information onCucurbifaria may be found in Mirza (1968) and Welch(1926).

Specimens examined: On Rhamnus frangula twigs, Hampstead,Middlesex, UK, Jan. 1866, M. C. Cooke, holotype of Hendersoniarhamnicola, K; on R. frangula, WesHalen, Komischer Wald, beiBottrop, Austria, April. 1923, Rupprecht, as Camarosporium frangulaePetrak, isotype IMI 24167; on R. [rangula, Margo, Italy, Oct. 1901,Bresadola, holotype of Hendersonula botryosphaerioides, S; on R.frangula, Trento, Italy, 14 April 1904, Bresadola, PAD no. 4249; onR. frangula, Sonntagsberg, Austria, 23 Feb. 1908, P. Strasser, S; onR. frangula, Mahr-Weisskirchen, Austria, April 1936, F. Petrak asDichomera rhamn; (West.) Sacc., K.

Dichomera neorharnni Sutton & Dyko, nom. nov.(Fig.5A)

Camarosporium rhamni Allesch., Hedwigia 33: 72 (1894), nonD. rhamni (Westend.) Sacc., Syll. fungo 3: 471 (1883).

Fruiting on dead twigs of Rhamnus [rangula. Myceliumimmersed, hyaline to brown, branched, septate. Conidiomata


Fig. 5. A, Dichamera nearhamni, conidia; B, Dichamera cang/abata, conidia.

473

B

25J.llllA

eustromatic, immersed, subperidermal, globose to irregular inoutline, dark brown to black, up to 0'8 mm long and 0'5 mmwide; wall composed of 5-10 cell layers, 62-190 ~m thick,outer layers of dark brown, thick-walled textura angularis, innerlayers of thin-walled, hyaline textura angularis. Ostioles singleto multiple, circular, papillate. Conidiophores absent. Conidiogenous cells doliiforrn to cylindrical, discrete, smooth, hyaline,arising from the inner cells of the locule, 5-7 x 2-2'5 ~m.

Conidia holoblastic, oval to obovate, brown, muriform with3-4 transverse and 1-8 longitudinal or oblique eusepta,sometimes slightly constricted at septa, smooth, base truncate,apex obtuse, 18-29 x 10-14 ~m.

This taxon, while occurring on the same host as D.rhamnicola, has wider and slightly longer conidia with moreoblique septa. The two species have slightly different growthhabits on the host; D. neorhamni tends to produce more linearconidiomata while D. rhamnicola produces conidiomata whichtend to be more circular in outline. Many of the conidiomataon the type material were effete, containing only a few matureconidia.

Allescher (1894) stated that C. rhamni was the pycnidialstate of Cucurbitaria rhamni Fuckel. Fuckel (1870) describedDiplodia frangulae which he reported to be the pycnidial stateof C. rhamni and cited his exsiccatus Fungi rhenani no. 1955Diplodia frangulae. Later, he cited the same exsiccatus andreported muriform septation present in the conidia (Fuckel,

1871). Material of C. rhamni and D. frangulae includingFuckel's exsiccatus no. 1955 was examined and the teleomorphand only one conidial state, referable to Diplodia, were found.This Diplodia is, in fact, identical in size, shape and conidialontogeny to Diplodia mutila Fr.; it differs only in havingslightly thinner walls and lacks a prominent vacuole andgranular cytoplasm.

The ascospore of Cucurbitaria rhamni are nearly the samelength, width (20'5-27 x 8'5-10'8 ~m) and shape as theconidia of D. neorhamni. The difference is that the ascosporesshow a constriction in the centre dividing the spore into twonearly equal halves. The spores examined were 5-7transversely septate and tapered more toward the apices thanis seen in conidia of D. rhamni.

Specimens examined: On Rhamnus frangula, Munchen waldung nearSolin, Germany, 23 May 1883, A. Allescher, holotype of Camarasparium rhamni Allesch., ex Pilzherb. Allescher, M.

Hendersonula cerberaeHendersonula cerberae F. Tassi, Rev. Myco/. 18: 170

(1896).

Illustrations: Allescher (1901).

Tassi (1896) described his taxon as

stromate sub-corticali, confluendo elongato, centro, plurilocellato;loculis globosis, intus albicantibus, 100-120 ~m aggregatis, apice


poro pertusis; sporulis anguste ellipticis, vel sub-clavatis, initiohyalinis vel chlorinis, stipitato-fasciculatis, conhnuis, endo-plasmagranuloso instrudis dein 2-septatis, ad septa non, vel raro constridis,fuligineis, 12-16 x 4'5-5 ~m.

Type material was located at SIENA and consists of fivesmall twigs of Tanghinia venenifera from India, 3-9'5 em, splitinto quarters. The conidiomata are immersed in the bark,eustromatic, 1'5-1 mm diam, not infrequently linearly arranged and with simple conidial locules and ascomatal initialsdeveloping in the upper half of the stroma. The stromaconsists of thick-walled, brown textura angularis which becomesthinner-walled and hyaline only very close to the regions ofconidiogenesis or ascomatal formation. Ostioles are not clearlydefined. The lageniform to cylindrical conidiogenous cells,8'5-12)( 2-3 i-\m, arise directly from the inner wall of texturaangularis or from short conidiophores which may be branched.Conidia are produced holoblastically, elliptical in shape,hyaline, with granular cytoplasm, aseptate and 13-18 x4-5'5 i-\m.

This fungus is referable to Fusicoccum Corda but as Sutton(1980) notes, although 200 taxa are currently ascribed to it,the genus is not currently accepted and is in need ofconsiderable revision.

Also present on the material are conidia of Lasiosphaeriatheobromae (Pat.) Griff. & Maubl. (Botryodiplodia theohromaePat.), ascospores of a taxon perhaps referable to DidymellaSacc, and the fusiform, 2-septate, brown conidia,12-16 x 4'5-5 i-\m, as described by Tassi which could not beidentified. It was not possible to associate any of these sporeswith a fruiting structure.

Petrak (1949) rejected H. cerberae as belonging toHendersonula on the basis of many locules opening through apore. His observations could not be confirmed but it issuggested that H. cerberae be considered of doubtful applicationas it is based on several discordant elements, the origins ofwhich cannot be ascertained.

Specimen examined: On branches of Tanghinia (Cerbera) oenenifera,. India, F. Tassi, SIENA, holotype.

HendersonulR dRlbergiRe

Hendersonula dalbergiae Ahmad & Arshad, Biologia, Lahore18: 118 (1972).

The species was described from branches of Dalhergia sissoofrom Pakistan (West Pakistan) as producing solitary togregarious, confluent, erumpent pycnidia up to 400 !-lm diam,composed of pseudoparenchymatous cells. The conidia weredescribed as yellow-brown, oblong-ellipsoid, 1-3 septate,15'5-20 x 7-8 !-lm.

No response to a request for type material deposited atLAH was received. The description is insufficient to enable anappropriate placement of this species to be made.

Hendersonula crRtaegi

Seimatosporium vaccinii (Fuckel) Eriksson, Svensk hot, Tidskr.68: 242 (1974).

Coryneum vaccinii Fuckel, ]b nassau ver. Naturk. 27-28: 81(1873).

474

Hendersonia fruetigena Sacco var. crataegi Allesch., Ber. Bayer.Bot. Gesell. 5: 20 (1895).

Hendersonula crataegi (Allesch.) V. Hahn., Ber. Deutsch. Bot.Gesell. 36: 315 (1918).

Coryneum microstietum Berk. & Broome var. staphyleae W. D.Davis, Phytopathology 21: 1172 (1931).

Coryneum rhododendri Schwein. var. fusoideum Dearness,Mycologia 33: 364 (1941), nom. inval., Art. 36.

Illustrations: Pirozynski & Shoemaker (1970), Sutton (1980).

Fruiting on stems of Vaccinium spp., Rhododendron spp.,5taphylea trifolia, Crataegus spp., and fruits of Crataegus spp.Mycelium immersed, hyaline to brown, branched, septate.Conidiomata acervular, immersed, subcuticular, becomingerumpent, circular in outline, 0-083 to 1'4 mm diam; wall 24cell layers, 5-17 !-lm thick, composed of hyaline to pale browntextura intricata. Ostiole absent, opening by rupture of overlyinghost tissue. Conidiophores cylindrical, septate, branched,hyaline, filiform, arising from conidiomatal cells, up to25 x 1'5-2. Conidiogenous cells cylindrical, filiform, percurrentlyproliferating, integrated or discrete, indeterminate, hyaline,0-3 annellations, 11-30 x 1'5-2 !-lm. Conidia holoblastic,fusiform, often with periclinal walls collapSing betweenthickened septa, 3 euseptate, smooth, septa darker thanpericlinal walls, upper three cells pale brown, basal cell subhyaline to pale brown, basal scar truncate, appendages absent,12-18 x 4'5-5'5 i-\ffi.

Hohnel (1918) had not seen Allescher's type material ofHendersonia fruetigena var. crataegi but he had examined afungus on Crataegus oryacanthus fruits from Austria which hethought to be identical with that described by Allescher. Heconsidered the taxon to be stromatic and accepted Allescher'staxon at species level and transferred the name to Hendersonula.Examination of Hohnel's Austrian material labelledHendersonula crataegi shows the fungus not to be congenericwith H. fruetigena var. crataegi Allesch. which is a synonym ofSeimatosporium vaccinii but identical with various forms ofHendersonula fruetigena (Sacc.) v, Hahn" a name which Hahnel(1920) later proposed.

Pirozynski & Shoemaker (1970) considered Coryneumrhododendri var. fusoideum to be a synonym of Seimatosporiumlichenicola. The conidia of 5. lichenicola, however, tend to bemore ellipSOidal in shape, slightly shorter in length and wider(13-15 x 5'5--6'5 !-lm). Coryneum rhododendri var. fusoideumhas a conidial shape and size identical with that of 5, vacciniiand does not belong with 5. lichenicola.

Seimatosporium Corda is an extremely variable and heterogeneous genus which is in considerable need of revision asmore species are known in culture. A review of the pertinentliterature may be found in'Sutton (1980) who provided shortdescriptions and a key to 38 species.

Specimens examined: On Rhododendron catawbiense, Roan Mountain,Tennessee, U.S.A., 19 June 1949, A.). Sharp, Flora of Tennessee,Fu,ngi, TENN 19447; on R. catawbiense, Roan Mountain, Carter Co.,Tennessee, U.S.A., 5 May 1934, A. J. Sharp, Flora of Tennessee,Fungi, holotype of Coryneum rhododendri var. fusoideum Dearn.,TENN 7321; on Vaccinium myrtillus. Neuchatel, Switzerland, Hieme,Morthier, Herbier Barbey-Boissier, Fuckel, Fungi rhenani ed. I, no.2849. Coryneum oaccinii Fuckel, G; on Vaccinium myrtillus, Neuchatel,


Switzerland, Morthier, Fuckel, Fungi rhenani no. 2405 Coryrzeumvaceinii Fuckel, IMI 74948; on Rhododendron sp., Andrews Bald, GreatSmoky Mountains National Park. Tennessee, U.S.A., 17 Oct 1940,E. K. Cash, TENN; on Staphylea trifolia. South Deerfield, Massachusetts, U.s.A., 10 May 1968, R. B. Livingston, IMI 181172; on S.trifolia, Amherst Massachusetts, U.s.A., W. H. Davis, authenticmaterial for Coryneum microstietum var. staphyleae W. H. Davis, IMI18II73; on Crataegus oryacanthus, Munich, Germany, May 1896,A. Allescher, holotype of Hendersonia fruetigena var. crataegi A1lescher,M; on Crataegus twigs, Magyarhon viranyabol, F. Hazslinszky, K.

Hendersonula conglohataDichomera conglobata (Sacc.) Sutton & Dyko, comb. nov.

(Fig.5B)

Sphaeropsis conglobata Sacc., Syll. fungo 3: 299 (1884).

Haplosporella conglobata (Sacc.) Allesch., in Rabenhorst, Kryptog.Flora Deutschl. 1. Abt. 7: 70 (1903).

Hendersonula conglobata (Sacc.) Hohn., in Strasser, Verhandl.Zool. bot. Gesell. Wien 55: 618 (1905).

Fruiting on twigs of Befula alba. Mycelium immersed, hyalineto brown, branched, septate. Conidiomafa eustromatic, immersed, subepidermal, black, globose to irregular, unilocular tomultilocular, up to 1'4 mm diam; wall 0-9 cell layers,35-56 ~m thick. outer wall composed of dark brown, thickwalled textura angularis, inner layers of hyaline thin-walledtextura angularis. Osfioles single to multiple, circular, papillate.Conidiophores absent. Conidiogenous cells doliiform, lageniformor cylindrical, percurrently proliferating, discrete, indeterminate, smooth, hyaline, arising from the inner cells of thelocules, 0-13 x 2'5-3'5 ~m. Conidia holoblastic, hyaline topale brown, ovoid, aseptate or with up to 4 transverse euseptaand/or 1-2 oblique eusepta, smooth, base truncate, apexobtuse, 13-22'5 x 5'5-11 ~m.

The conidia in this taxon are most unusual in appearance asthey are hyaline to pale brown and the septa are notinfrequently oblique. These characters usually indicate animmature or malformed Dichomera or Camarosporium andindeed, Petrak (1949) considered this fungus an abnonnalDichomera. Additional material was located and the same typeof conidium was found in these specimens, No Dichomera orCamarosporium has been previously reported from Betula albabut as this fungus appears to have some affiliation withDichomera, a new combination is proposed.

Saccardo (1884) proposed the name Sphaeropsis conglobafafor the supposed anamorph of Cucurbitula conglobata Fuckel(1870). Fuckel did not designate a type when he first providedthe description of the pycnidial and ascogenous states underthe name C. conglobafa. Later, Fucke! (1873) noted that he haddistributed material, from the originally described location, asFungi rhenani no, 2532 ed. I which contained the pycnidialfungus. Samples of this exsiccatus from herb. BM and Kcontained an ascomycete referable to Coniochaefa (Sacc.)Massee on a piece of white bark and Sphaeropsis conglobata ona piece of small twig. The specimen from herb. K is designatedlectotype of D. conglobata, since by inference it was thematerial (syntype) on which Saccardo based his description.

Hohnel (1918) noted C. conglobata was more appropriate toRosellinia de NoLor Coniochaeta than Curcurbitula and was not

475

the teleomorph of Sphaeropsis conglobata as Fuckel (1870) andSaccardo (1884) had supposed. Hohnel (1905, 1918), withoutseeing the type material, was certain he had collected S.conglobata and considered it to be a Hendersonula. He gaveDothidea virgultorum (Fr.) Winter ( == Anisogramma virgultorum(Fr.) Theissen & Sydow, fide Muller & V. Arx 1962) as thete!eomorph of 5. cong/obafa. Muller & V. Arx (1962) describedand illustrated the anamorph of A. virgultorum as havinghyaline, aseptate, ellipsoid conidia 12-15 x 4-6 ~m, apparently produced sympodially. This does not agree with S.conglobata. Hohne! (1918) placed Myxosporium lanceolum Sacco& Roum. forma befulae Fautrey and Dothiorella betulae (Preuss)Sacco in synonymy with H. conglobafa. Dofhiorella betulae andM. lanceolum f. betulae produce hyaline, aseptate, fusiformconidia which also do not agree with H. conglobata. Fries'5phaeria conglobata (Fries, 1823) could also be a possiblesynonym of the teleomorph and this possibility was noted byFuckel (1870) and Saccardo (1884). However, S. conglobata isnow known to be Melanomma pulvis-pyrius (Pers.) Fuckel(Chesters, 1938; Holm, 1957) producing an anamorph withhyaline, ellipsoid, aseptate conidia from long, cylindrical,percurrently proliferating phialides. This latter conidial statehas been referred to Aposphaeria agminalis SacCo (Chesters,1938) but needs reassessment in light of the redeSCription ofthe type species, A. pulviscula (Sacc.) Sacc., by Sutton(1980).

Specimens examined: On Betula alba, Frankensteiner Kopf, Austria,Fuckel, Fungi rhenani no. 2532, Cucurbitula conglobata Fckl. 1 Fung.stylosporifer, lectotype K, isolectotype BM; on B. alba, Konigstein,Gennany, 24 April 1905, W. Krieger, Krieger, Fungi saxonici no.1993. Haplosporella conglobata (Sacc.) Allesch., BM, S; on B. alba.Konigstein, Gennany, April 1905, W. Krieger, Kabat & Bubak, FungiImperfecti exsiccati no. 663. Haplosporella conglobata (Sacc.) A1lesch.,BM, S.

Hendersonula fructicolaHendersonula fructicola H. Sydow & Sydow, Annis mycal.

14: 369 (1916).

The fungus was described from mature fruit of a palm from MtUrdaneta, Mindanao, Philippines. The description was

Pycnidiis dense crustacee vel botryose aggregatis, superficialibus,globosis, 200-300 ~m diam, atris, glabris, coriaceo-carbonaceis,papillula minuta praeditis, basi hyphis pallide fuscidulis 4-5 ~ latisseptatis plus minus copiose cinctis, contextu parenchymatico excellulis 12-14 ~m diam. sordide fuscis vel olivaceo-fuscidulis interioribus minoribus et dilutioribus composito; sporulis oblongis, rectis,utrinque obtusis vel subinde basi attenuatis, 2-3 septatis, non velparum constrictis, diu hyalinis vel sub-hyalinis, tandem pallidesordideque fuscidulis, 50-70 x 14-18 IJm; sporophoris non visis.

The type material could not be located and without additionalinformation an appropriate taxonomic placement of thefungus cannot be made.

Hendersonula fructigenaHendersonula fructigena (Sacc.) V. Hohn., Hedwigia 62; 78

(1920).

Hendersonia frucfigena Sacc., Michelia 1: 212 (1878).

The original description of Hendersonia [rucfigena read


Peritheciis sparsis erumpentibus e globoso depressis, papillulatis,contextu minute parenchymatico dilute fuligineo; stylosporis cylindraceis initio stipitellatis, 23-25 x 6, 3- septatis, ad septa leniterconstrictis, minute guttulatis, dilute fuligineis, stipitibus stylosporabrevioribus. In ramis arescentibus Cerasi patavii, Julio, 1874, raro.

The species has appeared in various compilations oncoelomycetes (Diedicke, 1914; Allescher, 1903) but onlyv. Hahnel (1920) has suggested any revision. This was attempted in the context of his treatment of Hendersonula crataegi(Allesch.) v. Hahn., which was based on a misdetermination.Hahnel also did not see the type of H. fructigena and hisaccount is based on an Austrian collection in which conidiawere 1-2 celled and varied in size from 12 x 7 ~m to18-23 x 6-8 ~m. In addition to transferring Hendersoniafructigena to Hendersonula, v. Hahnel (1920) introduced fiveforms - f. crataegi Allesch. (it is not clear if this is based onHendersonia fructigena var. crataegi Allesch. or not), f. pruniv. Hahn., f. comi v. Hahn., f. sorbi v. Hahn. and f. cerasi v. Hahn.None of these names was accompanied by a description sothey are nomina nuda. In the absence of type material it is notpossible to determine either the generic relationships ofHendersonia fructigena or how the species relate to Hendersonulacrataegi which is shown here to be a synonym of Seimatosporiumvaccinii (Fuckel) Eriksson. From Saccardo's original descriptionit is strongly suspected that the species belongs inSeimatosporium and not either Hendersonia or Hendersonula.

Hendersonula konseiensis

Hendersonula konseiensis Kobayashi, Trans. mycol. Soc.Japan 4: 8 (1962).

This species was described as the anamorph of Cryptodiaporthekonseiensis Kobayashi (1962) but later, in a revision of JapaneseDiaporthaceae, C. konseiensis was reduced to synonymy withPseudovalsella thelebola (Fr.) v. Hahnel and H. konseiensis tosynonymy with H. thelebola (Sacc.) T. Kobayashi, which wasbased on Stilbospora thelebola. For further discussion seeHendersonula thelebola (p. 483).

Hendersonula leptosphaerioides

Hendersonula leptosphaerioides G. Frag., Mem. R. Acad.Cienc. Art. Barcelona 15 (17): 444 (1920), as 'leptosphaerioidea '.

The original account bore no accompanying illustration. Thespecies was described with .superficial, confluent, irregular,rough, black stromata, with minute locules and a punctiformostiole. Conidia were cylindrical fusoid, straight or slightlycurved, both ends attenuated, obtuse, 30-40 x 5-6 ~m, at firsthyaline then yellowish, 7-8 septate, often minutely guttulate,formed from short conidiophores. The host was given asAstragalus saxatilis ( = A. chlorocyaneus).

No material was available for study and it is not possiblefrom the generalized description to determine if the species iscorrectly placed in Hendersonula. It seems unlikely that it is thesame species as H. astragalina Petrak. also described from

476

Astragalus, because the conidial sizes and shapes are quitedifferent.

Hendersonula macrosperma

Toxosporiopsis macrosperma (Cavara) Sutton & Dyko,comb. nov. (Fig. 6)

Hendersonula macrosperma Cavara, Fungi Longobardiae Exsiccati,140 cum icon. (1891-1895).

Stilbospora salicella v. Hahn., nom. nov., Ost. bot. Zeitschr. 66:104 (1916), and Mitt. Bot. Inst. Tech. Hochsch. Wien 4: 96(1927), non Stilbospora macrosperma Pers.: Fr., Syst. Myc.3: 485 (1832); nec Stilbospora macrosperma Berk. &Broome, Hooker's J. Bot. 3: 320 (1851).

Fruiting in bark of Salix sp. and Robinia hispida. Myceliumimmersed, hyaline to brown, branched, septate. Conidiomataeustromatic, gregarious, immersed, subperidermal, irregularlymultilocular, becoming erumpent, 0'1-0'4 mm diam; wall 4-6cell layers, 4-6 ~m thick. composed of hyaline to pale brownthin-walled textura prismatica which becomes hyaline towardsconidiogenous regions. Ostiole absent, opening by rupture oflocular and host tissues. Paraphyses cylindrical becomingfiliform above, hyaline, simple, smooth, 1-4 septate, intermixed with conidiophores, 60-80 x I-I'5 ~m. Conidiophoresshort to absent, cylindrical, hyaline, smooth. Conidiogenouscells cylindrical to lageniform, discrete to integrated, determinate, hyaline, smooth, 3'5-6 x 3-3'5 ~m. Conidia holoblastic, clavate to fusiform, 3-euseptate, darker pigmentationabout septa, versicoloured with basal and apical cells palerbrown and verrucose and median cells brown, thick-walled,smooth, apex obtuse, base truncate with marginal frill,27-40 x 12'5-16 j..\m, developing in a gelatinous sheathwhich persists at maturity.

The conidia of this fungus are similar in size and shapeto Toxosporiopsis capitata Sutton & Sellar (1966)(29'5-37'5 x n'5-15 j..\m) but differ in several other features,Conidia in T. capitata are dark greyish brown with darkerbands of pigmentation at the septa, the basal and apical cellsare much smaller than the median cells, the apical cell tapersto a point often with a mucilagenous appendage attached andthe conidial walls are completely smooth.

The example of Cavara's exsiccatus no. 140 which wasexamined and which serve as the type for this taxon, aresomewhat immature. Most of the conidia lack the dark bandsof pigmentation at the septa and the thicker walls seen at alater stage of maturity.

Neohendersonia Petrak is similar to Toxosporiopsis. Neohendersonia produces stromatic conidiomata with holoblastic,cylindrical to fusiform or pyriform, versicoloured conidiaformed on small doliiform conidiogenous cells. It differs fromToxosporiopsis in that the conidia are euseptate, the septa lacka dark pigmented band and the conidiomata lack conidiophoresand paraphyses.

Specimens examined: On Salix bark, Italy, Cavara, Fungi LongobardiaeExsiccati no. 140. Hendersonula macrosperma nov. sp.. isotypes, K. B, L,S, BPI: on Robinia hispida, Tuskegee, Macon Co., Alabama, U.s.A.,26 May 1936, G. W. Carver no. 451, BPI.


Fig. 6. Toxosporiopsis macrosperma. A, Conidia; B, conidiogenous cells, developing conidia and paraphyses; C vertical section of aconidioma.

477

A

10 JllTI B

c

Hendersonula meridionalis

Stilbospora meridionalis (D. Sacc.) Sutton & Dyko, comb.nov. (Fig. 7)

Hendersonia meridionalis D. SacCo in Saccardo & Sydow, Syll.fung. 14: 956 (1899).

Hendersoniopsis meridionalis (D. Sacc.) V. Hohn., Mitt. Bot. Inst.Tech. Hachsch. Wien 5: 106 (1928).

Hendersonula meridionalis D. Sacc., Myc. Ital. nr. 302 (1898),nomen nudum.

Coryneopsis tamaricis Grove, ]. Bot. 70: 35 (1932).

Teleomorph: Leptasphaeria meridianalis D. Sacc., in Sacc., Syll.fung. 14: 563 (1899), fide Saccardo & Sydow (1899).

Fruiting on twigs of Tamarix gallica and T. africans. Conidiomataeustromatic, immersed, subperidermal. circular in outline,0'2-1'1 mm diam; wall 6-11 cell layers, 20-35 IJm thick,composed of brown thick-walled textura angularis. Ostioleabsent, opening by rupture of the overlying host tissue.Paraphyses filiform, hyaline, multi-septate, usually simple but

infrequently branched near base, 40-120 x 3 IJm, at base to1'5-2 IJm above. Conidiophores short to absent, cylindrical.hyaline, smooth. Conidiogenous cells cylindrical to lageniform,discrete, indeterminate, hyaline, arising from cells lining loculeor from conidiophores, 10-60 x 2'5-4 IJm. Conidia holoblastic,cylindrical to elliptical. brown to dark brown, 3 (4)-euseptate,thick-walled smooth, apex obtuse, base obtuse with truncatebasal scar, (21) 24-38 x 11-13 (16) IJm.

Paraphyses vary greatly in number and prominence betweencollections and sometimes may be overlooked in the presenceof very long conidiogenous cells. Ellis & Ellis (1985) suggestedthe species would be better accommodated in Stilbaspara.

Hendersania meridianalis was proposed by Saccardo inSaccardo & Sydow (1899) for a fungus occurring on Tamarixgallica and producing 3-septate brown spores, 30 x 15 IJm andfiliform 'basidia' of 30-60 x 2 IJm in length. Saccardo citedthe exsiccatus, D. Saccardo, Mycotheca italica no. 302.Leptasphaeria meridianalis D. Saccardo socia Hendersanula meridianalis D. Sacco The name Hendersanula meridianalis is a nomennudum as no description was provided. Hohnel (1928)


Fig. 7. Sti/bospora meridiona/is. A, Conidia; B, conidiogenolls cells, developing conidia and paraphyses; C vertical section of aconidioma.

478

A

10 tJffi

200 tJffi

suggested that H. meridionalis was congeneric with Hendersoniopsis lhelebola (Sacc.) v. Hohne! and made the appropriatecombination. The D. Saccardo exsiccatus, no. 302 was locatedin herb. K and it was found to contain only the conidial stateof a Bolryosphaeria. The exsiccatus, Savulescu, Herb. Mycologicum Romanicum fasc. 35, nr. 1740 Hendersonia meridionalis(herb. K, 1MI), however, does contain a fungus which closelyfits Saccardo's description. The holotype of Hendersonialamaricis Cooke (1885) described from Tamarix gallica wasfound to be a Dichomera species. Migula's (1921) use of H.lamaricis Cke is by his description and illustrations, amisapplication of Cooke's name. Later, Grove (1932) based hisname Coryneopsis tamaricis on Migula's description of H.lamaricis Cooke so it should be interpreted as a new species,attributable to Grove alone and not a combination based onH. lamaricis. Study of Grove's material labelled C. lamaricisshows the fungus to be consistent with the Savulescuspecimen. Grove (1932) thought he was making a newcombination so he did not deSignate a holotype but in herb.

K, one packet was annotated' Coryneopsis lamaricis Grove, sp.nov.' and this was the specimen first cited in his description;this specimen has been designated as the holotype.

It is clear from the original description by Saccardo inSaccardo & Sydow (1899) and subsequent accounts andillustrations by Grove (1932), Migula (1921) and Ellis & Ellis(1985) that the one species, referable to Slilbospora is involved.Additional examples of D. Saccardo, Mycolheca italica no. 302

should be examined to select a lectotype for the nameHendersonia meridionalis. Sutton (1975) reviewed the literaturefor Stilbospora while monographing the related genus,Coryneum Nees. Slegonsporium Corda is similar to Slilbospora,producing acervular conidiomata with paraphyses, septatebranched conidiophores and holoblastic percurrently proliferating conidiogenous cells. It differs from Slilbospora in thatit produces conidia with transverse and longitudinal distosepta.Coryneum Nees differs from both Slegonsporium and Slilbosporain the absence of paraphyses and it produces conidia which areonly transversely distoseptate.


Specimens examined: On twigs of Tamarix gallica, lasia, Romania, 22July, 1951, C. Sandu-viIle, Herb. Mycol. Romanicum, Dr T. R. Savulescu,

fase. 35, nr. 1740, Hendersonia meridionalis Saccardo, E, BM, IMI115591; on T. africana, Dalmatia from Island Arbe, Yugoslavia, 3 May,1912, O. Jaap, Otto Jaap, Fungi Selecti exsiccati no. 595, Hendersoniatamaricis Cooke, K; on T. gallica, near Barmouth, Merioneth, u.K., 29Aug. 1929, P. G. M. Rhodes, no. 4339, holotype of Coryneopsis

tamaricis Grove, K; on T. gallica, Cobo, Guernsey, 16 Sept. 1931,P. G. M. Rhodes, no. 4950, K; on T. gallica, Seaton, Devon, U.K., 18Mar. 1929, P. G. M. Rhodes, no. 3846, K; on T. gallica, DunwichBeach, Suffolk, U.K., 25 May 1977, MB. & J. P. Ellis, 1MI 214210; onTamarix, Houghton, near Heme Bay, Kent, UK, 24 Sept. 1929, exherb. W. B. Grove, K; on Tamarix gallica, Minsmere Level, Suffolk,U.K., 15 April 1977, M.B. & ]. P. Ellis, IMI 236998; on Tamarixgallica, West Wittering, Sussex, u.K., 5 Oct. 1985, B. C. Sutton, IMI298103.

Hendersonula mohgaoensis

Hendersonula mohgaoensis R. B. Singh & G. V. Patel,Botanique, Nagpur 9: 18, 1978 (1980), as 'mohgaoense'.

This species was described with

Pyenidia in chains, embedded in stroma in host tissue, oval orirregular each 126-145 x 85-140 j..lm in size, thin-walled; conidiogenous cells 4 j..lm long; conidia oval, 4 x l'5 j..lm, bicelled, thinwalled. Host dicot wood.

It was recovered from the Deccan Intertrappean beds ofMohgaonkalan, M.P. from the Cretaceous period.

Its position in Hendersonula is extremely doubtful if theconidial size is correct. Unfortunately the figures accompanying the account are too poor to elucidate its relationships withany degree of accuracy.

Hendersonula morbosa

Hendersonula morbosa Sacc., Syll. fungo 3: 445 (1884).

Illustration: Farlow (1876).

Saccardo (1884) cited Farlow's (1876) publication on blackknot of plum for the description and illustrations of thisfungus. Farlow described 'stylospores' which were in loculesinterspersed between those containing asci. He noted thespores arose in tufts or less frequently covering the wholesurface of the cavity; the conidia were borne on very slenderconidiogenous structures of different lengths, were oval,yellowish, 3-septate and 12 x 16 \.1m. Conidia were foundwhich fitted the description by Farlow: they are dilute brownin colour and some appear to have a small apiculus. Theseconidia only seemed to occur on older, weathered fungalstromata but it was not possible to associate them definitelywith any conidiomata or conidiogenous structures. Apiosporinamorbosa (Schwein.) v. Arx, the organism responsible for blackknot, is known to produce an anamorph referable to Fusic1adiumBonorden (Ellis, 1976). The Fusic1adium state is separate anddistinct from the fungus described as Hendersonula morbosa.Hyphomycetes colonizing old stromata of A. morbosa are notuncommon and it is quite probable that H. morbosa is one ofthem.

Specimens examined: On Prunus sp., Sphaeria morbosa, M. A. Curtis,Distr. W. G. Farlow, K; on Prunus pennsylvanica, Grand Marais, Minn.,

479

U.s.A., 2 Sept. 1891, F. W. Dewart, ex Herb. W. G. Farlow, asPlowrightia morbosa (Schw.) Sacc., K; on Prunus sp., Potsdam, NewYork, U.s.A., Ellis no. 336, ex H. W. Ravenel's herbarium, K.

Hendersonula morl

Hendersonula mori Sacco & Voglino, Annals mycol. 11: 232(1913).

The type was described from Morus alba collected in Italy andconsists of a glass slide with a small amount of fungus materialcontaining pycnidia but no conidia. The pycnidia had a darkouter rind and appeared unilocular. Notes on the packet statethe locular cavities were aggregated in a stroma and contained4-septate conidia 16-18 x 7 \.1m in size.

This description appears to be similar to that forCamarosporium mori Sacco before full maturity, but withoutadditional material, it is not possible to confirm the identity ofthis fungus.

Specimen examined: On Morns alba, Novara Pedemontii, Italy,Voglino, May, 1913, Herbarium Mycol. H. B. Patavinus, P. A.Saccardo, holotype, PAD.

Hendersonula phyllachoroides

Hendersonula? phyllachoroides Sacc., Bull. Soc. Bot. ltal. 4:219 (1893).

Saccardo (1893) was in some doubt as to the systematicposition of this species and whether the structures he termedspores were in fact spores. Pycnidia were described asproduced in stromatic colonies, variously shaped, black,phyllachoroid, covered by the cuticle, 3-5 mm long, joined,lacking ostioles, with a brown context. 'Spores' were referredto as densely parallel, stipitate, cylindrical, rounded at theapices, 40-45 x 7-9 \.1m, 4-septate, not constricted, each cellwith a large guttule, brown at the ends, paler.

Petrak (1949) could not place the species on the basis of theoriginal description and in the absence of any referencecollection it is not possible to confirm the species inHendersonula.

Hendersonula piniSphaeropsis sapinea (Fries) Dyko & Sutton in Sutton, The

Coelomycetes: 120 (1980).

Sphaeria sapinea Fr., Syst. Mycol. 2: 491 (1823).Diplodia sapinea (Fr.) Fuckel, ]. nassau. Ver. Naturk. 23-24: 393

(1870).Macrophoma sapinea (Fr.) Petrak, Sydowia 15: 309 (1961).Sphaeria pinea Desm., Ann. Sci. nat. Bot. 2 Ser., 17: 104

(1842).Sphaeropsis pinea (Desm.) Berk. & Broome, Ann. Mag. Nat.

Hist., 3 ser., 15: 401 (1865).Diplodia pinea (Desm.) Kickx, Fl. Cr. Fl. 1: 397 (1897).Botryodiplodia pinea (Desm.) Petrak AnnIs mycol. 20: 308

(1922).Macrophoma pinea (Desm.) Petrak & H. Sydow, Rep. Nov. spec.

Regni Veg. 42: 116 (1927), non Macrophoma pinea Pass.( = Dothiorella pinea (Pass.) Petrak & H. Sydow, fidePetrak, 1961).

Granulodiplodia pinea (Desm.) Zambettakis, Bull. trimestr. Soc.myco/. Fr. 70: 330 (1955), nom. inval. Arts 36, 41.


Granulodiplodia pinea (Desm.) Morelet & Lanier in Morelet.Bull. Soc. Nat. Archeol. Toulon Var 203: 12 (1973).

Diplodia conigena Desm., Mem. Soc. Sci. Agric. Lille 138(1846).

Macroplodia conigena (Desm.) Westend., Bull. Acad. Roy. Sci.Belg. 2 Ser., 2: 562 (1857).

Diplodia megalospora Berk. & Curtis, Grevillea 3: 3 (1874).Sphaeropsis pinastri Cooke & Ellis, Grevillea 7: 5 (1878), non

S. pinastri (Lev.) Sacc.Sphaeropsis ellisii Sacc. nom. nov., Syll. fungo 3: 300

(1884).Sphaeronema macrosperrna Karst., Hedwigia 23: 17 (1884).Phoma macrosperrna (Karst.) Sacc., Syll. fungo 3: 73 (1884).Macrophoma macrosperma (Karst.) Berlese & Voglino, Afti Soc.

Veneto Trent. 10: 178 (1886).Sphaeropsis p·tftemansii Henn., Hedwigia 48: 14 (1909).Hendersonula pini Diedicke, Krypt. Mark Brandenb. 9: 667

(1914), fig. 18, p. 640.?Sphaeropsis fragosiana Trav., Bull. Soc. Bot. Ital. 24 (1915), fide

Petrak & H. Sydow (1927).Phomatosphaeropsis pinicola Ribaldi, Ann. Spero Agr. N.5. 7: 847

(1953).?Sphaeropsis pinicola Speg., Anal. mus. Nac. Buenos Aires 20:

362 (1910), fide Petrak & H. Sydow (1927).Rejected synonymsPhoma pinastri Lev., Ann. Sci. Nat. Bot., 3 Ser., 5: 282

(1946), = Sclerophoma sp.

Sphaeropsis pinastri (Lev.) Sacc., Syll. fungo 3: 300 (1884),nomen sed non planta.

Diplodia pinastri (Lev.) Grove, ;. Bot. 54: 193 (1916), nomensed non planta.

Sphaeropsis micromegala Berk. & Curtis, Grevillea 2: 180(1874).

Phoma micromegala (Berk. & Curt.) Sacc., Syll. fungo 3: 73(1883).

Macrophoma micromegala (Berk. & Curt.) Berlese & Voglino,Atti Soc. Ven. Trent.: 185 (1886).

Illustrations: Punithalingam & Waterson (1970a), Petrak (1961),Ribaldi (1953), Sutton (1980).

Fruiting on bark. cones and needles of conifers. Myceliumimmersed, dark brown, branched, septate. Conidiomata pycnidial, globose, immersed, sometimes appearing superficial,separate or aggregated, dark brown to black. unilocular,0'3-0'5 mm diam, wall 6---8 cell layers, 30-60 ~m thick. outerwall of dark brown, thick-walled textura angularis, inner wall ofhyaline, thinner-walled textura angularis; cells darker aroundostiole. Ostiole central, circular, single. Conidiophores absent.Conidiogenous cells lageniform to cylindrical, occasionallypercurrently proliferating, discrete, indeterminate, hyaline,smooth, arising from inner wall of locule, 8'5-15 x 4'0 x 7'5~m. Conidia oblong to clavate, straight to slightly curved, atfirst aseptate, sometimes much later becoming 1-euseptate,walls O'5-1 ~m thick. outer surface of wall smooth, orappearing pitted, apex obtuse, base truncate, 30-55 x 11

18 ~m.

The conidiogenous cells are mostly lageniform andholoblastic in young pycnidia, but occasionally percurrentlyproliferating with the production of a second conidium. As

480

each conidium develops, a large vacuole is observed at firstnear the apex of the conidiogenous cell then later near thecentre of the developing conidium. The cytoplasm is granularin appearance; the conidium walls are hyaline and noticeablythickened in most specimens. As the conidium matures, itbecomes yellowish in colour, and the walls become differentially thickened. The conidium continues to mature to darkbrown and the apparent wall ornamentation becomes moreprominent; occasionally the central vacuole becomes biconcave at this time and eventually the conidium becomesseptate. Cheng-Guo, Blanchette & Palmer (1986) have shownconidia in two populations of S. sapinea to differ ultrastructurally. The deposition of electron-dense material in thecell wall layers is different and this is correlated with cultureappearance and infective ability. In older conidiomata, septateconidia may be as frequent as 90 %; the septa occur medianlyas well as one third the distance from the apex of theconidium. The conidia vary greatly in shape and size invarious collections made from the same host species, as wellas between different coniferous host species.

Smerlis (1970) reported that Botryosphaeria laricis (Wehmeyer) V. Arx & E. Muller produced an anamorph whichmatched the description of Macrophoma sapinea, a synonym ofS. sapinea. In addition to a redescription of both states of B.laricis, he gave the results of inoculation studies with isolatesfrom B. laricis and identified M. sapinea. He concluded thatwhile the anamorph of B. laricis was morphologicallyindistingUishable from M. sapinea, on the basis of pathogenicityof the latter isolate, they were not conspecific.

Sphaeropsis sapinea has been confused with the anamorphsof Botryosphaeria obtusa (Schw.) Shoemaker, Lasiodiplodiatheobromae (Pat.) Griff. & Maubl.. Aplosporella beaumontiiAhmad and Diplodia mutila Fr. on pine. The anamorph of B.obtusa can only be distinguished from S. sapinea on the basisof conidial size and host. for which an overlap exists, and bythe presence of the teleomorph B. obtusa. Lasiodiplodiatheobromae can be readily distinguished from S. sapinea by thepresence of paraphyses in the eustromatic conidiomata andthe ovoid, brown, striate conidia. The conidia of A. beaumontiiare very distinctive in the reticulate ornamentation present onthe external wall, the eustromatic conidiomata and paraphyses.Diplodia mutila produces thick-walled conidia, medianly 1euseptate, with walls that show no apparent ornamentationwith light microscopy. In identification of the various taxadiscussed, broken mature conidia provide a more reliableindication of the presence or absence and the location of wallornamentation: all of these taxa produce conidia with quitegranular cytoplasm.

Fries, Scleromyceti Sueciae Exsiccati no. 126. Sphaeria sapineaFries serves as the type for this taxon. The majority of thespecimens of this exsiccatus are effete and many of theconidiomata have been broken; several of the specimens alsocontained additional fungi. The collection from herb. Bseemed to be in the best condition and yielded one slidemount showing conidiogenous cells with conidia in variousstages of development. including the later stage withdistinctive wall ornamentation so this specimen has beendesignated the lectotype.

Diedicke's type material of Hendersonula pini was not


located but his description and illustrations seem adequate toidentify his fungus as S. sapinea.

Ribaldi (1953) described a new genus typified by Phomatosphaeropsis pinicola from Pinus nigra. Although the typematerial was not seen, from the description and excellentillustrations, it appears that he described S. sapinea and ahyphomycetous saprophyte or parasite. Fungi similar toRibaldi's descriptions and illustrations were found in Ravenel'scolledions of Diplodia megalosporus on cones of Pinus taedafrom herb. NYS and K. The hyphomycete produces hyaline,holoblastic, septate, tetraradiate conidia. We did not observethe hyaline, bacilliform, 2-7 x 1-2 J,1m conidia described andfigured by Ribaldi but they do not appear to belong to thehyphomycete.

The description of S. sapinea is cosmopolitan and it occurson a wide range of coniferous hosts, some of which are listedbelow. It has been implicated in various needle and stemblights and diebacks, usually after the tree has sustainedmechanical damage. It is a common saprophyte in forest litterand may cause a blue stain in freshly cut timber. A partialreview of the phytopathological literature may be found inPunithalingam & Waterson (1970a).

More than 300 specimens have been examined from herb.IMI, K. BPI, NY, CUP, UPS, B, G, L, S, DAOM, NYS whichoccur on cones, needles and bark of: Abies excelsa, A. procera,Cupressus lusitanica, Larix sp., Pinus austriaca, P. caribaea, P.coulteri, P. canariensis, P. elliotii, P. halepensis, P. laricionis, P.maritima, P. mugho, P. patula, P. pinastri, P. pinea, P. ponderosa,P. resinosa, P. radiata, P. sylvestris, P. strobus, P. taeda, Pseudotsugataxifolia, Thuja sp. from temperate and tropical regions inNorth and South America, Africa, Europe, Asia and Australia.Not all these samples have been included in the specimensexamined; it only includes type material, exsiccati and otherimportant colledions.

Specimens examined: On Pinus sylvestris twigs, C. Roumeguere, Fungi

Selecti Gallici no. 554 Sphaeria pinea Desm., K, NY; On P. montana,

Padova, Italy, Dec. 1876, Bizzozzero, Saccardo, Mycotheca Veneta no.1205 Diplodia pinea Kickx, BPI, NY, K; on P. pinea stems and needles,Roma, Italy, Mar. 1902, D. Saccardo Mycothem Italica no. 955Diplodia pinea (Desm.) Kickx, NY, BPI, L, S, K; on cone scales of Pinus

sp., Eastboume, Sussex, U.K. C. J. Muller, M. C. Cooke FungiBritannici Exsiccati, Ed. sec. no. 16 Phoma pinastri lev., K. E; on conescales of P. laeda, Society HilL South Carolina, U.s.A., Apr. 1555, no.5012 Diplodia megalospora B. & C, isotype of D. megalospora, K, FH,BPI, K, NYS; on P laricionis, Padova, Italy, Jan. 1550, Bizzozzero,Saccardo Mycalheca Venela no. 1531 Diplodia pinea (Desm.) Kickx f.slrobilicola, BPI. NY, K; on cone of Pinus sp., Chelwoodgate, Sussex,U.K., P. W. James & J. Cannon, 20 Nov. 1966, F. L. Balfour-Browne,Flora of the British Isles, as Sphaeropsis ellisii Sacc., BM; on cone ofP. auslriaca, London, Ontario, Canada, April 1915, J. Dearness, Fungi

Columbiani - Elam Bartholomew no. 4777 Sphaeropsis pinaslri (lev.)Sacc., K, IMI 6657; on P. halepensis, Fort Cunningham, South Africa,I. B. Pole Evans, Sydow Fungi exotici exsiccati no. 279 Sphaeropsispinicola Speg., L B, S, K, UPS, NY, IMI 6689; on conifer cone scales,France ex Hortus Mass, Castagne, Corsica, Herb. E. Fries, Diplodia

pinastri Montagne, UPS; on P. sylvestris, J. B. H. J. Desmazieres,Plantes Cryptogames de la France, Ann. 1525-51. Ann. 1534. no. 1277Sphaeria pinea Desm., isotype, PC, BR, NY, K; on P. sylveslris, Lebisey,France, April 1841, Roberge, no. 1559 Sphaeria pinea Desm., K, G, B;on P. sylvesfris, Austria, 10 June 1961, F. Petrak, Cryptogamae

481

exsiccatae editae a Museo Hist. Natur. VindobonensL no. 4548Macraphoma pinea (Desm.) Petro & Syd., PC, B, BPI, BR, L, G, S, UPS;on P. sylvesfris, Terrington, St Clements, Norfolk, England, Autumn,1874, C. P. Plowright, de Thiimen, Mycafheca Universalis no. 353Diplodia sapinea FckL K, B; on P. sylvestris, Desmazieres Planfescryptogames de France no. 777 Sphaeria pinea, L, K; on P. sylvesfris,

C. Roumeguere, Fungi Gallici Exsiccafi no. 842 Sphaeria pinea var. b.Desm., NY; on P. sylveslris, Aut. 1875, C. Roumeguere, Fungi SelectiGallici no. 584 Sphaeria pinea, Desm., L, NY, K; on Pinus, ex Herb. DrH. S. Billings, ex Herb. Desm. in authors' hand as Sphaeria pinea, NY;on Pinus sp., E. Fries, Scleromyceti Sueciae Exsiccati, Suecia Smaland,Femsjo, Sweden, no. 126 Sphaeria sapinea Fries, lectotype: B,isotypes, G, K, E, UPS, C, BR, FH; on Pinus sp. King's Lynn, Norfolk,U.K., M. C. Cooke, Fungi Britannici Exsiccati, Ed. Sec., no. 17, Dipladiasapinea, IMI 6875; on Pinus sp., J. E. Vize, Micrafungi Brilannici no. 8Diplodia sapinea FckL IMI 57008; on P. sylvesfris, Ticinensi, Italy,Cavara Fungi Longobardiae Exsiccati no. 95 Sphaeropsis ellisii Sacc., S, K;on P. maritima, Rabenhorst, Fungi Eurapaei Exsiccati no. 454Macroplodia canigena Westend., K; on Abies excelsa, London, Ontario,Canada, May 1915, J. Dearness, Fungi Columbiani, Elarn Bartholomewno. 4775 Sphaeropsis pinasfri (Lev.) Sacco f. abietis Dearness &Bartholomew, IMI 6655, K; on P. maritima, Westend. & Wall., Herb.Crypt. Belg. no. 1230 Macropladia conigena (Desrn.) Westend., K; onP. sylvestris, New Jersey, U.S.A., J. B. Ellis, no. 2917 Sphaeropsis

pinastri C. & E., K; on P. sylveslris, Newfield, New Jersey, U.S.A., Ellis,North American Fungi no. 525 Sphaeropsis pinasfri C. & E., isotype, K;on P. austriaca, Newfield, New Jersey, U.S.A., Jan. 1894, Ellis &Everhart, Fungi Columbiani no. 967 Sphaeropsis ellisii Sacc., K; onJuniperus sabina, Brandenburg, Germany, 29 Sept. 1904, P. Vogel.Sydow, Mycatheca germanica no. 377 Diplodia juniperi Westend., E;on P. ponderosa Dougl., Hayden Flat, Trinity County, California,U.s.A., 16 March 1960, J. O'Keefe, Herb. Univ. Calif. Fungi ofCalifornia no. 47316 Diplodia pinea (Desm.) Kickx, CUP, BPI; on P.sylvestris, Albion, Peel Co., Ontario, Canada, 26 Sept. 1954, R. F.Cain, Cryptogamic herb. Univ. Toronto, no. 30260. Diplodia pinea(Desm.) Kickx, DAOM 40824, UPS, S, NY, BPI; on P. sylvestris,

Brandenburg, Germany, 25 April 1937, P. Vogel. Sydow, Mycofhecagermanica no. 3373 Macrophoma pinea (Desm.) Pet. & Sydow, L, S, B;on P. sylvestris, Mito-City Ibanakii pref.. Japan, 3 Oct. 1966, T.Kobayashi. lab. Forest Path. Herb. Gov. For. Exp. Sta. Tokyo, Japan,no. 3060. Dipladia pinea, BPI. NY; on P. coulferi, Julian, San Diego Co.,California, U.S.A., 2 May 1967, D. R. Miller, Herb. Univ. Calif.. Fungi

of California, Diplodia pinastri Grove, IMI 142194; on Picea excelsa,Mustiala, Finland, 6 Dec. 1866, Karsten, as Sphaeronaema macro

spermum Karst. in Hohnel herb. under no. 1955 Diplodia pinea (Desm.)Kickx annotated in Hohners own hand, holotype as S. macrospermum,

FH; on P. canariensis, Teneriffe, Esperanoa, Canary Islands, June,1900, M. Bornmuller. type of no. 1621 Macrophoma canariensis P.Mag., S; on Pinus sp., S. Paulo, Brazil, April 1905, P. Hennings, no.1258 Sphaerapsis puttemansii, holotype, S; on P. sylvestris, Austria,1894, Fuckel. Herbier Barbey-Boissier, Fuckel. Fungi rhenani no. 2301Dipladia pinea (Desm.) Kickx, S, DAOM 126441; on Thuja sp., Berlin,Germany, Nov. 1895, P. Sydow, Sydow Mycotheca Marchica no.4496 Diplodia thujae Westend., NY, K; on P. sylvestris, Austria nearHom, 10 June 1967, F. Petrak, F. Petrak-Pilzherbariurn, Macrophama

pinea (Desm.) Petrak & H. Sydow, IMI 94421, BPI. B; on conifer sp..418 Link, no. 120 Sphaeria sapinea, B; on P. sylvestris, Prignitz region,East Germany, 2 Oct. 1906, 0 Jaap, Herbarium G. Bresadola no. 456Batryodiplodia pinea (Desm.) Pet. & Syd., BPI; on P. sylvestris, Berlin,Germany, June, 1558, P. Sydow, Sydow, Mycafheca Marchica no.2081 Diplodia sapinea K; on P. sylvestris, Padova, Italy, June 1897,D. Saccardo, Mycotheca italica no. 160 Dipladia sapinea (Fr.) Fckl. K;on P. sylvesfris, Hom, Austria, 10 June 1961, Cryptogamae exsiccataeeditae a Museo Hist. Natur. Vindobonensi no. 4548 Macrophama


pinea (Desm.) Petro & Syd., BPI, S, B, L, PC K; on P. sylvesfris, Abiesand Cedrus, Desmazieres, Crypt. France ser. i, no. 1482 Diplodia(Chromospora) Conigena, K; on P. sylvesfris, Parma, Italy, G. Passerini,C. Roumeguere, Fungi Gallici exsiccafi no. 2855 Diplodia conigena var.italica, K; on bark, Padova, Italy, D. Saccardo, Mycotheca italica no.355 Diplodia conigena Desm., K; on cones of Cedrus, Pisano, Italy,Arcangeli, Erb. Crill. Ital. ser. II, no. 1388 Diplodia conigena, K; on P.sylvestris, Budenheim, Germany, Autumn, L. FuckeL Fungi rhenani no.538 Diplodia pithyophila FckL K.

Hendersonula pinicola

Hendersonula pinicola Dearness, Mycologia 20: 236(1928).

Hendersonia dearnessii Darker, Can. ]. Bot. 45: 1410 (1967).

The type specimen of H. pinicola contains Bifusella linearis(Peck) V. Hohn., which produces a stroma, Lophadermiumbrachysporium Rostrup, Alternaria sp. and the brown, 2-septateconidia described by Dearness. These latter conidia cannot berelated to any definite conidiomata.

Darker (1967) was also aware of the composite nature ofthe type specimen and reasoned that since the 'stroma'described by Dearness belonged to the Bifusella, the remainingpycnidial fungus was obviously a Hendersonia. As Hendersoniapinicola Wehmeyer is a prior name in Hendersonia, Darkerproposed the nomen novum Hendersonia dearnessii.

It is impossible to select any of these elements as asatisfactory type which agrees with Dearness's originaldescription. Both Hendersonula pinicola and its later synonymHendersonia dearnessii are of doubtful application.

Specimen examined: On Pinus strobus, Davidson River, North Carolina,U.s.A., 8 June 1925, B. B. Hedgecock, det. Dearness no. 5864,Dearness colI. DAOM; on P. strobus, Roan Mtn, Tenn., U.S.A.,G. G. Hedgecock no. 8678, 16 Sept. 1918, det. Dearness no. 5864,Dearness col. DAOM; Davidson River, North Carolina, U.s.A.,G. G. Hedgecock, 6 Aug. 1925, det. dearness no. 5864. Holotype,Dearness col. DAOM, isotype BPI.

Hendersonula platani

Hendersonula platani Camara & Luz, Agron. Lusit. 1: 187(1939).

This species was described from Platanus orientalis in Portugal.The type specimen was not located, but a specimen was sentfrom LISE, on Platanus hybrida, which appears to fit theoriginal description and illustrations very well. It contains thefruiting structure of a Massaria species which cannot beidentified further because it is effete. The ascospores agreeclosely with the measurements (28-34 x 12'5-14 I-lm) andillustrations of the conidia described for H. platani. Macrodiplodiopsis desmazieresii (Mont.) Petrak is also present.

The name H. platani is regarded as of doubtful applicationas the ascomycete is inadequate for identification.

Specimen examined: On branches of Platanus hybrida, Travos bomtese alto Dauno-Rogua, Portugal, April 1952, H. T. Lucas, no. 1591LISE.

482

Hendersonula pterosparli

Hendersonula pterosparti Camara & Luz, Agron. Lusi!. 5:138 (1943).

This fungus was described from Pterospartum tridentatum inPortugal. The type specimen could not be located. Theauthors described the fungus as 'pycnidiis in acervulusstromaticus corticem perforantibus, botryose aggregatis' andnoted the 'sporophores' were not visible. The conidia weredescribed as oblong-ellipsoid, 3-septate occasionally with alongitudinal septum, eguttulate, brown, 14-22 x 6-8 I-lm.

In the absence of a specimen, the data are insufficient todetermine the identity of the fungus. The presence oflongitudinal septa in the conidia suggest that the species isinappropriately placed in Hendersonula.

Hendersonula romagnesii

Hendersonula romagnesii Negru, Bull. trimestr. Soc. mycal.Fr. 80: 324 (1964).

This organism was described from branches of Laburnumanagyroides from Romania. It was described as producing adothideaceous, erumpent, black multiloculate stroma withshort simple conidiophores and yellow brown, elliptical toelongate, 3-septate conidia, 15-16 x 5-6 I-lm long.

Herb. BUC does not hold the type specimen and no replywas received from a query to the author. The originalillustrations are very general and the description is insufficientto identify this fungus further.

Hendersonula saccharina

Hendersonula saccharina Ahmad, Biologia, Lahore 17: 17(1971).

This species was described from culms of Saccharum sp. inWest Pakistan (Pakistan) and described as producing globosepycnidia in erumpent stromata, 110--250 I-lm diam, composedof parenchymatous tissue; conidiophores papillate, hyaline;conidia cylindric with slightly narrowed ends, 3-5 septate, notconstricted at septa, yellowish brown in mass, 15-20 x 2'6I-lm. Type material was requested but not received and thedescription is insufficient to confirm correct placement inHendersonula.

Hendersonula tecomae

Hendersonula tecomae Ahmad & Arshad, Biologia, Lahore18: 118 (1972).

This species occurs on branches of Tecoma sp. in WestPakistan (Pakistan) and was described as producing anerumpent, multiloeulate stroma composed of pseudoparenchymatous tissue, 1'26 mm diam, with yellow-brown, ellipsoid oroblong ellipsoid, 3-septate conidia, 16'5-21 x 6'5-8 I-lm.

Type material was requested but not received and thespecies cannot be placed with any accuracy from the publisheddescription.


Hendersonula thelebola

Hendersoniopsis thelebola (Sacc.) v. Hahn., Annis mycol.16: 124 (1918).

Stilbospora thelebola Sacc., Michelia 2: 542 (1882).

Hendersonula thelebola (Sacc.) Kobayashi, Bull. Govt for. Exp. St.226: 32 (1976), non rite publ. (Art. 33.2).

Hendersonula konseiensis Kobayashi, Trans. mycoI. Soc. Japan 4:8 (1962).

Illustrations: Sutton (1980).

This binomial was introduced for the anamorph of Pseudovalsella thelebola (Fr.) v. Hahn. and several synonyms werelisted, including Hendersonula konseiensis Kobayashi. Amongstthese was also Hendersoniopsis thelebola (Sacc.) v. Hohn., thetype species of Hendersoniopsis v. Hohn. If this species isindeed correctly placed in Hendersonula then by applying theprinciple of priority it would mean that Hendersoniopsis is theprior generic name. Wehmeyer (1941) and Sutton (1980) haveredescribed Hendersoniopsis and it is clear from the latteraccount that Hendersoniopsis is quite inappropriate for thespecies accepted in Hendersonula. The correct name for theanamorph of Pseudovalsella thelebola is Hendersoniopsis thelebolaand Hendersonula konseiensis and Hendersonula thelebola aresynonyms.

483

Hendersonula toruloidea

Nattrassia Sutton & Dyko, gen. nov.

Conidiomata eustromatica, erumpentia, uni- vel multilocularia: parietesex textura angulari brunnea incrassata compositi; loculi per ostiolosseparatos dehiscentes. Conidiophora absentia. Cellulae conidiogenae

lageniformes vel ampulliformes, discretae, ex cellulis interioribusparietum oriundae, seriem conidiorum ad unum planum producentia,collo absenti, spissescenti periclinali et canali cytoplasmico lato.Conidia holoblastica, demum 1-2 euseptata, versicoloria, laevia.

Sp. typ.: Naftrassia mangiferae (H. & P. Syd.) Sutton &

DykoConidiomata eustromatic, erumpent, uni- to multi-loculate;

walls composed of thick-walled, brown textura angularis;locules opening by means of separate ostioles. Conidiophoresabsent. Conidiogenous cells lageniform to ampulliform, discrete,arising from the inner wall of the stroma, producing asuccession of conidia at one level. collarette absent, periclinalthickening and cyptoplasmic channel wide. Conidia holoblastic,eventually 1-2 euseptate, versicoloured, smooth.

Naftrassia differs considerably from Hendersonula eventhough both genera have eustromatic conidiomata andpigmented conidia. In Naftrassia conidiomata are erumpentwith many separate locules each of which is ostiolate but in

Fig. 8. NaUrassia mangiferae. A, Conidia; B, conidiogenous cells and developing conidia; C vertical section of a conidioma.

A

oOu

\JlOJiffi

B

32

500 Jlffi

c

MYC 93


Fig. 9. Nattrassia mangiferae. Arthric synanamorph.

~...

~ ".' '.

':'"

••••••••••••

fJ.·:.:···

CJ

10.um

(\U

Hendersonula sensu stricto conidiomata are mycoparasitic on orin ascomycete fruit bodies of stroma tissue. There is asimilarity in conidiogenesis in that both genera form conidiafrom what have been termed 'phialides' (sensu Sutton, 1980)but in NaUrassia percurrent proliferation of the conidiogenouscell produces a succession of conidia which secede at the samelevel whereas in Hendersonula proliferation may result in someconidia seceding at successively higher levels. Conidialmorphology is quite different. In NaUrassia conidia take sometime to mature so from a single conidioma hyaline, palebrown, and versicoloured smooth conidia may be present andthese may be aseptate, with a single septum towards the baseor 2-septate and the median cell larger than the cell at eachend. Conidial cells in Hendersonula are of similar pigmentationand even though 0-3 septa are produced the verruculose orsmooth conidia do not become versicoloured. The septa areprominent and appear thicker than the periclinal walls whichis not the case in NaUrassia. Another feature differentiatingNaUrassia is the presence of toruloid arthric vegetative cells,often developing in the absence of conidiomata and by whichthe species may be readily identified.

Naltrassia mangiferae (H. Sydow & Sydow) Sutton & Dyko,comb. nov. (Figs 8, 9)

Dothiorella mangiferae H. Sydow & Sydow in H. Sydow,Sydow & But!., AnnIs mycol. 14: 192 (1916).

Fusicoccum eucalypti da Camara, An. Inst. Super. Agron. 3: 32(1929).

Hendersonula toruloidea Nattrass, Trans. Br. mycol. Soc. 18: 197(1933).

Hendersonula cypria Nattrass, Cyprus Fungi, Nicosia: 43(1937).

Hendersonula agathidis Young, Qd II. Agric. Sci. 5: 12 (1948), as,agathi'.

Arthric synanamorph:

Scytalidium dimidiatum (penz.) Sutton & Dyko, comb.nov.

Torula dimidiata Penz., Michelia 2: 466 (1882).

Exosporina fawceUii Wilson, Hilgardia 17 (12): 427 (1947).

Scytalidium lignicola Pesante, Ann. Spero agron., N.5., 11: supp!.p. cclxv (1957), as 'lignicolum'.

Fruiting on woody tissue of Prunus spp., Malus sp., Citris spp.,Iuglans spp., and a wide range of other hosts (Punithalingam& Waterston, 1970b). Mycelium superficial, immersed, branched, septate, hyaline to dark brown. Conidiomata eustromatic,immersed, eventually erumpent, dark brown to black,unilocular to multilocular, globose, up to 2 mm diam; wall of7-12 cell layers, up to 20-43 ~m thick, outer wall of irregular,thick-walled, dark brown textura angularis; inner wall ofhyaline, thinner-walled textura angularis. Ostiole central to eachlocule, circular, papillate. Conidiophores absent. Conidiogenouscells lageniform to ampulliform, hyaline, discrete, collaretteabsent, periclinal thickening and cytoplasmic channel wide,arising from inner wall of the locules, 6'5-14 x 2'5-4 ~m.

Conidia holoblastic, ellipsoid to nearly fusiform, hyaline, atfirst aseptate, then becoming 1-2 (3) euseptate, central celldark brown, end cells hyaline to light brown, 10-16 (21)x 3'5-6'5 ~m. Arthric state frequently present, cylindricaltruncate, oblong-obtuse to doliiform, dark brown, thickwalled, arising from disarticulating hyphae, 0-1 septate,4-16'8 x 8'5 ~m.

In a revision of some stromatic coelomycetes on Acacia andEucalyptus species from Western Australia, Sutton & Davison(1983) re-examined Fusicoccum eucalypti da Camara and pointedout that it provided an earlier name for the species known as


Hendersonula toruloidea. They also alluded to other nameswhich might apply to the same fungus. Dothiorella mangiferaeH. Sydow & Sydow in Sydow, Sydow & Butler (1916) is acase in point. This was described from dead branches ofMangifera indica with stromatic multilocular conidiomata andoblong to subfusoid hyaline aseptate conidia measuring9-13 x 3'5-4'5 ~m. The type confirms this information and inaddition many 2-septate versicoloured conidia were also to befound on the conidiomata and bark surface. These measured12-13 x 5 ~m and clearly represent the mature conidia. Itwould be reasonable to assume that even earlier names existfor this seemingly cosmopolitan species but so far none hasbeen found so the new combination into NaUrassia isaccordingly made.

Another species examined in the same context is Hendersoniacreberrina Sydow, Sydow & Butler (1916) which was originallydescribed from Mangifera indica fruit and later reported byBrodrick & van der Westhuizen (1976) to cause soft brown rotof this fruit in South Africa. Sutton (in litt.) pointed out thatthe S. African fungus differed from H. creberrima in havingversicoloured conidia rather than the evenly pigmented onesdescribed and figured in the original account. Brodrick & vander Westhuizen (1976) suggested that their material named H.creberrima may be the same species as Hendersonula cypriaNattrass. Comparison of the S. African fungus with theoriginal account clearly indicate that it is not H. creberrimabecause the latter shows no accompanying arthric state,conidiomata are simple pycnidia, the conidia are of evenpigmentation and septa not prominent and the conidial size of17-21 x 6-8 ~m and conidiophore length of 20 ~m are quitedifferent from that known for H. toruloidea or H. cypria. It isconcluded that the material described by Brodrick & van derWesthuizen (1976) as H. creberrima is in fact H. toruloidea andthat the relationships of the true H. creberrima do not impingeon the status of H. toruloidea.

Nattrass (1937) proposed the name Hendersonula cypria fora fungus on Prunus armeniaca from Cyprus which producedslightly larger conidia than H. toruloidea (12-16 x 5-7 ~m v.12-13'5 x 4-5'5 ~m) and were overall brown. A Scytalidiumstate was not seen in H. cypria. After study of several isolatesof H. cypria and many of H. toruloidea, it is concluded that thedifference between the two taxa is not sufficient to warrantmaintaining them as separate. The ability to producearthrospores appears to be a variable trait, with most isolatesshowing abundant production.

Young (1948) proposed H. agathidis for a fungus similar toH. toruloidea occurring on Agathis palmerstoni from Australia.The species separation was made on the basis of largerconidiomatal size and slightly longer and wider conidia whichoccasionally become 3 septate. The fungus in the typematerial fits comfortably within the range of variation seen inH. toruloidea and while the conidiomata tend to be morepycnidial, this is not uncommon for H. toruloidea onmonocotyledonous hosts. The difference between these twotaxa is insufficient to maintain them as separate.

The arthric state is most frequently seen in or recoveredfrom the diseased host and is referable to Scytalidium Pesante.This state has been described in some detail by Sigler &

Carmichael (1976) and Hughes (1953).

485

NaUrassia mangiferae (under the name H. toruloidea) has beenisolated as a dermatophyte on people immigrating fromtropical countries to the U.K. Moore (1988) found thatprimary isolates from 128 cases of H. toruloidea infection ofhuman skin and nails were invariably of the Scytalidiumsynanamorph. Three distinct stable colonial forms weredescribed and conidiomatal formation was observed inrepresentatives of two of the three colonial forms recognized.There was a significant relation between colonial form andgeographicalj racial origins of the patients. While conidiomataare rarely observed in culture, Campbell et al. (1973), Campbell(1974) and Sigler & Carmichael (1976) have been successful ininducing their production.

In culture, isolates of N. mangiferae from plant and humanhosts produced abundant, felty to fluffy, grey to brownishblack aerial mycelium. In some isolates, single conidiomata oraggregations of black conidiomata, 500 ~m diam, wereformed after incubation of cultures for 6 wk to 2 monthsunder near uv light. Conidia extruded from the ostioles are atfirst ellipSOid and hyaline, much later becoming 2 septate witha brown median cell or all three cells becoming brown. Betterresults were obtained by inoculating sterilized Malus sp. twigsand subjecting them to near uv light. Abundant fruitingoccurred from most isolates in 3-6 months.

No holotype for H. toruloidea was specially designated byNattrass (1933) or in material sent to herb. 1M!. Material inherb. IMI which has been regarded as type and accessed asIMI 8628 a-g, consists mostly of isolates from various stonefruit hosts made by Nattrass from 1929 to 1932. One of thesespecimens, however, was collected on a natural substrate(Prunus armeniaca) and shows the characteristic coelomycetestate of the fungus with both immature and mature conidia.This is deSignated as lectotype.

Originally described by Nattrass on plum, apricot andapple this taxon has now been reported from tropical andsemi-tropical countries of Europe, Africa, Asia, North andSouth America causing gummosis, die back, wilt and canker ofnumerous, mostly woody hosts (Punithalingam & Waterston,1970 b). It is a wound-invading pathogen on branches andtrunks damaged by sunburn, freezing, or pruning (Calavan &

Wallace, 1954). The fungus attacks the cambial region of thetree and progresses downwards causing wilt and branch dieback (paxton & Wilson, 1965; Khan, 1955). Later the barkblisters and cracks exposing the black spore masses to winddissemination. The wood below shows brown staining.

Selected specimens examined: Several hundred collections in herb. IMIshow the Scytalidum state; the following is a representative selectionof material showing the coelomycete state. On Prunus armeniacaCairo, Egypt, Aug. 1930, R. M. Nattrass, lectotype of Hendersonulatoruloidea Nattrass, IMI 8628 g; on Ficus carica. Nicosia, Cyprus, Nov.1933, R. M. Nattrass, IMI 8629; on Populus alba, Nicosia, Cyprus,Nov. 1933, R. M. Nattrass, IMI 8620; on Prunus armeniaca, Kythrea,Cyprus, July 1934. R. M. Nattrass no. 442. holotype of Hendersonulacypria Nattrass, IMI 42047, K; on P. armeniaca, Kythrea. Cyprus, Dec.1937, R. M. Nattrass, IMI 2138; on Fucraea sp., Fed. expo Sta.Serdang, Malaya, March 1955, A. Johnston, IMI 60094; on Ipomoeabatatas, Fed. expo Sta. Jalea Kebum, Malaya, March 1955. A. Johnston,IMI 60093; on Malus pumila. Iran, 27 Jan. 1976, D. Ershad, IMI200622; on Melia aUiderach, Pirzwala, Punjab. Pakistan, 22 Feb. 1952,


A. H. Khan, IMI 55277a; on Ananas comosus, Pulan Lekong BesarIso!. Sta. Malaysia, 1961, A. Johnston, IMI 62494a; on A. comosus,Selangor, Malaysia, Dec. 1960, S. J. Navaratnam, IMI 84390; onSansevieria guineensis, Kindia, Guinea, March 1962, J. Kranz, IMI93227; on Plumeria obtusa, Mandalay, Burma, 9 Oct. 1972, M.Thaung, IMI 10555; on Philodendron bipinnatijidum, Jabalpur, India,July 1964, U. P. Sahni, IMI 108124; on Sansevieria sp., Jaipur, India,1969, S. Sharma & M. L. Sonar, IMI 143900; on Mangifera indica,Dambata, Kano Province, Nigeria, 20 June 1960, E. Harris. IMI81789; on Musa (Cavendish), Newcastle, Jamaica, 1961, D. Meredith,IMI 95575; on Manihot utilissima, Tamale. Northern Territories,Ghana, 16 April 1949, S. J. Hughes, IMI 39364; on M. utilissima,Njala, Kori, Sierra Leone, 5 July 1949, F. C. Deighton, IMI 37368; onAgave sisalana, Moor plantation, Ibadan Prov., Nigeria, 13 June,1961, A. G. Bailey, IMI 99532; on ]uglans regia, Visalia, Tulane Co.,Calif., US.A., May 1948. E. E. Wilson, authentic material forExosporina fawcettii Wilson, IMI 31449; on Ananas sativa, Njala, Kori,Sierra Leone, 26 Jan. 1955, A. B. Katta, IMI 60248; on Citrus sinensis,Lyallpur, West Pakistan, Feb. 1957, A. H. Khan, IMI 68859; on A.comosus, Turufia plantation, Rendova, BS.J.P., 20 May 1959. A.Johnston, IMI 77097; on Agave americana. India, 1977, R. C. Rajak,IMI 2II913; on Acacia auriculiformis, Dehra Dun, India, 1968, B. K.Bakol. IMI 136774; on Morus alba, Punjab. Pakistan, 22 Feb. 1952,A. H. Khan, IMI 49167; on M. alba, U.S.A., 1953, H. Dill, IMI51975; on Eucalyptus sp., Portugal. 1969, collection no. 35a, 36, IMI139172; on Malus fruit, Allahabad University, India, 1962, R. Tandon,IMI 95023; on Malus fruit, College of Agriculture, Univ. of Baghdad.Iraq, 1961, K. A. Debbagh, IMI 86983; on Manihot utilissima,Tamale, Ghana. 16 April 1949, S. J. Hughes, IMI 39473; on Mangiferaindica, NelspruiL E. Transvaal. South Africa, 1972, G. van Westhuizen.IMI 168955; on Agathis palmerstoni, Gadgarra, Atherton Tableland,North Queensland, Australia, 9 July 1936, H. E. Young, holotype ofHendersonula agathidis, IMI 12279; on Agathis sp., Nadaevatu, VitiLevre, Fiji, 9 June 1977, A. Frith, IMI 214069; on Eucalyptus globulus,Lisboa, Portugal. April 1928, B. Oliveira 36, holotype of Fusicoccumeucalypti da Camara, LSVA; on Mangifera indica, Lucknow, India, 22Oct. 1908, F. Bahadur (E. J. Butler 1724), holotype of Dothiorellamangiferae H. Sydow & Sydow, HCIO.

Misdetermination: On Sequoia gigantea, Riverside, California, U.S.A., 7March 1957. L. Bonar, California Fungi no. 1036 Hendersonulatoruloidea NatL ex LPS, CUP, NY, S, K. B, = Hormonema sp.

Hendersonula varians

Hendersonula varians Curzi & Barbiani, Inst. Bot. R. Univ.

Pavia Lab. CriN. Ital.: 184, fig. XL fig. 4-6 (1927).

This fungus was described from Salix sp. in Italy but it was notpossible to locate the type specimen. The description is asfollows:

'Stromatibus forma varia, saepe erumpentibus, contextu superne atro,parte inferiori loculis pallide fuligineo praeditis; sporulis oblongis.piriformibus v. ovalibus, denique plerumque 3-septatis, raro 1-2septatis. loculis extremis amplioribus, 10-12 x 4'5 11, fuligineis'.

The figure shows a multiloculate stroma producing conidiaaround the full circumference of the locule. No ostiole can beseen and the conidia are shown to be clavate to cylindrical,brown and 1-3 septate.

Neither the diagnosis nor the illustration are sufficient toidentify the species with certainty.

486

Scolecosporella varians (Sacc.) Sutton & Dyko, comb.nov.

5tagonospora varians Sacc., Annis myco!. 12: 310 (I914).

Fruiting on leaves of 5ymplocus obtusa and 5. whitfordii.

Mycelium immersed, hyaline to brown, branched, septate.Conidiomata pycnidial, immersed, subepidermal, globose toirregular, dull black, unilocular, 0'3-1 mm diam; wall 4-9 celllayers, 15-37 ~m thick, composed of pale brown, thick-walledtextura angularis; causing area of discoloration on leaf up toI-I'5 mm diam. Ostiole circular, central, indistinct, sometimespapillate. Conidiophores absent. Conidiogenous cells lageniform,discrete, determinate, smooth, hyaline, arising from cells lininglocule, 5-14 X 2-3 ~m. Conidia holoblastic, fusiform tosubulate, hyaline to pale brown, 3-euseptate, thin-walled,smooth, apex acute, base truncate, (24) 30--38 X (4) 5-6'5 ~m.

The conidiogenous cells are very short and appearsomewhat similar to the texfura angularis from which theyarise, hence they are easily overlooked. The conidia arecharacteristic in this genus as their pigmentation is quite diluteexcept at the septa which are pale to medium brown. Thedilute pigmentation and the fusiform to subulate shape of theconidia distinguishes taxa in this genus from those in5tagonospora Sacco nom. cons.

This fungus was found on the type specimen of Hendersonia

symploci but is quite distinct. The conidiomata of 5. varians aresubepidermal, dull black and larger in diameter than those ofthe parasitized Dermatodothis with its subcuticular, shiny blackascostroma. The conidia of 5. varians are also substantiallylonger and lighter in colour than those seen in H. symploci.

Petch (1924) suggested that the name Hendersonia symploci

was based on the conidia of 5. varians and the stromata ofDermatodothis zeylanica. As previously shown, this is incorrect.5tagonospora varians was not found on the Petch specimens.

Saccardo (1914) described this taxon from 5. whitfordii

growing in the Philippines with fusiform, 3-septate, hyalineconidia 22-24 X 3'5 ~m. The holotype, however, contains not

\only the conidia of 5. varians but also a stromatic ascomycetewith pseudoparaphyses, bitunicate asci, sub-hyaline to dilutebrown, 5-septate ascospores, 27-31 X 6'5-7'5 ~m. The ascomycete while perhaps not referable directly to Leptosphaeria

Ces. & de Not. may belong to a related genus. Measurementsof conidia from the type are somewhat larger than those givenby Saccardo (24-30 x 4--6 ~m). The conidial state is bestshown on Berkeley's specimens but the conidiomata and hostsymptoms appear identical to those seen on the Saccardotype. The conidial state does not appear to be hyperparasiticand it seems plausible that the ascomycete may be theteleomorph of 5. varians. This would not be unusual as theteleomorph of 5colecosporiella fyphae (Oudemans) Petrak isknown to be Leptosphaeria typharum (Desm.) Karsten (Webster,1955).

5colecosporiella Petrak was redescribed by Sutton (1968) andthe genus now contains five species which, until the additionof 5. varians, was limited to monocotyledonous hosts.

Specimens examined: On Symplocus whitfordii, Mt Banajao, Provo ofLanguna, Luzon, Phillippines, Jan. 1913, M. Ramos, Flora of thePhilippines no. 19586a, holotype of Stagonospora varians, K; on


S. obtusa, Nuwara Eliya, Sri Lanka, 7000 ft, no. 592. ex Herb.Berk., K.

A large number of curators made this study possible bylending material for study. They are thanked for sendingmaterial from the following herbaria B, BM, BPI, CUP,DAOM, E, G, HCIO, K L, LISE, LISVA, LPS, M, NY, NYS,PAD, S, SIENA, TENN, TFM, UPS and W. Thanks are alsodue to Dr E. Punithalingam who, over the years, hasmaintained a proprietary eye on developments in Hendersonula,and has brought much useful information to our notice.

Miss Helen Simonelli has coped industriously and expertlyin typing successive drafts and due thanks to her areexpressed.

REFERENCES

ALLESCHER, A (1894). Einige hir das siidliche Bayern neueSphaeropsideen, Melanconieen und Hyphomyceten. Hedwigia 33,70-75.

ALLESCHER, A. (1901). Rabenhorst's Kryptogamen-[/ora von Deutsch

land, Oesterreich und der Schweiz 1 Abt. 6, 1-1016.ALLESCHER, A (1903). Rabenhorst's Kryptogamen-[/ora von Deutsch

land, Oesterreich und der Schweiz 1 Abt. 7, 1-1072.BERKELEY, M. J. & BROOME, C. E. (1873). Enumeration of the

fungi of Ceylon. Part II. Containing the remainder of theHymenomycetes, with the remaining established tribes of Fungi.Journal of the Linnean Society 14, 29-140.

BRODRICK, H. T. & VAN DER WESTHUIZEN, G. C. A (1976).Hendersonia creberrima, the cause of soft brown rot of mango inSouth Africa. Phytophylaetica 8, 13-16.

BUCHANAN, P. K. (1984a). Systemic growth of Ascochyta paspaliin paspalum. New Zealand Journal of Agricultural Research 27,

451-457.BUCHANAN, P. K. (1984b). Ascochyta paspali, a fungal parasite of

Paspalum dilatatum. New Zealand Journal of Botany 22, 515-523.BUCHANAN, P. K. (1987). A reappraisal of Ascochytula and

hcochytella (Coelomycetes). Mycological Papers (CMf) 156, 1-83.

CALAVAN, E. C. & WALLACE, J. M. (J954). Hendersonula toruloidea

Nattrass on Citrns in California. Phytopathology 44, 635-639.

CAMPBELL, C. K. (1974). Studies on Hendersonula toruloidea isolatedfrom human skin and nail. Sabouraudia 12, 150-156.

CAMPBELL, C. K., KURWA, A, ABDEL-AZIZ, A-H. M. &

HODGSON, C. (1973). Fungal infection of skin and nails byHendersonula tornloidea. British Journal of Dermatology 89, 45-52.

CHENG-GUO, W., BLANCHETTE, R. A & PALMER, M. A (1986).Ultrastructural aspects of the conidium cell wall of Sphaeropsis

sapinea. Mycologia 78, 960-963.CHESTERS, C. G. C. (1938). Studies on British Pyrenomycetes. II. A

comparative study of Melanomma pulvis-pyrius (Pers.) FuckeLMelanomma fuscidulum Sacc. and Thyridaria rubro-rotata (B. & Br.)Sacc. Transactions of the British Mycological Society 22, 116--150.

CLEMENTS, F. E. & SHEAR, C. L. (1931), Genera of Fungi. New York,U.S.A: Hafner.

COOKE, M. C. (1885). New British fungi. Grevillea 14, 1-7.DARKER. G. D. (1967). Revision of the genera of the Hypo

dermataceae. Canadian Journal of Botany 45, 1399-1444.DIEDICKE, H. (1914). Kryptogamen[/ora der Mark Brandenburg. Pilze

VII. Sphaeropsideae, Melanconieae, pp. 641-800.ELLIS, M. B. (1976). More Dematiaceous Hyphomycetes. Kew, U.K.:

CAB International Mycological Institute.ELLIS, M. B. & ELLIS, ]. P. (1985). Microfungi on Land Plants. London

and Sydney: Croom Helm.

487

FARLOW, W. G. (1876). The black knot. Bulletin Bussey Institute,

Botanical Articles 1, 440.FRIES, E. M. (1823). Systema Mycologicum 2, 1~21. Lund,

Sweden.FUCKEL, L. (1870). Symbolae Mycologicae. Jahrbuch des Nassauischen

Vereins fur Naturkunde 23-24, 1-459.FUCKEL, L. (1871). Symbolae Mycologicae. Jahrbuch des Nassauischen

Vereins fur Naturkunde 25-26, 289-346.FUCKEL, L. (1873). Symbolae Mycologicae. Jahrbuch des Nassauischen

Vereins fur Naturkunde 27-28, 3-99.GROVE, W. B. (1932). New or noteworthy Fungi. Part XII. Journal of

Botany (London) 70, 1-38.GROVE, W. B. (1937). British Stem and Leaf Fungi, 2. Cambridge,

UK: Cambridge University Press.HOHNEL, F. VON (1905). Dritter Nachtag zur Pilzflora des

Sonntagberges (N.-o) 1904. Verhandlungen der Zoologisch-botani

schen Gesellschaft in Wien 44, 600-621.HOHNEL, F. VON (1918). Dritte vorlaiifige Mitteilung mycolo

gischer Ergebnisse (Nr. 201-304). Berichten der Deutschen

botanischen Gesellschaft 36, 309-317.HOHNEL, F. VON (1920). Fungi imperfecti. Beitrage zur Kenntnis

derselben. Hedwigia 62, 56--89.HOHNEL, F. VON (1923). System der Fungi Imperfecti Fuckel.

Mykologische Untersuchungen u. Berichte 1, 301-369.HOHNEL, F. VON (1928). Ober Hendersonia meridionalis D. Sacc.

Mitteilungen aus dem Botanischen Institut der Technischen Hochschule

in Wien 5, 103-106.HOLM, L (l957). Etudes taxonomiques sur les Pleosporacees.

Symbolae Botanicae Upsalienses 14, 1-188.HUGHES, S.]. (1953). Conidiophores, conidia and classification.

Canadian Journal of Botany 31, 577-659.KHAN, A H. (l955). Some new diseases of Citrus in Pakistan.

Mycopathologia 10, 289-302.KOBAYASH!, T. (l962). A new species of Cryptodiaporthe on Sorbus.

Transactions of the Mycological Society of Japan 4, 6--8.KOBAYASHI, T. (1976). Taxonomic studies of Japanese Diapor

thaceae with special reference to their life-histories. Bulletin ofthe Government Forest Experiment Station, Meguro 226, 1-242.

MIGULA, W. (1921). Kryptogamen-flora von Deutschland, Deutsch

dsterreich und der Schweiz 3 Pilze 4 Tiel (1). Fungi imperfecti:Sphaeropsidales, Melanconiales. 1~14.

MIRZA, F. (1968). Taxonomic investigations on the ascomycetegenus Cucurbitaria S. F. Gray. Nova Hedwigia 16, 161-213.

MOORE, M. K. (l988). Morphological and physiological studies ofisolates of Hendersonula tornloidea Nattrass cultured from humanskin and nail samples. Journal of Medical and Veterinary Mycology

26,25-39.

MORGAN-JONES, G., NAG RAJ, T. R. & KENDRICK, B. (1972).leones generum coelomycetum. University of Waterloo Biology

series 7, 1-52.MOLLER, E. & v. ARX, J. A (1962). Die Gattungen der didymosporen

Pyrenomycete. Beifriige zur Kryptogamen[/ora Schweiz 11, 1-922.

NATTRASS, R. M. (1933). A new species of Hendersonula

(H. tornloidea) on deciduous trees in Egypt. Transactions of the

British Mycological Society 18, 189-198.NATTRASS, R. M. (1937). A First List of Cyprus Fungi. Nicosia:

Government of Cyprus.PAXTON, J. D. & WILSON, E. E. (l965). Anatomical and phy

siological aspects of branch wilt disease of Persian walnut.Phytopathology 55, 21-26.

PETCH, T. (l924). Revisions of Ceylon fungi. Part VII. Annals of the

Royal Botanic Garden, Peradeniya 9,119-184.

PETRAK, F. (l949). Beitrage zur Pilzflora Irans. Sydowia 3, 268332.


PETRAK, F. (1951). Uber die GaHungen Hendersollula Speg. undHendersonulina n. gen. Sydowia 5, 418-422.

PETRAK, F. (1961). Macrophoma sapinea (Fr.) Petr. in den Fiihrenwaldern des niederosterreichischen Waldviertels. Sydowia 15,309-316.

PETRAK, F. & SYDOW, H. (1927). Die Gattungen der Pyrenomyzeten, Sphaeropsideen und Melanconieen. Repertorium

specierum novarum regni vegetabilis 42, 1-551.PETRAK, F. & SYDOW. H. (1935). Kritisch-systematische Original

untersuchungen uber Pyrenomyzeten, Sphaeropsideen undMelanconieen. VI. Annales mycologici 33, 157-193.

PIROZYNSKI, K. A. & SHOEMAKER, R. A. (1970). Seirnatosporium

leaf spot of Ledum and Rhododendron. Canadian journal of Botany48, 2199-2203.

PUNITHALINGAM, E. & WATERSTON, J. M. (1970a). CM]

Descriptions of Pathogenic Fungi and Bacteria No. 273. Dip/odia

pinea.

PUNITHALINGAM, E. & WATERSTON, J. M. (1970 b). CM]

Descriptions of Pathogenic Fungi and Bacteria No. 274. Hendersonu/atoruloidea.

RIBALD), M. (1953). Necrosi corticale e aeterazione cromatica delligno prodaHe dal Pino austriaco da Phomatosphaeropsis pinicola n.gen. et n. sp. Annali deIIa sperimentazione agraria N.S. 7, 837-851.

SACCARDO, P. A. (1884). SyIIoge fungorum 3, 1-860.SACCARDO, P. A. (1893). Mycetes sibirici. Pugillus alter. Bolletino

della Societa botanica italiana 213-221.

SACCARDO, P. A. (1914). Notae mycologicae Series XVIII. Annalesmycologici 12, 282-314.

SACCARDO, P. A. & SYDOW, P. (1899). Sylloge fungorum 14,1-1316.

SIGLER, L. & CARMICHAEL, J. W. (1976). Taxonomy of Mal

branchea and some other hyphomycetes with arthroconidia.Mycotaxon 4, 349-488.

SMERLIS, E. (1970). Botryosphaeria laricis and its relationship toMacrophoma sapinea. Canadian journal of Botany 48, 1888-1901.

SPEGAZZINI, C. (1880). Fungi argentini. Pugillus III. Anales de laSociedad Cientz'fica, Buenos Aires 10, 122-142.

SPRAGUE, R. (1950). Diseases of Cereals and Grasses in North America.

New York, U.s.A.: Ronald.STEVENS, F. L. (1913). Plant Disease Fungi. U.s.A.: Berwick &

Smith.STEVENS, F. L. (1919). The Fungi which Cause Plant Disease. New

York u.s.A.: Macmillan.

(Received for publication 31 August 1988)

488

SUTTON, B. C. (1968). KeIIermania and its generic segregates.Canadian Journal of Botany 46, 181-196.

SUTTON, B. C. (1975). Coelomycetes. V. Coryneum. MycologicalPapers (CMf) 138, 1-224.

SUTTON, B. C. (1980). The Coelomycetes. Kew, UK: CAB International Mycological Institute.

SUTTON, B. C. & DAVISON, E. M. (1983). Three stromaticcoelomycetes from Western Australia. Transactions of the BritishMycological Society 81, 291-301.

SUTTON, B. C. & SELLAR, P. W. (1966). Toxosporiopsis n. gen., anunusual member of the Melanconiales. Canadian Journal of Botany44, 1505-1513.

SYDOW, H. (1923). Novae fungorum species. XVIII. Annalesmycologici 21, 342-344.

SYDOW, H., SYDOW, P. & BUTLER, E. J. (1916). Fungi Indiaeorientalis. Pars V. Annales mycologici 14, 177-220.

TASSI, F. (1899). Novae micromycetum species descriptae et iconibusillustratae. Bolleltino del Laboratorio ed Orto Botanico, Siena 2,103-105.

TASSI, F. (1902). I generi Phyllosticta Pers., Phoma Fr., Macrophoma

(Sacc.) Berl. & Vogl. e i loro generi analoghi. giusta la legged'analogia. SoIIeltino del Laboratorio ed Orto Botanico 5, 1-72.

TOMMERUP, I. C. & LANGDON, R. F. N. (1970). Morphologyand development of Hendersonula monochaetiellae sp. nov. Transactions of the British Mycological Society 55, 477-482.

TRACY, S. M. & EARLE, F. S. (1895). Mississippi fungi. MississippiAgricultural and Mechanical College Experiment Station Bulletin 34,8-124.

WEBSTER, j. (1955). Graminicolous Pyrenomycetes. V. Conidialstates of Leptosphaeria michotii, L. microscopica, Pleospora vagans andthe perfect state of Dinemasporium graminum. Transactions of theBritish Mycological Society 38, 347-365.

WEHMEYER, L. E. (1941). A revision of Melanconis, Pseudovalsa.Prosthecium, and Titania. University of Michigan Studies. ScienceSeries 14, 1-161.

WELCH, D. S. (1926). A monographic study of the genus Cucurbitariain North America. Mycologia 18, 51-86.

WESTENDORP, G. D. (1859). Sur quelques cryptogames inedites ounouvelles pour la Flore Beige. BuIIetin de /'Acadimie r. de BelgiqueN.S., 7, 77-94.

YOUNG, H. E. (1948). Hendersonula agathi n. sp., the cause of leafcast of Kauri Pine in Queensland. Queensland journal of AgriculturalScience 5, 9-12.

Documents

Revision of Hendersonula