JAMES REID Laboratory of Forest Patltology, llIaple, Ontario

AND

COLIN BOOTH Co~12111o1~zvea1th ;IIycological Institz~te, R e w , Sz~rrey, England

Received September 27, 1965

Abstract The genus name Flageoletia (Sacc.) Hohn. is shown to be a synonym of the

penus narrie Plronzatospora Sacc. Three of the four described species, Flageoletia leptnsca (Pk. & G. LV. Clint.) Hbhn., Flageoletia tenz~is (Sacc.) Ilohn., and Flageoletia relr~ilialza Hohn. are synonymous and assigned to Phontatospora leptasca (Pk. & G. LV. Clint.) J. Reid & Booth conib. nov. The fourth species, Flageoletia balrrbusina Syd. represents an undescribed genus and is herein desig- nated nTeoflageoletia ba?ilbztsina (Syd.) J . Reid & Booth, comb. nov., the type species of the new genus, Neoflageoletia J. Reid & Booth.

Historical Peck and Clinton (1878) described Valsa leptasca P. & C., an organism

\vhicll had been found growing on dead branches of Rhus typhina L. near Buffalo, New York. Saccardo (1885) transferred this species to the genus Cryptosporella Sacc., making the new coillbination Cryptosporella leptasca (P. & C.) Sacc. Cool~e (1885), who recognized Cryptosporclla Sacc. only as a subgenus of Valsa Fr., designated it 17aIsn (Cryptosporella) leptasca C. & P. Ellis and Everhart (1892) regarded Cryptosporella Sacc. as a section of the genus CryPtospora Tul. and made the new cornbillation Cryptospora Ieptasca (C. & P.) Ell. & Ev.

Later, Saccardo (1896) erected Flageoletia as a subgenus of Cryptosporella based on an organisill collected on Coryll~s avellana L. near Rigily in France, \vhich he cited thus: iiCryptosfiorelba (Flageoletia) leptasca (P. & C.) Sacc. *C. ten.1li.s Sacc." This was described as differing from the type in the long- stallied asci. Oudeinans (1920) subsequently designated this second organism Crypto.sporella leptnscn (P. 8 C.) Sacc. subsp. t e n ~ ~ i s (Sacc.) Oud.

Iiehm (1913) identified a fungus collected on P r l ~ n z ~ s spinosa L., gro\ving near \Vellia/NIal~risch-Weissl~ircl~en, Czechosloval;ia, as C. leptasca var. fclrl~is and issued it as n~~ll lber 2033 ol his Asconlycetes exs., Fasc. 52. noted that it "Stimmt zehr g ~ ~ t zu cler Beschreibung und zu Flageoletia wegeil cler zj,Iindrischen Schlauche L I I I ~ zylindrischen zwei Oltropfeil enhaltenden, I-reihigen sporen."

Hohnel (1916) raised Flngcolctia to the generic level, as Flageoletia (Sacc.) Hohn., and stated that it was not a illember of the 31elanconidaceae, but related to 17alsaria. I-Ie then listed three, what he termed, closely related species: (1) Flngeoletia lcptasca (C. & P.) I-Iohn. on R. typhina, (2) Flageoletia telzl~is (Sacc.) I-Iohn. on C. nvcllann, and (3) Flngeolctia rehrniana I-Iohn. on P. spdnosa, for which no validating diagnosis was given. Subsequeiltly I-Iohnel

'Joint contribution from the Department of Forestry of Canada, and the Commonwealth llycological Institute, I<ew, Surrey, England.

Cariad~ar~ Journal of Botan~,. \rolume 44 (1966)

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446 CANhDI.AS J O U R N A L O F BOT.AST. VOL. 44, 1966

(1918) gave a description of F. rehmiana and stated it was based on the fungus issued in Rehm's Ascom~-cetes eus. as Xo. 2033. He also provided the iollowing generic diagnosis.

I.'lageoletia (Sxcc.) I-161.1~. Stroma erumpent, not valsoid. Perithecia irregularly arranged, 114th

papillate ostioles (without long necks). Paraphyses numerous, typical. Asci cylindric, stalked. Spores uniseriate, hyaline, one-celled, oblong, small. Imperfect state unlcnown. i\/lelogrammaceae. I-Ie suggested that Crypfosporelba z~mbilicata (Pers.) Berl. c% Vogl. and Cryptosporella innata (Berli. & Curt.) Sacc. might also be species of Flageoletia (EIohneI 1916, 1915).

Sydow (1832) added a fourth species t o the genus, flageobetia barnblrsi.na Syd., a fungus found on dead stems of Bamblrsa sp. in the Philippines.

FIG. 1. P l ~ o ~ ~ ~ a t o s p o r a berkleyi. From HOLOTIPE, I<. ,-\sci and ascospores. FIG. 2. Flageoletia leptasca. Fro111 HOLOTYPE, SYS. Asci and ascospores. FIG. 3. Flageoletia te t~uis . Fro111 HOLOTYPE, PAD. Asci and ascospores. FIG. 4. Flageoletia rel~minna. From Rehm's Xsco. ess. So . 2033, Herb. Sacc., P-AD.

Asci a11d ascospores.

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KEII) X S D BOOTII: FLAGEOLI.:TI.-1 447

Taxonomy Flageoletin leptasca (Plt. & G. W. Clint.) Igohn., Ost. Bot. %. 66: 57. 1916.

IFigs. 2 and 7. Authority given as (C. cG P.) by Hohnel. ITalsn leptasca P. & C. Rept. N.\-. St. Bot. 29: 59. 1878. TTalsu (Cryptosporella) leptasca C. & P. (Cool~e?), Grevillea, 14: 53. 1885. Cryptospora ieptasca (C. & P.) Ell. & Ev., K. ;lm. Pyr. p. 532. 1892. CryptosPorella (Flageoletia) leptasca (P. & C.) Sacc., Bull. Soc. Mycol. Fr.

12: 64. 1896. 'I'he origiilal authorities for this fungus are given as P. Sr C., but have

frequently beell changed to C. & P. In additioil the C. has often beell taken to mean I I . C . Coolte. I-lowever, the origiilal collectio~l was made by G. W. Cliilton and is listed under his contril~utions in the 'I'wenty-Ninth Report of the S e w Yorl; State Museum, page 37. Furthermore, no collection of this fungus is to be found in the herbariun~ of the Royal Botanic Gardens, I k w , nor is there any evidence that one was ever present there. 'I'hus it is illost ~~nlil;ell, that Coolte ever had anything to do with this species and Clinton should be considered the joint author.

Examination of the type speciilleil of V. leptasca indicated that it coilsisted of a species of Phomatospora Sacc. parasitic on, or siinply growing on or in the stroina of a pyrenomycete. 'I'his is clearly shown in Fig. 7, where the remains of part of the strorna can be seen below.

The discolored, blacltened margin bounding the stroina of the originaI fungus on the host plant suggests this may have been a Diaporthe sp.

Coinparison of sections and squash inounts prepared froin the holotype of Phomatospora berkleyi Sacc. (Figs. 1 and 6) with sections and squash inounts

FIG. 5 . lVeofEageolelia batizbzrsina. From ISOTYPE. I M I . Asci and ascospores.

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4-1-8 CASADI.\S JOURSAL O F BOTt\NY. VOL. 44, 1966

prepared from the type of V. leptasca (Figs. 2 and 7) leave no doubt that 1'. Ieptasca is a Phomato~pora sp. and, except for the larger perithecia, it agrees closely with P . berkleyi. In V. leptasca the perithecia vary between 210 and 368 p in diameter, while the perithecia measured froin the type of P . berkleyi were rarely more than 180 p in diameter. Arx and Miiller (1954) gave the perithecial measurements of P . berkleyi as 140-210 p. Thus, in spite of rather similar asci and ascospores, V. leptasca is considered a distinct species of Phomatospora and it is redescribed as Phomatospora leptasca (PI;. & G. \T'. Clint.) J . Reid 8 Booth, comb. nov.

Specimens examined: Cryptosporella leptasca Sacc., Buffalo, S .Y. , leg. G. W. Clinton, ( Valsa leptasca P. & C.), on Rhus, NYS2 (IIOLOTTPE).

Flageoletia tenz~is (Sacc.) I-Iijhn., Ost. Bot. Z. 66: 57. 1916. Figures, this paper 3 and 8, and Sacc., Bull. Soc. Mycol. Fr. 12: 1896, Plate V, Fig. 2.

Cryptosporella (Flageoletia) beptasca (P. & C.) Sacc. *C. tenz~is Sacc., Bull. Soc. Mycol. Fr. 12: 64-65. 1896.

Cryptosporella leptasca (P. & C.) Sacc. subsp. tenllis (Sacc.) Oud., En~um. Syst. Fungorum 2: 354. 1920.

Based on examination of the type and comparison of sections of this species with those of V. leptasca (compare Figs. 2 and 7 ~vi th 3 and S), this f~ungus is undoubteclly identical with 1.'. leptasca.

The black line bounding the stroma clearly indicates it is also parasitic on, or growing on, a pyrenomycete similar to that encountered in the North American material.

Specimeils examined: Cryptosporella (Flageoletia) leptasca (P. & C.) var. tenz~is, on Corylus avelanae, Rigny, France, Saccardo I-Ierb., PAD (I-IOLOT~PE).

Nageoletia rehmiana I-Iohn., Ann. Mycol. 16: 126. 1918. Figs. 4 and 9. Cryptosporella (Flageoletia) leptasca (P. & C.) Sacc. var. t e n ~ ~ i s Sacc. serLslL

Rehrn, in Rehm, Ann. Mycol. 11: 167. 1913. Cryptosporella leptasca (P. & L.) Sacc. var. valsoides Rehm, a noillen dubium

in Petrak's Flora PIoravica exs. (no number). Based on an examination of sections and preparatioils froin txvo copies of

Rehm's Ascornycetes eus. No. 2033 and comparing them wit11 those from the type of V. leptascn (coinpare Figs. 2 and 7 with 4 and 9), this is also identical with V. leptasca.

Specimens examined: (1) Cryptosporella (Flageol~tia) leptasca (P. & C.) Sacc. var. tenzlis Sacc., Rehm Ascom. exs. No. 2033, on Prunus spinosa (BPI and Saccardo I-Ierb, PAD); (2) Cryptosporella leptasca (P. cG L.) var. valsozdes Rehin (a nomen dubium), Petrak Flora Moravica eus. (no number) two copies BPI (apparentlv part of the same collection as that distributed by Rehm); (3) Flageoletia rehmiana I-Iijhn. 11. sp., Petrak, Flora Moravica (no number), I< (apparently part of the saille collection as that distributed by Rehm).

Phomatospora leptasca (Pl;. & G. W. Clint.) J. Reid & Booth, comb. no\-. Valsa leptasca P. 8 C., Rept. N.Y. St. Bot. 29: 59. 1878. Valsa (Cryptosporella) leptasca C. & P. (Coolie?), Grevillea, 14: 53. 1885.

2Herbariun~ designations are those of Lanjouw and Stafleu (1961).

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REID A S D BOOTH: FL.4GEOLETI.l 419

Cryptospora leptasca (C. & P.) Ell. Sr Ev., N. Am. Pyr. p. 532. 1892. Cryptos;horella (Rageoletia) leptasca (P. & C.) Sacc., Bull. Soc. Mycol.

Fr. 12: 64. 1896. Cryptosporella (Flageoletia) leptasca (P. & C.) Sacc. *C. tenliis Sacc., Bull.

Soc. NIycol. Fr. 12: 64-65. 1896. Cryptosporella (Flageoletia) leptasca (P. Sr C.) Sacc. var. fenmis Sacc. sensu

Rehm, in Rehm, Ann. Mycol. 11: 167. 1913. FLageoletia leptasca (C. & P.) I-Iohn., Ost. Bot. Z. 66: 57. 1916. Flageoletia tenuis (Sacc.) Hohn., Ost. Bot. 2. 66: 57. 1916. Flageoletia rehmiana I-Iohn., Ann. Mycol. 16: 126. 1918. Cryptosporella leptasca (P. & C.) Sacc. subsp. tenliis (Sacc.) Oud., Enum.

Syst. Fungoruln 2: 354. 1920. Cryptosporella leptasca (P. & L.) Sacc. var. valsoides Rehm, a norneil clubiurn

in Petrak's Flora AIIoravica exs. (no number). Perithecia einbedded in the circular to effuse stroinn of an old pyrenoinycete;

smooth, blacli, shiny, with a short conic ostiole up to 150 ,u wide a t its base and 135 ,u high which is excessively thicliened tolvards its tip and with the cells beconling occluded and heavily carbonized; not collectively erumpent, periphysate. They are oval to occasionally flattened through mutual pressure when strongly clustered; 250-400 X 200-300 ,u; paraphyses present.

Asci unitunicate, long cylindric, rounded above with a thic1;ened apex, a i d a t times, a rudiinentary apical structure consisting of two sinall, indistinct refractive bodies lying beside one another; not blueing in NIelzer's reagent; tapering below to a long narrow stipe, eight-spored; 3.00-135 X 3.5-4.5 ,u (pars sporifera 65-80 ,u, stipe 35-45 ,u long, narrowing to 0.5 ,u).

Spores one-celled, hq-aline, oval, biguttulate; 6.5-8.5 X 2.5-3.5 ,u; uniseriate in the ascus.

I-Iost: Associated with the strollla of pyrenoinycetes growing on various woody hosts.

There is no justification for placing C. z~mbilicata or C. innata in the salne genus as P . leptasca as suggested by I-Iohnel (1916, 1918).

The specilnen of C. innata in the herb:~riun~ of the Royal Botanic Gardens, Kew, as Sphaeria (versafilis) innata B. & C. on Castanea vesca, R'Iountains, N . Y . , No. 4, 496 Berliley I-Ierb., HOLOTYPE is an Anthostoma sp. and thus the organisin has nothing to do with Flageoletia or Phonzatospora.

The speciinen of Sphaeria zimbi1,icata Pers. in the Persoon herbarium a t Leiden consists of old, enlpty perithecia einbedded in a black carbonaceous stroma. No spores were seen, but the ostioles appear to be sulcate, suggesting a ~neinber of the Diatrypaceae. 4 second collection in the Persoon herbariuln as Sphaeria zlmbilicata P 3Iederae consists of scattered superficial to slightly e~nbedcled fruiting bodies \vithout spores. As a consequence this naine is a nomen confusuin and should be dropped.

Flageolet,ia bambusina Syd., Ann. Mycol. 30: 96. 1932. This last species to be placed in Rageoletia bears no relationship to the

foregoing species or the diagnosis of Flageoletia. I t represents an undescribed genus and as Flageoletia (Sacc.) Hohn. nlust be considered a synonym of Phomatospora Sacc., it is proposed as the type species of a new genus Nee- flageoletia J. Reid & Booth.

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450 C.-\ShDIAN JOURSAL OF BOTANY. VOL. 44, 1966

Neoflageoletia J . Reid & Booth, gen. nov. (order Sphaeriales, Family Pol~r- stigmataceae) Figs. 5, 10, and 11.

Perithecia in stromate sub epidernlide collocata; aggregata vel ordinibus disposita; collis brevibus, conicis vel lateribus parallelis. Ascis unitunicati, cylindracei vel cylindraceo-clavati, caulibus brevibus et sessilibus, 8-spol-i. Xscospori 7 cellulis, fusiforines vel lateribus a l iq~~ando non aequis, hyalini.

Perithecia embedded in a subepidermal s t ro~na; clustered to linearly ar- ranged; beaks short, conical to parallel-sided; asci unitunicate, cylindric to cy- lindric-clavate, short-stallred, remaining attached a t their base: 8-spored; asco- spores hyaline 2-celled, spindle-shaped to fusiiorrn to occasionally slightly inequilateral.

Type species: N. bnnzbusina (Syd.) J . Reid & Booth. Et!~mology: From TzeoJ = new + the generic name Flageoletia.

Neoflageoletia bambusina (Syd.) J. Reid & Booth, comb. nov. Figs. 5, 10, and 11.

Flageoletia barnbusinu Sydow, Ann. Mycol. 32: 96. 1930. The embedded stronia gives the substrate a blackish-grey color; it develops

under the epidermis and frequently penetrates three or four cell layers deeper into the underlaying cortical tissue; it is particularly well developed around the perithecia where it forins a loose to compact mass of subhyaline to dark- brown hyphal elements 2-4 p thick which separates the epiderinis from the underlying tissue to forin the elongate pustules on the host surface.

Perithecia embedded in the pustulate areas of the stroma, arranged in short, parallel rows, or irregularly clustered; inore or less spherical or flattened froin below and usually lying in the stroina between the epidermis and the under- lying host parenchyma; 80-160 p diameter; when aggregated they are fre- quently flattened laterally by mutual pressure and the walls may fuse. Walls up to 15 p thick, of two layers; outer layer of dark-brown, globose to slightly elongate, thick-walled cells 3 to 5 p long and 3 p wide; inner layer of thin- walled, elongate, strongly compressed cells. Beaks, short, not converging, parallel-sided or conic to occasionally flaring a t the ostiole, up to 60 p high and 60 p wide; formed of globose to elongate, thick-walled cells; central canal lined with hyaline, septate, thin-walled periphyses.

Asci unitunicate eight-spored, cylindric to cylindric-clavate, occasionally tapering slightly a t both ends, thickened and rounded above, tapering to a short stall; below and remaining attached to the perithecial wall; 50-75 X 6-8 p ; ascus tip not blueing in iodine. Paraphyses lacking or gelatinizing earlv.

Spores elongate spindle-shaped to fusiforin, straight to only occasionally inequilateral and then appearing slightly curved; tapering slightly towards the rounded ends; irregularly biseriate in the ascus; finally two-celled with a median septum ; 8-14 X 2-3 p.

Specimens examined: (Flageoletia bambz~sina) on dead stems of Bamboo, Mt. Maquiling, Laguna province; 37. 111. 1920; leg. E. Quisumbing No. 9153. IMI 15,069, ISOTYPE.

Sydow (1932, p. 97) stated that this material, though very abundant, was rather immature, and he could not be sure whether the ascospores finally

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Frc.. 6. Plrol?rnlospora beikleyi. From HOLOTYI'E, I<. Tran5versc section ol a pcriiheciu111. X270. . .-

F I ~ ; . 5. FI(~ltrgeolel~ir1 leplnscn. From Hor.o.ryr,ri, SYS. ?'ransversc scction ol a pcritlicci~~m showi~ig its relationship to the stroma of thc pyrcnom).cetc on \vhich it grcw.

FIG. 8. Flageolelia t e ~ ~ l r i s . Fro111 Iror.oTurlli, I':\I). ?'rans\rcrse scction of scvcral per- ithccia shoming thcir rcla~ionship to thc stroma of the pyrenomycetc on \vllich thcy grew. X63.

FIG. 9. Flageoletin rel~rrlia?en. From Rchm's Asco. ess. No. 2033, Herb. Sacc., I'AD. 'Tr;~ns\:crsc scction of scvcral pcrithccia showing their rclatioliship to thc stroma of the pyrenomyccte on \vI~ich they grew. X63.

Rcid and Booth-Can. J. Botany

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FIGS. 10 ancl 11. Neqflageolelia ba~~lbtr.si?~a. From ISOTYPE, IMI. Transvcrsc sections through clusters of perithecia. X63.

Reid and Booth-Call. J. Botany

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REID .AND BOOTII: FL.AGEOLET1h 45 1

became two-celled. I-Iis placing of it in Flageoletin was coilditional on the spores remaining one-celled.

-4s abundant two-celled spores were observed, there can be no doubt that this fungus is not a species of Phomatospora and, because of the nature of the stroma and the lacli of a blue pore reaction in iodine, must be placed in the l'olystigmataceae as a new genus.

Acknowledgments The authors thanlc Dr. R. W. G. Dennis, Royal Botanic Gardens, I<ew;

Professor C. Cappelletti, University of Padova; Mr. S. J. Smith, New Yorlc State NIuseuin, Albany, New York; and Dr. C. I<. Benjamin, Tlle Kational Fungus Collections, Beltsville, h/Iaryland, lor the loan of specimens under their care. \;lie also thank Miss S. Daniels for preparing the Latin diagnosis.

References vos ~ \ R s , J. A. and MULLER, E. 1954: Die Gattungen der amerosporen Pyrenomyceten.

Beitr. ICryptogamenflora Schwelz. 11(1), 1-434. COOKE, M. C. 1885. Synopsis Pyrenornycetum. Grevillea, 14, 46-56. ELLIS, J. B. and EVERHART, B. &I. 1892. The North American Pyre~~onlycetes. Ellis and

Everhart, Newfield, New Jersey. I-IGHNEL, F. 1916. Mykologisches XXIV Vorlaulige NIitteilungen. 0sterr. Botan. %. 66,

51-60. -- 1918. Mykologische Fragmente 254. Uber Valsa leptasca C. Pc P. Ann. i\lycol. 16,

125-127. LANJOUW, J. and STAFLEU, F. A. 1964. Index Herbar ior~~~n. Part I. The herbaria of the world.

5th ed. Regnum Vegetabile, 31, 1-251. OUDEMANS. C. A. I. A. 1920. Enunleratio Svstematica Funeorum. Vol. 2. Hacrae. - - - -

PECK, C. H. ~ I ~ ~ ~ C L I N T O N , G. \V. 1878. keport of the ~ ; a t e Botanist. Iieupt. N.Y. State Botanist, 29, 29-82.

REHM, H. 1913. Ascornycetes exs. Fasc. 52. Ann. hIycol. 11, 166-17 1. SACCARDO, P. A. 1882. Sylloge fungorum. Vol. 1.

1896. Notes rnycologiques. I. Fungi nevi gallici, germanici et capenses. Bull. Soc. Mvcol. France. 12. 64-69.

189G. Sylloge fungo;um. Vol. 14. Sy~om, H. 1932. Novae fungoruru species-XXI. Ann. Mycol. 30, 91-117.

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