ANNALES MYCOLOGICIEDITI IN NOTITIAM

SCIENTIAE MYCOLOGICAE UNlVERSALIS

HERAUSGEGEBEN UND REDIGIER'l'VON

H. SYDOW~'TR MITWIRKUNG vo~ ABATE J. BRESADOLA (TRIENT), PROFESSOR DR. FR. CA V ARA

APEL), PROFESòOR DR. P. DIETEL (ZWICKAU), DR. A. GUILLIERMOND (LYON),BOPBSSORDB.E.KÜSTER(G IESSEN),PROFESSOH Du.RENÉ MAIRE(ALGER), DR.F.PETRAK.. R:-WEISSKIRCHE:-),E.S. SALMON (WY E, NEAR ASHFORD, KENT), DR.A.SARTORY

ANCY)j .PBOFESSOR Dll. P. VUILLEMIN (NANCY), Dll. A. ZAHLBRUCKNER (WIEN)UND ZAHLREICHEN ANDEREN GELEHRTEN

DREISSIGSTER JAHRGANG - 1932

.BERLIN

VERLAG VON R. FRIEDLAENDER & SOHN

lÐ32

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In halt (Band XXX).

i. Originalarbeiten.

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B 0 e d i j n, K. B. Einige Bemerkungen zu del' Abhandlung von S.C. Teng, "Fungi of Nanking I" . . . . . . . . . . . . . . . 4í8

C i fer l' i, R. The criteria for definition of species in mycology . . 122G l' ego 1', Mar y J. F. A study of heterothallism in Oeratostomella

pluriannulata, Hedgcock . . . . . . . . . . . . . . . . . .. 1

G l' ego 1', Mar y J. F. Observations on the structure and identityof Tulasnella anceps Bres. et Syd.. . . . . . . . . . . '.' . . 463

G y e i n i k, V. Enumeratio lichenum europaeorum novorum rario-rumque. I .... . . . . . . . . . . . . . . . . . . . . .. 442

G y e i n i k, V. Über einige Arten del' Gattung' Parmeliopsis (Stizenb.)NyL. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 456

M a i k 0 v s k ý , K are i M. Über die europäischen Arten del' GattungPanus . . . . . . . . . . . . ~ . . . . . . . . . . 10

P a 1m, B. T. On Cyttaria Berk. and Cyttariella n. gen. . 405P a 1m, B. T. Biological notes on Albugo. . . . . . 421Pet c h, T. Some Philippine Entomogenous Fungi. . 118Pet c h ,T. Gibellula. . . . . . . . . . . . '.' . . . 386P 6 t l' a k, F. und C i fer l' i, R. Fungi dominicani. II 149Pi i á t, A i bel' t. Species nova generis Ganoderma Karst. e yulcano

Kilmandjaro: Ganoderma Baumii sp. n.. . . . . . . . . . . . 460

Pi 1 á t, A. et Ve s e 1 ý, R. Species nova vernalis generis Tricholoma:Tricholoma Kavinae ...................... 476

Po eve l' lei n, H. 'Die Gesamtverbreitung del' Uropyxis sanguinea

in Europa . . . . . . . . . . . . . . . . . . . . . . . . '.' . 402

Sartory, A., Sartory, R. et Meyer, J. Etude d'un nouveau

Scopulariopsis: Scopulariopsis gryll n. sp.. . . . . . . . . . 466

Sartory, A., Sartory, R. et Meyer, J. Etude d'un nouveau

champignon du genre Fusarium: Fusarium eucheliae . . . . . 471

S a v u Ie s c u, T r. et Ray s s, T. Nouvelle contribution à la con-naissance des Péronosporacées de Roumanie . . . . . . . . . 354

S y d 0 w , H. Fungi chileuses a cl. E. Werdermann lecti. Pars secunda 81By d 0 w, H. Novae fungorum species - XXI . . . . . . . . . 91

By d 0 w. Mycotheca germanica. Fasc. L-LII (no. 2451-2600) 394Z a h i b r u c k 11 e 1', A. Neue Flechten. -- XI . . . . . . . . . 427

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Burg U. 1\1.

Some Philppine Entomogenous Fungi.By T. Pet c h.

Through the kindness of Dr. Sydow, I have had the opportunity ofexamining numerous specimens of entomogenous fungi on leaves, collectedby Mary Strong Clemens in the Philippines. Though the collection con-sisted of seventy-two numbers, it contained only nine species, and with

one exception (Aschel'sonia Coffeae P. Henn.), these are all parasitic onAleyrodidae. Except where otherwise stated, the localities cited are inLuzon. The list may serve to give some idea of the relative abundance ofthe diffei;ent species of Aschersonia parasitic on Aleyrodidae in that country.

The chief interest of the collection lies in the occurrence of many spe-cimens of a new species, which is described below as Aschersonia phi-lippinensis, and its perithecial stage, Hypocrella philippinensis. This hasbeen known to OCCur in the Philippines for many years, but the specimens

collected previously have never been in a fit condition for description.The species is remarkable in that it often forms flat, circular, tomentose Orradially fibrilose, white or cream-coloured stromata, up to 7 mm diameter,with a fimbriate margin, without any pycnidia or perithecia. With the help

of the present specimens, it is now possible to refer the following previous

gatherings to Aschel'sonia philippinensis, - On Schizostachyuni, N Olzag'aray,

Provo Bulacan, May 1910, Rb. Bureau of Science, no. 10838; on Astroniasp., Mt. Maquiling, Luzon, September 1910, Merrill no. 7150; on Astronia,l\it. l\iaquiling, February 1914, C. F. Baker, Fungi Malayana, no. 8; onDerris heptaphylla, Lamao, H. A. Lee, no. 59. Some of these specimens

were referred to under Hypocrella Molli in the G e n era H y p 0 c r e II aand Aschersonia, Annals of the Royal Botanic Gardens,Per ad e n i ya, VII, 240. Immature specimens resemble immature Hypo-

crella Mollii, but are more tomentose and usually have a fimbriate margin.In general, the stromata of Aschersonia philippinensis are at first plane

and circular. Subsequently they develop, in the centre, a gToup of co-

lumnar tubercles, each of which bears a single, widely open, circular pyc-nidium at the apex. When, however, the tubercle is oblique, the pycnidiumopens obliquely and the orifice is usually vertically compressed into anarrow slit. In some stromata, the centre is occupied by a raised disc upto 2 mm in diameter, instead of a group of tubercles, and the pycnidiaopen on the vertical face of the disc, their orifices being overhung by itsupper edge. Other stromata may have neither tubercles nor disc, the pyc-nidia occurring in the stroma, vertical or oblique, the orifice being

surrounded by a slightly raised rim.

In the dimempressed orificeresembles the As

ture of the pycndisc is lacking l'nidia are not reg

Hypocrella ph

flavis, planis, teusque 3 mm diacylindraceis vel c

tomentosis, sing);ampullaceis, 450

ascis 240¡. longisporarum angiistiAleyrodid on und

1923, no. 3144; è

January 1924, ne

Petch in the sam

Cebu Island, MayAschersonia p

fJavis, planis, tenfimbriato, centro 1

0,25 mm alt., rect:tibus; vel strom:

margine disci; VE

marginatis. Pycr

rotundatis vel coi

fusoideis, fine actThe pycnidial stRourea erecta (El

Also no. 217, (D.ry 1923; no. 72(

Memecylon, loco ;no. 806, on unknoknown leaves, loc.Pampanga Prov.,Za.mbales Prov., 1

Merr., loc. cit., ~no. 2440, on unk? Guioa, loco cit., jSolana, Cagayangayan Prov., J anTiiguegarao, Cag:

Asingan, Pangasi

:ne EntomogenOusFungi.:y T. Pet c h.

'1'. Sydow, I have had the opportunity ofof entomogenous fungi on leaves, collected~ Philippines. Though the collection con-it contained only nine species, and with

'feae P. Henn.), these are all parasitic on

ierwise stated, the localities cited are inve some idea of the relative abundance of

ia parasitic on Aleyrodidae in that country.iction lies in the OCcurrence of many sp~-is described below as Aschersonia phi-

;age, Hypocrella philippinensis. This hasJpines for m.'ny years, but the specimens

been in a fit condition for descriptio~.

it often forms flat, circular, tomentose orcoloured stromata, up to 7 mm diameter,ny pycnidia or perithecÏa. With the helpT possible to refer the fOllowing previousensis,-On Schizostachyum, Norzagaray,

'eau of Science, no. 10838; on Astroniabel' 1910, iVlerril no. 7150; on Astronia,F. Baker, Fungi Malayana, no. 8; on

Lee, no. 59. Some of these specimens

M ollii in the G e n era H y p 0 c r e ii aif the Royal Botanic Gardens,re specimens resemble immature Hypo-ie and usually have a fimbriate margin.ersonia philippinensis are at first planeevelop, in the centre, a group of co-

il'S a single, widely open, circular pyc-the tubercle is oblique, the pycnidium

usually vertically compressed into a~entre is occupied by a raised disc upgroup of tubercles, and the pycnidiai, their orifices being overhung by itsTe neither tubercles nor disc, the pyc-

rtical or oblique, the orifice being

Some Philppine Entomogenous Fungi.119

In the dimensions of itspycnospores and paraphyses, and in. the com-pressed orifice of some of its pycnidia, Aschersonia philippinensis

resembles the Aschersonia of Hypocrella tubulata, but the internal struc-ture of the pycnidium is different. Specimens in which the tubercles ordisc is lacking resemble forms of Aschersonia placenta in which the pyc-nidia are not regularly arranged.

Hypocrella philppinensis Petch, n. sp. - Stroniatibus albis vel palldo-

fIavis, planis, tenuibus, interdum margine fimbriato vel membranaceo,usque 3 mm diam., tubercula spars

a vel conferta ferentibus; tuberculiscylindraceis vel ovoideis vel subglobosis, usque 0,5 mil alt., 0,35 mm diam.,tomentosis, singulo perithecio in quoque tuberculo; peritheciis angusto-ampullaceis, 450 .¡, alt., 150 ¡, diam., ostiolis minutis, impressis, flavis;ascis 240 '¡. long'is, 8 ¡. diam., cylindraceis, capitatis, octosporis; articulissporarum angnsto-ovalibus, rectis v81 curvatis, 8-12 ¥ 2-2,5 ¡,. On anAleyrodid on undetermined leaves, Asingan, Pangasinan Prov., November1923, no. 3144; ditto, on Apocynaceae, Bambàng, Nueva Vizcaya Prov.,January 1924, no. 3458; in both cases with Aschersonia philippinensisPetch in the same or separate stromata. Also no. 6214, on Ficus, Cebu,

Cebu Island, May 1924.

Aschersonia pliilppinensis Petch, n. sp. Stromatibus albis vel palldo-

flavis, planis, tenuibus, tomentosis, UJsque 8 mm diam., margine saepiusfimbriato, centro tubercula sparsa vel conferta, cylindracea, 0,36 mm diam.,

0,25 mm alt., recta vel obliqua, singulo pycnidio in quoque tuberculo, fereii-tibus; vel stromatibus centro' discoideis, ostiolis pycnidiorum obliquismargine disci; vel stromatibus omnino planis, pycnidiis sparsis immersismarginatis. Pycnidiis concavis, 0,25 mm diam., 0,15 mm aItis, ostiolisrotundatis vel compressis, sporis coacervatis flavis. Pycnosporis angusto-fusoideis, fine acutis, 9-11 ¥ 1,5-2¡.. Paraphysibus usque 150 ¡, longis.The pycnidial stage of Hypocrella philippinensis. On Aleyrodidae. OnRourea erecta (Blco.) Merril, Isabela Prov., December 1923, no. 2921.

IAlso no. 217, on unknown leaves, Mt. Arayat, Pampanga Prov., Febru-:Jry 1923; no. 720, on Lunasia amara, loco cit., April 1923; no. 744, onMemecylon, loc. cit., April 1923; no. 785, Eugenia, loco cit., April 1923;

no. 806, on unknown leaves, Orion, Bataan Prov., May 1923; no.

834, on un-known leaves, loc. cit., March 1923; no. 2199" on unknown leaves, Stotsenberg,Pampanga Prov., March 1923; no. 2411, on leguminous leaves, Olongapo,Zambales Prov., March 1924; no. 2412 (part) on Hydnocarpus subfalcataMerr, loc. cit., March 1924; no. 2420, on Ficus, loc. cit., March 1924;no. 2440, on unknown leaves, loc. cit., March 1924; no. 2472 (part) on? Guioa, loc. cit., March 1924; no. 2808, on Symphorema luzonicum (Blco.),Solana, Cagayan Prov., January 1924; no. 2827 (part), on Zingiber, Ca-g-ayan Prov., January 1924; no. 2862, on Aganosma acuminata (Roxb.),Tuguegarao, Cagayan Prov.,January 1924; no. 3118 (part), on Ficus,Asingan, PangaRinan Prov., November 1923; no. 3167, on llligera luzo-

120T. Petch

uensis (Pres I) JVJerr., loco cd., November 1923; no. 3298, 011 Callicarpa,Castilejos, Zambales Prov., March 1924; no. 3378, on Adenia Coccinea

(RIco.), Angeles, Pampanga Prov., October 1923; no. 4749, on unknownleaves, Oagayan Prov., January 1924; no. 6257, on Premna odorata, Ge-

rona, Tarlac Prov., January 1925; no. 6375, on PterospeJ'num niveuni

Vahl, La Paz, Tarlac Prov., December 1924; no. 6381, on Allaeanthus luzo-nicus (Blco.), loc. cit., December 1924; no. 6496, on Jasininuin Sambac,Calumpit, Bulacan Prov., October 1924; no. 7113 (part), on lchnocarpus

1;0Iubilis, Florida Blanca, Pampanga Prov., October 1925; no. 22 119 (part),on Cyathea, Mt. Isarog, Camarines Prov., colI. M. Ramon, November 1913.

Hypocrella Raciborskii Zimm. No. 1855, on unknown leaves, Manila,li'ebruary 1924; no. 2275 (part), on Premna, Castilejos, Zambales Prov.,March 1924; no. 4697, on Ficus, Isabela Prov., January 1924; no. 5656, on

St7"ngylodon, Mt. Apo, Davao, Mindanao, June 1924; no. 6210, on Ficus,Cebu, Cebu Island, May 1924.

In nos. 1815, 2275, 4697, with Aschersonia placenta B. et Br.Aschersonia placenta B. et. Br. No. 1847 (part), on Premna, Manila,

February 1924; no. 2341 (part), on unknown Rubiaceae, Stotsenberg, Pam-panga Prov., October 1923; no. 2909 (part), on Zizyphus talanai, Nueva

Ecija Prov., December-January 1923-24; no. 2639, on Ficus, Polo, Bu-lacan Prov., September 1923; no. 2646 (part), on Phyllanthus reticulatus,loc. cit., September 1923; no. 2649, on Preinna odorata Blco., loco cit.,~eptember 1923; no. 2657, on Premna, loco cit., September 1923: no. 2972,on Ficus ulmifolia Miq., Del Norte, Manila, September-October 1923;

no. 3364, on Pl'emna odorata, Angeles, Pampanga Prov., October 1923;

no. 3430 (part), on 01'eocnide tl'inervis (Wed

d.), Bambang, Nueva VizcayaProv., January 1924; no. 4906 (part), on Lepidopetaluin Per7"ttetii, Tarlac,Tarlac Prov., December 1924; no. 5392, on Cleinensia inacrantha, Nit. Apo,Davao Prov., Mindanao, June 1924; no. 6028, on Glochidium rub

rum BL.,Santa Maria, Bulacan Prov., November 1924; no. 7102 (part), on Ficus,Florida Blanca, Pampanga Prov., October 1925.

Aschersoiiia badia Pat. No. 2275 (part), on Preinna, Castilejos, Zani-bales Prov., .March 1924; no. 2945, on Preinna iiauseosa, Iligan, IsabelaProv., December 1923; no. 3151 (part.), on Piper, Asingan, PangasinanProv., November 1923; no. 4503, on Ficus, Iba, Zambales Prov., February1924; no. 6156 (part), on Preiniia iiauseosa, Angat, Bulacan Prov., No-vember 1924; no. 6571, on Preinna nauseosa, Paniqui, Tarlac Prov., Ja-nuary 1925; no. 7308 (part), on Pilea inelastoinoides, Baguio, Benguet

Prov., Decemher 1925.Aschersonia samoensis P. Henn. No. 922 (red and yellow forms) on

Canarium villosuin (Bl.) Vìl., Orani, Bataan Prov., May 1923; no. 1847(part), on Premna, .Manila, February 1924; no. 2341 (part), on unknowiiRubiaceae, Stotsenberg, Pampanga Prov., October 1923; no. 2408 (part),on Ardisia, Olongapo, Zambales Prov., March 1924; no. 2646 (part) on

P/i'h

on jon JZin~

bun!on 1

1924

1922Oreo1924Octo1924DeceProv.MarÜbilifecarpi.

(par(vembi

A~

berg,As

IsabelTo

Strebl:S einec

Pe~

berg, J

Lour.,Ae~

BataanProv.,lac. citno. 29C

Januar:l\ anila,(Cham.on Pipion Prer,no. 6151

1924; niJanuaryOctoberBaguio,

T. Petch

ovember 1923; no. 3298, on Callicarpa,~h 1924; no. 3378, on Ad'enia coccinea

., October 1923; no. 4749, on unknown~924; no. 6257, on Preinna odorata, Ge-

5; no. 6375, on Pterospermuii niveuni

bel' 1924; no. 6381, on Allaeanthus luzo-1924; no. 6496, on Jasiiinuii Sambac,

1924; no. 7113 (part), on Ichnocarpus

a Prov., October 1925; no. 22119 (part),Prov., colI. M. Ramon, November 1913.No. 1855, on unknown leaves, Manila,ri Premna, Castilejos, Zambales Prov.,tbela Prov., January 1924; no. 5656, on

ridanao, June 1924; no. 6210, on Ficus,

:schersonia placenta B. et Br.

No. 1847 (part), on Preiina, Manila,

llknown Rubiaceae, Stotsenberg, Pam-)9 (part), on Zizyplzus talanai, Nueva

l23-24; no. 2639, on Ficus, Polo, Bu-

346 (part), on Phyllanthus reticulatus,i, on Premna odorata BIco., loc. cU.,ia, loco cit., September 1923; no. 2972,3, Manila, September-October 1923;

eles, Pampanga Prov., October 1923;iis (Wedd.), Bambang, Nueva Vizcayaon Lepidopetaluin Perrottetii, Tarlac,

92, on Cleinensia inacrantlza, Mt. Apo,no. 6028, on Glochidium rubruni BL.,bel' 1924; no. 7102 (part), on Ficus,tober 1925.

(part), on Premna, Castilejos, Zam-on Premna nauseosa, Iligan, Isabelatrt.), on Piper, Asingan, Pangasinanr?icus, Iba, Zambales Prov., Februaryauseosa, Angat, Bulacan Prov., No-

'luseosa, Paniqui, Tarlac Prov., Ja-~ea melastomoides, Baguio, Benguet

No. 922 (red and yellow forms) on

Bataan Prov., May 1923; no. 1847

1924; no. 2341 (part), on unknowiirov., October 1923; no. 2408 (part),T., March 1924; no. 2646 (part) on

o',\',

Some Philppine Entomogenous Fungi.121

pliyllantiZus reticulatus, Polo, Bulacan Prov., September 1923; no. 2740,on Homonoia riparia Lour., Bosoboso, Rizal Prov., February 1924; no.

2803,on Ficus, Tuguegarao, Cagayan Prov., January 1924; no. 2827 (part), onZingiber, Cagayan Prov., .January 1924; no. 3060 (part), on Antidesrabunius (L.), Del Norte, Manila, January-February 1924; no. 3088 (part),on Psycliotria luzoniensis (Cham. & SChlecht.), loco cit., January-February1924; no. 3118 (part), on Ficus, Asingan, Pangasinan Prov., November1923; no. 3169, on Litsea, loc. cit., November 1923; no. 3430 (part), onOreocnide trinervis (Wedd.), Bambang, Nueva Vizcaya Prov., January1924; no. 4653 (part), on Streblus asper Lour., Porac, Pampanga Prov.,October 1923; no. 4846, on unknown leaves, Nueva Ecija Prov., January1924; no. 4906 (part), on Lepidopetalum Perrottetii, Tarlac, Tar~ac Prov.,December 1924; no. 5944, on Celastrus paniculata, Rosales, PangasinanProv., February 1925; no. 6019 (part), on Premna nauseosa Blco., SantaMaria, Bulacan Prov., Noyember 1924; no. 6566 (part), on Flemingia stro-bilifei-, Paniqui, Tarlac Prov., January 1925; no. 7113 (part), on Ichno-carpus volubilis, Florida Blanca, Pampanga Prov., October 1925; no. 22 119(part), on Cyathea,Mt. Isiuog, Camarines Prov., colI. M. Ramon, No-vember 1913.

Aschersonia Oava Petch. No. 2584, on unknown Urticaceae, Stotsen-berg, Pampanga Prov., October 192R.

Aschersonia Coffeae P. Henn. No. 2959, on Psidium Guajava 1., Ilgan,

Isabela Prov., December 1923.Torrubiella luteorostrata Zimm. Sterile stromata only. No. 609, on

Streblus asper Lour., Tarlac Prov., March 1923; no. 2583 (part), onSemecarpus, Stotsenberg, Pampanga Prov., October 1923.

Peziotrichum Lachnella Sacco No. 2583 (part), on Semecarpus, Stotsen-berg, Pampanga Prov., October 1923; no. 4653 (part), on Streblus asperLour., Porac, Pampanga Prov., March. 1923.

Aegerita Webberi Fawcett. No. 970, on unknown Rubiaceae, Orani,Bataan Prov., May 1923; no. 2408 (part), on Ard'isia, Olongapo, ZambalesProv., March 1924; no. 2412 (part), on Hydnocarpus subfalcata Merr.,lac. cit., March 1924; no. 2472 (part), on ?Guioa, loc. cit., March 1924;no. 2909 (part), on Zizyphus talanai, Nueva Ecija Prov., December-January 1923-24; no. 3060 (part), on Antidesma bunius (L.), Del Norte,Manila, January-February 1924; no. 3088 (part), on Psycho

tria luzoniensis(Cham. & Schlecht.), loc. cit., January-February 1924; no. 3151 (part),on Piper, Asingan, Pangasinan Prov., November 1923; no. 6019 (part),on Premna nauseosa Blco., Santa Maria, Bulacan Prov., November 1924;no. 6156 (part), on Premna nauseosa, Angat, Bulacan Prov., November1924; no. 6566 (part), on Flemingia strobilifera, Paniqui, Tarlac Prov.,January 1925; no. 7102 (part), on Ficus, Florida Blanca, Parnpanga Prov.,October 1925; no. 7308 (part), with sporodochia, on Pilea melastomoides,

Baguio, Benguet Prov., December 1925.

The criteria for definition of species in mycology.

By H. C if e rr i.It is by no means the author's intention to discuss again the apparently

insoluble problem of what really constitutes a plant species. From

what experience he has had with phanerogamic plants, he is forcedto admit the existence of those units called "Linnean" species, although

it seems to him that taxonomists only too often have to depend upoii

trained eyesight in order to recognize them. Nevertheless, in some groups,

.,species" in the ordinary sense scarcely exist. If the conception of

species may thus fail when applied to certain phanerogamic genera, it iseasy to imagine what wil happen when it is applied to the lower crypto-g'ams. The further one descends into the realms of the microscopic orga-nisms, the vaguer becomes the definition of species, until at last all endsin chaos. What one actually has to deal with is not a number of well-defined species or genera in the phanerogarric sense, but rather groups of

innumerable organisms, closely allied the one with the other, with almostonly hypothetical links between them, or none at alL. Every cryptogamistadmits the essential truth of this broad statement, as applied to the lowercryptogams.

The necessity of some kind of nomenclature, however, even for themost primitive or rudimentary organisms, obliges one to cling to thecnstomary taxonomic conceptions, be they ever so difficnlt of application.

A p r i 0 ri, as in the case of the phanerogams, the basic unit of taxo-

nomy applied to fungi is always the species, but a kind of "species",

which is not necessarily the same as among the phanerogams. Evidentlymycologists seem to have extended the conception of species to the fungiwithout questioning the feasibility of so doing. For two phanerogamicplants to be recognized as belonging to separate species, certain require-ments must be fulfilled. Primarily, they must differ in a certain, but nottoo small number of characteristics. In addition there should be no inter-mediate forms. Furthermore, hybrid forms, if any exist, should be moreor less sterile. When the levelling effect of intraspecific crossfertilization isexcluded, there appears the phenomenon known as J ordanism, i. e. theLinnean species resolve into numerous smaller, more or less recognizableunits, which are necessarily constant, since reproduction is apogamous.

How much of all this is applicable to fungi? Evidently the fathers ofmycology, being fundamentally phanerogamists, having in most cases onlywidely distinct forms and isolated specimens for study, described as

:ion of species in mycology.. Ciferri.ntention to discuss again the apparentlyy constitutes a plant species. From

th phanerogamic plants, he is forcednits called "Linnean" species, althoughonly too often have to depend upon

ize them. Nevertheless, in some groups,scarcely exist. If the conception of

I to certain phanel'gamic genera, it iswhen it is applied to the lower crypto-itO the realms of the microscopic orga-nition of species, until at last all ends

GO deal with is not a number of well-

3,eroganiic sense, but rather groups of~d the one with the other, with almost:m, or none at all. Every cryptogamistoad statement, as applied to the lower

nomenclature, however, even Ïor the~'anisms, obliges one to. cling to the

ie they ever so difficult of application.) phanerogams, the basic unit of taxo-~he species, but a kind of "species",s among the phanerogams. Evidentlythe conception of species to the fungiof so doing. For two phanerogamic

r to separate species, certain require-

Ghey must differ in a certain, but not

In addition there should be no inter-forms, if any exist, should be more

ect of intraspecific crossfertilization isLenon known as Jordanism, i. e. theus smaller, more or less recognizable

it, since reproduction is apogamol1s.

Ie to fungi? Evidently the fathers ofirogamists, having in most cases onlyspecimens for study, described as

The criteria for definition of species in mycology.123

species everything which they could recognize on "habitus"1). In the case

of parasitic fungi, similar forms were lumped together with small attentionto the host plant, in accordance with the admitted universal pleophagy.The growing amount of material for study tempered the second require-ment for phanerogamic species, and forms not so well marked weredescribed as, first, forms, or varieties, then, as species. When the ten-dency to distinguish the fungi according to their host plant came into

vogue, the application of the first requirement was also greatly limited, atleast for certain groups of fungi. As far as we know, the third require-ment for specific rank in the higher plants (surely, the crucÎal criterion inthe taxonomy of the phan:erogams), is of but doubtful application inmycology. A strict interpretation of this law for fungi is nonsense. Onthe other hand, the true significance, in Nature, of heterothallism and ofpleomorphism is still unknown. B ri e l' 1 e y' s point of view on the in-applicability to fungi and bacteria, of genetic concepts derived from thestudy of the highest organisms, is to a high degree justified. We canadmit in fungi the existence of pure lines or cross-fertiized generations,

alone or combined, with every transitional stage between these extremes.The transference of the conception of species from the phanerogamic

plants to the fungi, leads to the conclusion that true Linnean species or.Tordanian species, in a phanerogamic sense, do not exist in mycology.The last recognizable group in mycology is that of a more or less arti-ficially distinguished, small aggregate of individuals, resembling, in mostcases, Jordanons rather than Linneons, to use the traditional phanero-

gamic terminology.From this fundamental ambiguity in the interpretation and, of course,

i~ the limitation of the specific unit in mycology, comes the greatest indi-vidual variation in the selection and use of criteria for specific rank increating species of fungi, polarized in two opposite extremes, the "pure"morphologic, and the "pure" biologic, with- every intermediate shading.The first and oldest was gradually subjected, according to the criterion ofmost mycologists, to the interference of the second, but the tendency tomultiply species according to the host plant has reached such proportionsthat a general review of the situation may well be demanded. Dr. But Ie l'recently ilustrated (Proc. Tnt. Congr. Plant Sc., VoL II, pp. 1590-1597;1929) the confusion produced by the multiplicity of the criteria employed,which has given rise to the species created during the last twenty years.

The critical point in. the classification in mycology is the taxonomyof parasitic species of fungi, involving the biologic interpretation of thespecies, the "cultural" criteria being derived from and included in theformer. For these, the problem can be reduced to a fundamental question:Must we accept a biologic interpretation in the taxonomy of the para-sitic fungi? If so, can specific rank be maintained for biologic units?

1) "Habitus" is unterstood both from macroscopic and microscopic point of view.

124 R. Ciferri

An exclusive morphologic application applied to obligate parasiticfungi, as defined by the group of neo-morphologists (C u n n i n g ham, e ta. 1.), is, according to Our views, scarcely justified. One cannot return tothe primordia of mycology, and renounce the application of those prin-ciples in classification derived from fifty or more years of investigation ofthe mutual relation between fungus and host plant. In the parasitic fungi,as well as in fungi with highly developed biochemical activities (such asthe yeasts), all phases of their life are subordinate to their biological func-tions, a complex morphologic development frequently being sacrificed toa more perfect adaptation to the host. The consequence is the primitive-ness of many parasitic fungi (caused by a regressive evolution or by a nol1-evolution), whose uniformity of morphology is not correlated with therange and exaltation of biological function. A classification of parasiticfungi on purely morphological grounds takes a one-sided view, perhaps

that of the least important side, compared with the synthetic picture offeredby utilizing all characteristics of the organisms

1).In conclusion, the writer's opinion is that no rules can be concerted

for, nor limitations imposed upon the use of one or another or all criteriafor the definitioIt and delimitation of systematic entities in mycology. Thepresent chaotic situation in this science, as ilustrated by But Ie r, is not

due to the multiplication of systematic units (this multiplication being

the natural outcome of the development of Our knowledge), but to theirinadequate taxonomic expression. In the solution of the latter problem,two courses are open: (1) The adoptIon of specific rank for all systematicunits in mycology, independent of the criterion or criteria employed, butwith an expressed distinction between different kinds of species; or (2) theadoption of specific rank for morphological units only, with one or moreinferior ranks for units established from criteria other than morphologic.

The adoption of one of these two solutions, sanctioned by an inter-national agreement, while'not bearing upon the actual relationship betweenthe species, would, at least, tend to make their taxonomy less beWildering.

1) An exclusive application of morphologic characteristics to the classification

of parasitic fungi, e. g., to the Ustilaginales, as demanded by C u n n i n g ham,beginning with the aggregation of the loose smuts of barley and ,vheat, might endin the partial reconstruction of Us t i lag 0 c a I' b 0 and a few old "Sammel-species", which would include most of the smut species, and affect members ofall familes, from Graminaceae to Compositae. Furthermore, if host specializationshould be valueless for the classification of parasitic fungi, it is very doubtfulwhether any more weight should be attached to the localization of the parasiteof the plant host. As rightly expressed by Dr. But i e r: "If we refuse to acceptthe one in the classification, it is difficult to justify the use of the other". Con-sequently, the classification of certain groups, e. g., Dothideales, as presented byThe i Sse nand S y d 0 w, should not be considered.

Theeach di¡Ascomyiand, forhost pIa

applied.of the inSpecific igical diffon matrÙare chamor animalsymptom:of the boi

of Ascom:than thost

(e. g., Tabiochemic:

Thus i1of what sh

as in thost

gronp. ADr. But i.

But the!!''neity of :tlir dispari

In the jsiiut fungi

particular ¡¡

his reason in

Even thEnirieties, an

the fact tha1

is by no me:

taxonomic u:Sub-groups, :

') The mphological diff,great difficultybe based on b

eta 1. Morphc

and size of Sc

changes in env

126 R. Ciferri

A. Morphological characteristics upon which the originally recog-nized and more easily distinguished species were based. This is the oldestas well as the most important criterion, when dealing with herbarium

materiaL. All morphological characteristics, however, are not of the sameimportance. One can distinguish between:

1. Mac r 0 - m 0 r p h 0 log i c a I c h a r act e ri s tic s, based on well-defined and easily recognized structural differences.

2. M i c r 0 - m 0 r p h 0 log i c a I c h a r act e ri s tic s, based oil rela-tively small differences in shape of any formed element.

3. B i 0 met ri c c h a r act e ri s tic s, based on small, but stil stati-stically recognizable differences in size only.

B. Biological characteristics, including all other than morphological

ones. The biological criteria, originally most neglected, are now widelyused in distinguishing species").

They are of several kinds:1. M a tr i c a I c h a r act e r i s tic s, based upon the study of the

symbiotic phenomenon between fungus and living host plant. Ac-cording to the nature of the different manifestations of their pa't"a-sitism, they may be subdivided into:A. Specialization characteristics, based on the range of the parasitic

adaptation of each fungus. According to the nature of the

methods employed, we can distinguish:a) S p e cia i i z a t ion cap a b i e 0 f d ire c t proof from

the outcome of cross-inoculation with positive result.b) S p e cia i i z at ion cap a b i e 0 fin d ire c t proof by

cross-inoculation with negative result (inability to infect a

definite host plant under experimental conditions), or byfailure to produce natural infection under natural conditions(e. g., in the field).

c) Sup p 0 sed s p e cia i i z a t ion, not directly or iiidirectlyproved, but based on a probable or possible analogy of beha-vior with a specialized fungus of known host range.

These three criteria are arranged in order of their im-

portance.B. Ecological characteristics, based on the study of the localization

of the fungus in or upon definite organs of the plant host.C. Pathographic characteristics, based upon the study of the effect

of the parasitic fungus (mechanical, teratological, metamorphic,

hystolytic, etc.) on the host plant, as well as on the reaction ofthe same to the parasite. This characteristic is, for the most

") A classification based upon the interrelation between two organisms, andtakig into consideration the complex fungus plus the host plant, is a phyto-

pathological classification rather than a mycological one. Of course, this remarkmay affect the classification only from a philosophical point of view.

paph

2. C u It

behavteristiartifi csite oj

with ~

TheseA. Ph;

natB. Bio

catranana

C. Ph~

theresi

Geographisitic fungi asdistribution 0

Most of tltaxonomy of tof the twent

morphologic (proposed on n

a rule, with iused, and, as

to that of the

with the matr

found. No Spi

alone, but seVt

liarities. Veryalone, and non

Taxonomicmuch employeehut varieties h,

teristics (i. e.variety TOU1'E

slig'htly larger

4) These tliri

of instruments u

physical, chemic~

cases the division

128 R. Ciferri

characteristics combined (e. g., U stilago longissiina iSchlecht. J Mey., typeon Glyceria aquatica; the variety dubiosa Liro on Triticum repens with

the additional characteristic of producing perforation of the leaves).Some varieties have been based on ecological characteristics (e. g.,

Ustilago tritici iPers.J Jens. on the spikelets of Triticum vulgare, with thevariety foliicola P. Henn. on the leaves of the same plant host). Thesub-variety as a taxonomic unit has not been employed in the Ustilaginales,nor the sub-form. Only a few forms have been proposed, more or lesswith the same criteria as those of the varieties, generally based on ecolo-gical and matrical characteristics (e. g., Entyloma linariae Schr. on Linaria1Jllgaris, with the form veronicae Wint. on Veroiiica, with amphigenoussori as an additional characteristic).

Only one author has used race as a taxonomic entity, based on micro-morphologic and matrical characteristics combined (e. g'., U1'ocystis ane-mones iPers.J Wint., on Anemone neinorosa, with the slightly morpholo-gically distinct race ranunculi-1'epentis Bub., on Ranunculus repens).

Nom i n a n u d a were frequently created by the earlier authors as\-arieties or forms, based, as a rule, only on matrical characteristics or onsome supposed specialization.

In conclusion, new entities have been described promiscuously as

species more often than units of inferior rank, with the result that thereis a considerable amount of confusion as to the real meaning of "species"as applied to the Ustilaginales.

In order to ilustrate recent splitting of Some classical species, the

writer has arranged the elements diagrammatically, according to the

principal characteristics upon which they have been based. Figure 1shows the species derived from Ustilago violacea iPers.J Rouss., asrecently described, chiefly on the basis of experimental cross-inoculation

as presented by L i l' o. Of eleven species listed, including U. violacea s.stricto reduced to the smut attacking the type host plant, nine were basedupon the results of cross-inoculations, and two 'are micro-morphologic andmatrical speci8s (Fig. 1). Of the six species derived from U. hypodytesiSchlecht.J Fr. (Fig. 2), three were based on morphologically differentiatedcharacteristics and on a supposed specialization on some unstated hostplant; one on pathographic characteristics combined with the variationin the host plant; one on ecological characteristics and host plant; one ona supposed host plant specialization. Of the many species derived fromUstilago striaeformis iWest.J Niess!. or the cycle thereof, today dividedalmost entirely according to the host plant parasitized, Figure 3 ilu-

strates the actual situation of ten species. Three species were based onmorphological differences, as well as on a supposed host plant speciali-zation; four species were isolated upon the basis of ascertained hostspecialization; all other species upon a supposed specialization. 801'0-sporiimi saponariae Rud. was recently sub-divided into four species, in-

Ii/I"'i/,I'

11 ,,''i' ,,IIii'i1'

8 II! i~ :11 ivi ,/I I

'i I~ "ii;j !, io .,1 ,~ Ji' /-- fl /'J T/ /

~ itl/- ~,ct ~

~ ¡'~ ~\\o ~\ \.- _II \;; 'ii \ \o iii \bJ iii ,ct " \:- Iil i~ 1\ \;: ,I, 1

1"1\'

'1\'iIi, \I, 'II i

I'II

\1I

IIIIi

I

I

'Jngissima iSchlecht.J Mey., typez Liro on Triticum repens with

. perforation of the leaves).Jcological characteristics (e. g.,its of Triticum vulgare, with theof the same plant host). The

m employed in the Ustilaginales,"e been proposed, more 01' less;eties, generally based on ecolo-ityloma linariae Schr. on Linariam Veronica, with amphigenous

:onomic entity, based on micro-~ombined (e. g., Urocystis ane-~a, with the slightly morpholo-

l., on Ranunculus repens).tted by the earlier authors as

1 matrical characteristics or on

ri described promiscuously asank, with the result that there, the real meaning of "species"

of some classical species, themmatically, according to the, have been based. Figure 1i violacea iPers. J Rouss., as

experimental cross-inoculationisted, including U. violacea s.pe host plant, nine wer.e based

wo are micro-morphologic andes derived from U. hypodytes

morphologically differentiatedzation on, some unstated hostcombined with the variation

iristics and host plant; one onie many species derived fromcycle thereof, today divided

it parasitized, Figure 3 ilu-

Three species were based onsupposed host plant speciali-Iie basis of ascertained host

.pposed specialization. Soro-

divided into four species, in-

The criteria for definition of species in mycology.

I U. antherarum (Fries) Liro. . " ",

Ii,

: / U. coronariae Liro " ." ..".".'/Ii""'i"JI I U. diamhorum Liro . . . . .'II,II'I

;,," i

'': i U. lychnidis - dioicae (D. C) Liroo ""~ :'''cI 'ii I

'i I~ "ii:: ~/' . I" L'o Ii' I U. niva is iro............~ ~/' io t' iVJ Ii:u 'I

~ it- -- U. pallida Liro . . . . .

td;\.. ~i\o ~\ \.- \1 I \

;; 11 I ,

o 1)\\ 'u silenes - nutantis (D. C) Liro . . .bf "I \C' " i:- ',',t: 1\ \;: ,I, I

I" i U."silenes-inflarae (D.C) Liro '."1,1

II''1\'i II \ .1\ \ U. stellariae (Sow) liro . . . . . , .

\1II

III

III,i U. superba Liro .........II

I

iI U. violacea (Pefs.) Rouss. s" stricto ..

TBmiTn:)

)!qduSoqiBd

)!.á°10)3:

uO!iBZ!TB!);id s

p;isoddns

uO!iii!TB!);ids

p;iU!Bii;i)SV

J!.á°ToqdJoW

129

::::o~~'"..'"c.'"'"'"u'"

o~

oQI..

'";:2

-0'"

-;õ

Eo

J:-0'"~..'"

-0'"'"

'ü'"a.vi

,.0.~

9

t, )~i, _i

-r ~:j

.: (

'Ii,

130 R. Ciferri The critE

pUTIin:)

I U. aculeata (Ul,

i U. agrestis Syd.,.,I ,

I I, I, . I U. agrostidis-p,, i Ii

J!qdUJÉ'oqiud .. i Ii ¡i I,, , ,I .. , II ~ 'I, u l' i U. airae-caespit, .. 1:1:a 0, u. amplexa Syd. ..

, u ~ 'i', I ~ :' i0 , , vi ,I i..

, I '" .'1~ 1 i.. Vi ,', U. alopecurivor:

I I ;; V" I I I Ivi V ,'. I

, I -0Zvi I I 0 ,ii ,

,,, I.. II 0. ,', I~ , I '" 0 ,,, ,, i

J!É'OIOJ3:.. .. ,'i i.. V"

" 0 Q) "" , U. brizae (Ule).. /I U. athenae Maire òO ~ "iI ~.. , . . tq I "U II i '".. './ i~ II I

,/ i .. rJ ).1'.. '1 i '"

'êt-~~~~ !l/ ~U. bromina Syd~( 8 i. II

.. ~I\ I

U l,' -0 1\ i

t.li '- ~ iU '\\'ii , '" .~ \\\ \'" ii 'u d rd' i. ,,\U. phlei - praten~Maire .. .. II i 'o i i . . acry i 15 0 f/ I' I

0. I,S 0 " .~ II " ",.. \ ' UOPUZ!IUpddS0 0. ,\ '

I' oJ ~ II III ,0 \ ' pdsoddns -0 '0 1\ U. milii (Fuck.), ..b. i'

"tf/ I'

C' I , ;: , ,, .. I... -0 . I'0 ,

I , '" I iVl , I U. Iygei '(Maire) .. \ ,~ i Maire .ü , U. Salvei B. et,

.. I0. iI CJi

,

uo!iuZ!rE!Jdds,

i i, ,i

PdU!UlldJSVN U. scaura Liro .

\ oj1¡.\

,i U. valesiaca (SchelL.) Maire .

~

J!É'°roqdJow

ri The criteria for definition of species in mycology. 131

pllnlin:)

U. aculeata (Ule) Liro.piJniin::

i '.

I.,II,I U. agrostidis - paluscris Dav. . oni I on

OJi')!adiiJâ'oqiiid Z)!adiii~oalTld ~ i II i

" ' , ":, , '".! . ,

~v l' , U. airae-caespitosae (Lind.) LiroOJ 0 1/1 'Y:a .. '"v ~ " Ien

:' i 8..vi iI I ~

on I: ' 0OJ -; U. alopecurivora (Ule) Liro . '-'; '" ::,' i OJ

.~ os-0 , I I ~0 Z

,I, ,J!â'°IOJ3:0. ''i J '"

,', I~ ~ ,,, I 0

)!~OJO)g:.. .. 'I' J b/'"0 Q) "" l U. brizae (Ule) Liro . .- os~ '',1 ,b/

:.' "~

os ~ './, '"

~t;:u

'"

'S ,-~~~8;:

U. bromina Syd. -08

i- fl' ~i. i'" ..-0 1\ '

~V 'ii' uo!iiiz!pipdds ..

. ~ \\ \ \ 0.. ~ ,,'

u. phlei - pratensis Dav. pdsoddns a.. .. II I \ 00 ~ 11, ~6 0 ,I.

\' " -00. ''. ' OJ

UO!lTlz!JTlpdds0 .~~ C1 I, '- ii \ OJ

pdsoddns -0 'C II U. milii (Fuck.) Liro -0OJ ~ I' on~.:; ~ ' \ OJ

'0" , . OJ-0 , \ 0.on

J

' , enOJ \ \

U. Salvei B. et Br.'ü \uO!iiiz!ITlpddsOJ ,0. , '"(/

, pdU!iiiJd)SV 0.uO!liiZ!ITl!)dds

,\ ¡E,

PdU!Tlll;))SVN U, scaura Liro .oiii

J!â'oioqdJoW

)!~OloadlOW

9*

132

, S. alsinearum Cif. . . . .,,,/,/,,,,I/ I S. dianrhi Rabenh., i, /, ,

i Ii I/ ,/ """ / I;: 'iii , i

v Î/ , S. dianrhorum Cif.~ /1 ,. -- I' ./~ ~i/,/C ~'o t,~Il'"~ II,rJ ~\ ',.

\\ "\ \ ,\ .. S. gypsophilae Cif. . . . .. \. ,I I\ \i \I \, iI \I \i \\ \\ i S. silenis - inflatae (2.) Cif.ii\Ii,,II\\,

: S. saponariae S. striCto

o~~in

8~.-l.o0.Vlol.o

C/

R. Ciferri

pmUjnJ

J!i:duá'oi:iud

J!É'OjOJ3:

uori¡¡z!jupdds

pdsoddns

uopuzt¡U!Jdds

PdU!UiJdJSV

J!É'Ojoi:diow

-0::i:lU'"

..'"coa.'"'"

E::..oa.'"o..ovi

8-0~"õ

o

Eo

c.-0lU

'E;lU

-0colU

'ülUa.

vi

~öO

¡i

The (

i E. ancej

,

I' E. arnic

I',1,

;, I E. arno~"

i,'

,,' I E. as(eri",i,',"

I,,: i E. bavar"i,,'

'::;; i E. bolro','" ¡

o 1',"~ ).~/,',' J E. catan;

,1,/1, i

r. '','' JI,il,' " E. Davis

;; ""i"i: ~I,,' i

~ ,",,,~ ii', i' ' E. erigero l~i' I"" ~¡// l;: 1~11

o .t,i / E. eupa(i"t;',,/~r--- E. heleni--f'~ t"",

"1. i,\ '- " E. hieracC ''1,"~. 111\\~ 'i 'u II I i E. incern

\ II~ i'll8 1111'.o i i . E. leontc__ 1\:: Ii \.i 1\ \C E. lepachrT' \ i,. 'i

Ii E. Maire¡

\\

\ E. madia,

E. parthe

E. Saccan

E. Scaljan

E. \Visean

The criteria for definition of species in mycology. 133

i E. anceps Cif.pmu¡nJ i

I

II E. arnicae Syd. . puniinJi'"i

E. arnoseridis Syd. .. II'"

'"

"'i E. asteris - alpini Syd.

)!4duJ~04lUd -c "ihi, ".

:: ¡:i: "Syd.

/,': iE. bavaricum )!4duJ~04lUd v

ll , c:'" ,"i,.. ""

E. boltoniae Cif. ..-

" ','1,1 -0c ::

\0 '.'"i 0c. 1'''1

V'" 0

1" "OJ'" ..

E. catananches Cif. "~ "I' J

E 1,/1' I :::: rn 'Ii" 1 -0

)!~oIO)3: .. 1'11; i iE. Davisii Cif. .

c;; '"0¡: f'i' I J

c.(¡I,',' i

"'" )!~oIO)3:

v0 Q) "'1, /.. Cl E. erigerontis Cif. "0 ii'" i i

ECf 0 l~,/ / 08 'i Wi I

è';: I,11E. eupatorii Cíf.

-0 o 'L11 // c~

f /"'V' 1/

-; 1/ 8il _ Cif.Õ

t' ' ---. E. helenii-J¡' -0

E :: til", ~UO!iuz!lul);)ds 0 '". ~,i\ '.

E. hieracii Syd.-;

c. C'ii," Õp;)soddns -0 ~ Ill"'.'" t' 'i i UO!iuz!¡ul);)ds

E::.¡: U II I i E. incertum Cif. 0'" I II c.

-0 t' ,Ii' p;)soddns -0" 8 \ i '"'" III i E. leootodontis Syd. ,t'ü 0 \ I ''" , I '"c. - -0

Cf ~ IIi.¡ I I \'"

c: E. lepachydis Cif. '"

ii Û~ '".. 'I 0.

CfUO!iuz!lul);)ds bJ \1 E. Maireanum Cif. .

¡; \UO!iuz!lul);)dsP;)U!UlJ;nsy ,

\E. madiae Cif. . oj, P;)U!Uii;))sy ¡;

E. parthenii Cif.

E. Saccardianum

J!~o¡oqdJoW E. Scalianum Cif.

E. wisconsiniense Cif. )!i?oloqdJoW

134 R. Ciferri

eluding S. saponariae s. stricto and the old but very little differentiatedS. dianthi Rabenh.; six in all. (Fig. 4). Four species were isolated uponthe basis of micro-morphologic characteristics, and on the basis of hostplants attacked; one on ecological characteristics and a supposed hostspecialization.

Under the caption, Entylonia calendulae (Oud.) De Bary, Europeanmycologists included all Entyloma living on the members of the Com-positae family. On the other hand, American mycologists assigned the

same function to E. coinpositaruni Farl. The two species, from the morpho-

logical standpoint, are identicaL. S y d 0 w sub-divided the Europeanspecies according to the. genus of the host plant parasitized, and thewriter the American species, chiefly upon a similar basis. The result wasthat both species are at present represented by nearly twenty-five species,

of which twenty-one are sketched in Figure 5. Besides the supposed host

specialization, common to all derived species, eight were based on smallmorphological differences; none on ascertained host specialization, nor onecological, pathographic or cultural characteristics.

The writer trusts that the ilustrations given above wil show in howmany different senses the term "species" has been used in mycology. Nodoubt, mycologists would welcome any effort toward a standardization ofthis term, although it must be admitted that a wholly consistent nomen-

clature, closely reflecting the true value of any taxonomic unit" is anunattainable goal.

Some relief from the present confusion may be obtained by the simpleexpedient of always indicating, by means of some abbreviation, w hatkin d 0 f s p e c i e s one has in mind. For instance, m. would indicate amorphologic species; ec. an ecologic species; pa. a pathographic species;cu. a cultural species. A double or multiple abbreviation would indicatet,hat a species has been proposed on two or more characteristics; e. g.,m. ma. would indicate that the species was proposed on morphologic

(micro- or macro-morphologic or biometric) and matrical grounds, directlyor indirectly ascertained, or predicated on some supposed specialization.

Applying this method to the species embraced in Figure 2, we should

have:Ustilago valesiaca (Schell.) Maire, ma. pa.U. dactylidis Maire, ma. m.U. lygei Maire, ma. ec.U. athenae Maire, ma. m.

U. aniplexa Syd., mai m.U. agrestis Syd., mai

lt is clear that this suggestion has the advantage of not changing thestatus of species already described, nor the rank, leaving on one side theproblem of elevating all varieties and forms to the dignity of species. Thelevelling of all entities to the same specific rank, independent of the cri-

terion or ification. ,,:hand, thisspecies", idytes, s

The searrange thspecies", (some scalenomic valtgiven the Jsecond pIaicharacteriscies", andperts, char

A tentaTaxonomicTaxonomic

metiTaxonomic

specTaxonomicTaxonomicTaxonomicTaxonomic

ApplyiiiFigure 4, tl

SoraSoroSoraSoroSo 1'So 1"

The inc(in rank illwould not Jzoological iinferior uni

A thirdnomenclatubecame gen

yan Halltake rank ii

old but very little differentiatedFour species were isolated uponristics, and on the basis of hostacteristics and a supposed host

ulae (Oud.) De Bary, European

5 on the members of the Com-erican mycologists assigned the'he two species, from the morpho-o w sub-divided the European

Qost plant parasitized, and thea similar basis. The result was

~d by nearly twenty-five species,re 5. Besides the supposed host

cies, eight were based on small'tined host specialization, nor oncteristics.given above wil show in how

tias been used in mycology. Noort toward a standardization of

hat a wholly consistent nomen-

of any taxonomic unit" is an

may be obtained by the simple; of some abbreviation, w hat, instance, m. would indicate aes; pai a pathographic species;

Ie abbreviation would indicateor more characteristics; e. g.,was proposed on morphologic

and matrical grounds, directlysome supposed specialization.braced in Figure 2, we should

Maire, ma. pa.

,dvantage of not changing' the

rank, leaving on one side theto the dignity of species. Therank, independent of the cri-

The criteria for definition of species in mycology. 135

terion or criteria upon which the species is based, is but a formal uni-fication. "Species" wil always be as heterogenous as before. On the otherhand, this does not consider the "mother species" from which the "minorspecies", if any, was taken, or to which it is alled (U s t i lag ok y po _d Y t e s, s ens u 1 a to, in the example given in Figure 2).

The second (and considerably more revolutionary) proposal, is toarrange these different kinds of "species" as in.ferior units of the "motherspecies", or the 'old morphological "S am m e Is p e c i e s", according tosome scale supposed to express con v e n t ion all y their relative taxo-nomic value. Evidently, morphological characteristics would have to beg'iven the main consideration, while matrical characteristics would take asecond place, and so on. Only units based on broadly different morphologiccharacteristics (macro-morphologic species), would be maintained as "spe-cies", and these should be selected in each group by a committee of ex-perts, charged also with ranging the units inferior to the species.

A tentative scheme of arrangement could be as follows:Taxonomic entities based on macro-morphologic characteristics: SPECIESTaxonomic entities based on micro-morphologic characteristics or bio-

metric characteristics: ............... SUB-SPECIESTaxonomic entities based on matrical characteristics and ascertained

specialization:. . . . . . . . . . . . . ... . . . . . .' VARIETY

Taxonomic entities based on ecological characteristics: . SUB- VARIETYTaxonomic entities based on pathographic characteristics: . . .. FORMTaxonomic entities based on cultural characteristics: . . . . SUB-FORMTaxonomic entities based on some supposed host plant specialization: RACE

Applying these criteria to the binomial indicated in the diagram

Figure 4, the new arrangement would be:Sorosporium saponariae Rud. (species).S01"sporium saponariae Rud. subsp. silenis-inflatae (Zigno).Sorosporium saponariae Rud. subsp. gypsophilae (Cif.).S01"sporium saponariae Rud. subsp. dianthorum (Cif.).S01"sporium saponariae Rud. subsp. diantki (Rabenh.).Sorosporium saponariae Rud. var. alsinearum (Cif.).

The inconvenience of this method would be the very numerous changesin rank involved. However, "new" combinations for these inferior unitswould not necessarily have to be used, and the application of the rule ofzoological nomenclature for new combinations, eventually limited to theseinferior units, could be accepted in a botanical nomenclature.

A third and last suggestion is the adoption of the so-called trinarynomenclature, adopted by zoologists and some American botanists, whichbecame generally applicable in botanical taxonomy, under the auspices ofvan H a i 1. In this case all derived units inferior to the species wouldtake rank in order of their fundamental macro-morphologic species, selected

:I

136 R. Ciferri, The criteria for definition of species in mycology.

as proposed in the preceding paragraph, without considering the criterionemployed in the definition of these units.

Applying the trinary nomenclature to the entities indicated in Figure Lwe should have:

Ustilago violacea (Pers.) Rouss.

U. violacea antherarum (Fries).U. violacea coronariae (Liro).U. violacea dianthoritm (Liro).U. violacea lychnidis-.dioicae (DC.).U. violacea nivalis (Liro).U. violacea pallida (Liro).U. violacea silenes-in(latae (DC.).U. violacea silenes-nutantis (DC.).U. violacea stellariae (Sow.).

U. violacea superba (Liro).In the writer's opinion, this third suggestion is the most simple and

would be productive of the least radical change. If accepted, the homo-

geneity of the "species" would be maintained, the first line containing themorphological (macro-morphologic) characteristics, but without the sacri-fice of all other taxonomic entities established on different criteria, andwith a minimum of change of the present nomenclature.

A b bot t, E. V.

XXI, 1931,Abbott, E. V.

XXIV, 1932

Agostini, AnIstit. Bot. L

Agostini, Aibarinus COl

Agostini, Aionicomicosi

A p pel, O. Die

(Rep. Proc-401.)

Arnaud, G. Jsites: CalI(

XVI, 1930,Arnaud,G. LE

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