ANNALES MYCOLOGICI EDITI 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 In halt (Band XXX).

i f i. Originalarbeiten. B 0 e d i j n, K. B. Einige Bemerkungen zu del' Abhandlung von S. Seite C. Teng, "Fungi of Nanking I" ...... 4í8 C i fer l' i, R. The criteria for definition of species in mycology . . 122 ~ G 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 identity l! . i 1 of Tulasnella anceps Bres. et Syd...... '.' . . 463 " r . I 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' Gattung Panus ...... ~ ...... 10 P a 1m, B. T. On Cyttaria Berk. and Cyttariella n. gen. . 405 P a 1m, B. T. Biological notes on Albugo...... 421 Pet c h, T. Some Philippine Entomogenous Fungi. . 118 Pet c h ,T. Gibellula...... '.' . . . 386 P 6 t l' a k, F. und C i fer l' i, R. Fungi dominicani. II 149 Pi 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 81 By d 0 w, H. Novae fungorum species - XXI ...... 91 By d 0 w. Mycotheca germanica. Fasc. L-LII (no. 2451-2600) 394 Z a h i b r u c k 11 e 1', A. Neue Flechten. -- XI ...... 427 Burg U. 1\1. In the dimem pressed orifice resembles the As ture of the pycn Some Philppine Entomogenous Fungi. disc is lacking l' By T. Pet c h. nidia are not reg Through the kindness of Dr. Sydow, I have had the opportunity of Hypocrella ph examining numerous specimens of entomogenous fungi on leaves, collected flavis, planis, te by Mary Strong Clemens in the Philippines. Though the collection con- usque 3 mm dia sisted of seventy-two numbers, it contained only nine species, and with cylindraceis vel c one exception (Aschel'sonia Coffeae P. Henn.), these are all parasitic on tomentosis, sing); Aleyrodidae. Except where otherwise stated, the localities cited are in ampullaceis, 450 Luzon. The list may serve to give some idea of the relative abundance of ascis 240¡. longi the diffei;ent species of Aschersonia parasitic on Aleyrodidae in that country. sporarum angiisti The chief interest of the collection lies in the occurrence of many spe- Aleyrodid on und cimens of a new species, which is described below as Aschersonia phi- 1923, no. 3144; è lippinensis, and its perithecial stage, Hypocrella philippinensis. This has January 1924, ne been known to OCCur in the Philippines for many years, but the specimens Petch in the sam collected previously have never been in a fit condition for description. Cebu Island, May The species is remarkable in that it often forms flat, circular, tomentose Or Aschersonia p radially fibrilose, white or cream-coloured stromata, up to 7 mm diameter, fJavis, planis, ten with a fimbriate margin, without any pycnidia or perithecia. With the help fimbriato, centro 1 of the present specimens, it is now possible to refer the following previous 0,25 mm alt., rect: gatherings to Aschel'sonia philippinensis, - On Schizostachyuni, N Olzag'aray, tibus; vel strom: Provo Bulacan, May 1910, Rb. Bureau of Science, no. 10838; on Astronia margine disci; VE sp., Mt. Maquiling, Luzon, September 1910, Merrill no. 7150; on Astronia, marginatis. Pycr l\it. l\iaquiling, February 1914, C. F. Baker, Fungi Malayana, no. 8; on rotundatis vel coi Derris heptaphylla, Lamao, H. A. Lee, no. 59. Some of these specimens fusoideis, fine act were referred to under Hypocrella Molli in the G e n era H y p 0 c r e II a The pycnidial st and Aschersonia, Annals of the Royal Botanic Gardens, Rourea erecta (El Per ad e n i ya, VII, 240. Immature specimens resemble immature Hypo- Also no. 217, ( crella Mollii, but are more tomentose and usually have a fimbriate margin. D.ry 1923; no. 72( In general, the stromata of Aschersonia philippinensis are at first plane Memecylon, loco ; and circular. Subsequently they develop, in the centre, a gToup of co- no. 806, on unkno lumnar tubercles, each of which bears a single, widely open, circular pyc- known leaves, loc. nidium at the apex. When, however, the tubercle is oblique, the pycnidium Pampanga Prov., opens obliquely and the orifice is usually vertically compressed into a Za.mbales Prov., 1 narrow slit. In some stromata, the centre is occupied by a raised disc up Merr., loc. cit., ~ to 2 mm in diameter, instead of a group of tubercles, and the pycnidia no. 2440, on unk open on the vertical face of the disc, their orifices being overhung by its ? Guioa, loco cit., j upper edge. Other stromata may have neither tubercles nor disc, the pyc- Solana, Cagayan nidia occurring in the stroma, vertical or oblique, the orifice being gayan Prov., J an surrounded by a slightly raised rim. Tiiguegarao, Cag: Asingan, Pangasi 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 or :ne EntomogenOusFungi. disc is lacking resemble forms of Aschersonia placenta in which the pyc- :y T. Pet c h. nidia are not regularly arranged. '1'. Sydow, I have had the opportunity of Hypocrella philppinensis Petch, n. sp. - Stroniatibus albis vel palldo- of entomogenous fungi on leaves, collected fIavis, planis, tenuibus, interdum margine fimbriato vel membranaceo, ~ Philippines. Though the collection con- usque 3 mm diam., tubercula spars a vel conferta ferentibus; tuberculis it contained only nine species, and with cylindraceis vel ovoideis vel subglobosis, usque 0,5 mil alt., 0,35 mm diam., 'feae P. Henn.), these are all parasitic on tomentosis, singulo perithecio in quoque tuberculo; peritheciis angusto- ierwise stated, the localities cited are in ampullaceis, 450 .¡, alt., 150 ¡, diam., ostiolis minutis, impressis, flavis; ve some idea of the relative abundance of ascis 240 '¡. long'is, 8 ¡. diam., cylindraceis, capitatis, octosporis; articulis ia parasitic on Aleyrodidae in that country. sporarum angnsto-ovalibus, rectis v81 curvatis, 8-12 ¥ 2-2,5 ¡,. On an iction lies in the OCcurrence of many sp~- Aleyrodid on undetermined leaves, Asingan, Pangasinan Prov., November is described below as Aschersonia phi- 1923, no. 3144; ditto, on Apocynaceae, Bambàng, Nueva Vizcaya Prov., ;age, Hypocrella philippinensis. This has January 1924, no. 3458; in both cases with Aschersonia philippinensis Jpines for m.'ny years, but the specimens Petch in the same or separate stromata. Also no. 6214, on , Cebu, been in a fit condition for descriptio~. Cebu Island, May 1924. it often forms flat, circular, tomentose or Aschersonia pliilppinensis Petch, n. sp. Stromatibus albis vel palldo- coloured stromata, up to 7 mm diameter, flavis, planis, tenuibus, tomentosis, UJsque 8 mm diam., margine saepius ny pycnidia or perithecÏa. With the help fimbriato, centro tubercula sparsa vel conferta, cylindracea, 0,36 mm diam., T possible to refer the fOllowing previous 0,25 mm alt., recta vel obliqua, singulo pycnidio in quoque tuberculo, fereii- ensis,-On Schizostachyum, Norzagaray, tibus; vel stromatibus centro' discoideis, ostiolis pycnidiorum obliquis 'eau of Science, no. 10838; on Astronia margine disci; vel stromatibus omnino planis, pycnidiis sparsis immersis bel' 1910, iVlerril no. 7150; on Astronia, marginatis. Pycnidiis concavis, 0,25 mm diam., 0,15 mm aItis, ostiolis F. Baker, Fungi Malayana, no. 8; on rotundatis vel compressis, sporis coacervatis flavis. Pycnosporis angusto- Lee, no. 59. Some of these specimens fusoideis, fine acutis, 9-11 ¥ 1,5-2¡.. Paraphysibus usque 150 ¡, longis. M ollii in the G e n era H y p 0 c r e ii a The pycnidial stage of Hypocrella philippinensis. On Aleyrodidae. On if the Royal Botanic Gardens, Rourea erecta (Blco.) Merril, Isabela Prov., December 1923, no. 2921. re specimens resemble immature Hypo- IAlso no. 217, on unknown leaves, Mt. Arayat, Pampanga Prov., Febru- ie and usually have a fimbriate margin. :Jry 1923; no. 720, on Lunasia amara, loco cit., April 1923; no. 744, on ersonia philippinensis are at first plane Memecylon, loc. cit., April 1923; no. 785, Eugenia, loco cit., April 1923; evelop, in the centre, a group of co- no. 806, on unknown leaves, Orion, Bataan Prov., May 1923; no. il'S a single, widely open, circular pyc- known leaves, loc. cit., March 1923; no. 2199" on unknown leaves, Stotsenberg,834, on un- the tubercle is oblique, the pycnidium Pampanga Prov., March 1923; no. 2411, on leguminous leaves, Olongapo, usually vertically compressed into a Zambales Prov., March 1924; no. 2412 (part) on Hydnocarpus subfalcata ~entre is occupied by a raised disc up Merr, loc. cit., March 1924; no. 2420, on Ficus, loc. cit., March 1924; group of tubercles, and the pycnidia no. 2440, on unknown leaves, loc. cit., March 1924; no. 2472 (part) on i, their orifices being overhung by its ? Guioa, loc. cit., March 1924; no. 2808, on Symphorema luzonicum (Blco.), Te neither tubercles nor disc, the pyc- Solana, Cagayan Prov., January 1924; no. 2827 (part), on Zingiber, Ca- rtical or oblique, the orifice being 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- 120 T. 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 unknown P/i'h leaves, Oagayan Prov., January 1924; no. 6257, on Premna odorata, Ge- on j rona, Tarlac Prov., January 1925; no. 6375, on PterospeJ'num niveuni on J Vahl, La Paz, Tarlac Prov., December 1924; no. 6381, on Allaeanthus luzo- Zin~ nicus (Blco.), loc. cit., December 1924; no. 6496, on Jasininuin Sambac, bun! Calumpit, Bulacan Prov., October 1924; no. 7113 (part), on lchnocarpus on 1 1;0Iubilis, Florida Blanca, Pampanga Prov., October 1925; no. 22 119 (part), 1924 on Cyathea, Mt. Isarog, Camarines Prov., colI. M. Ramon, November 1913. 1922 Hypocrella Raciborskii Zimm. No. 1855, on unknown leaves, Manila, Oreo li'ebruary 1924; no. 2275 (part), on Premna, Castilejos, Zambales Prov., 1924 Octo 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, 1924 Dece Cebu, Cebu Island, May 1924. Prov. In nos. 1815, 2275, 4697, with Aschersonia placenta B. et Br. MarÜ Aschersonia placenta B. et. Br. No. 1847 (part), on Premna, Manila, February 1924; no. 2341 (part), on unknown Rubiaceae, Stotsenberg, Pam- bilife carpi. panga Prov., October 1923; no. 2909 (part), on Zizyphus talanai, Nueva Ecija Prov., December-January 1923-24; no. 2639, on Ficus, Polo, Bu- (par( vembi lacan Prov., September 1923; no. 2646 (part), on Phyllanthus reticulatus, loc. cit., September 1923; no. 2649, on Preinna odorata Blco., loco cit., A~ berg, ~eptember 1923; no. 2657, on Premna, loco cit., September 1923: no. 2972, As on Ficus ulmifolia Miq., Del Norte, Manila, September-October 1923; Isabel no. 3364, on Pl'emna odorata, Angeles, Pampanga Prov., October 1923; To no. 3430 (part), on 01'eocnide tl'inervis (Wed Strebl: Prov., January 1924; no. 4906 (part), on Lepidopetaluind.), Bambang, Per7"ttetii, Nueva Vizcaya Tarlac, S einec Tarlac Prov., December 1924; no. 5392, on Cleinensia inacrantha, Nit. Apo, Pe~ Davao Prov., Mindanao, June 1924; no. 6028, on Glochidium rub Santa Maria, Bulacan Prov., November 1924; no. 7102 (part), on Ficus, berg, J rum BL., Lour., Florida Blanca, Pampanga Prov., October 1925. Ae~ Aschersoiiia badia Pat. No. 2275 (part), on Preinna, Castilejos, Zani- Bataan bales Prov., .March 1924; no. 2945, on Preinna iiauseosa, Iligan, Isabela Prov., Prov., December 1923; no. 3151 (part.), on Piper, Asingan, Pangasinan lac. cit Prov., November 1923; no. 4503, on Ficus, Iba, Zambales Prov., February no. 29C 1924; no. 6156 (part), on Preiniia iiauseosa, Angat, Bulacan Prov., No- Januar: vember 1924; no. 6571, on Preinna nauseosa, Paniqui, Tarlac Prov., Ja- l\ anila, nuary 1925; no. 7308 (part), on Pilea inelastoinoides, Baguio, Benguet (Cham. Prov., Decemher 1925. on Pipi Aschersonia samoensis P. Henn. No. 922 (red and yellow forms) on on Prer, Canarium villosuin (Bl.) Vìl., Orani, Bataan Prov., May 1923; no. 1847 no. 6151 (part), on Premna, .Manila, February 1924; no. 2341 (part), on unknowii 1924; ni Rubiaceae, Stotsenberg, Pampanga Prov., October 1923; no. 2408 (part), January on Ardisia, Olongapo, Zambales Prov., March 1924; no. 2646 (part) on October Baguio, o T. Petch ',\', Some Philppine Entomogenous Fungi. ovember 1923; no. 3298, on Callicarpa, 121 ~h 1924; no. 3378, on Ad'enia coccinea pliyllantiZus reticulatus, Polo, Bulacan Prov., September 1923; no. 2740, ., October 1923; no. 4749, on unknown on Homonoia riparia Lour., Bosoboso, Rizal Prov., February 1924; no. 2803, ~924; no. 6257, on Preinna odorata, Ge- on Ficus, Tuguegarao, Cagayan Prov., January 1924; no. 2827 (part), on 5; no. 6375, on Pterospermuii niveuni Zingiber, Cagayan Prov., .January 1924; no. 3060 (part), on Antidesra bel' 1924; no. 6381, on Allaeanthus luzo- bunius (L.), Del Norte, Manila, January-February 1924; no. 3088 (part), 1924; no. 6496, on Jasiiinuii Sambac, on Psycliotria luzoniensis (Cham. & SChlecht.), loco cit., January-February 1924; no. 7113 (part), on Ichnocarpus 1924; no. 3118 (part), on Ficus, Asingan, Pangasinan Prov., November a Prov., October 1925; no. 22119 (part), 1923; no. 3169, on Litsea, loc. cit., November 1923; no. 3430 (part), on Prov., colI. M. Ramon, November 1913. Oreocnide trinervis (Wedd.), Bambang, Nueva Vizcaya Prov., January No. 1855, on unknown leaves, Manila, 1924; no. 4653 (part), on Streblus asper Lour., Porac, Pampanga Prov., ri Premna, Castilejos, Zambales Prov., October 1923; no. 4846, on unknown leaves, Nueva Ecija Prov., January tbela Prov., January 1924; no. 5656, on 1924; no. 4906 (part), on Lepidopetalum Perrottetii, Tarlac, Tar~ac Prov., ridanao, June 1924; no. 6210, on Ficus, December 1924; no. 5944, on Celastrus paniculata, , Pangasinan Prov., February 1925; no. 6019 (part), on Premna nauseosa Blco., Santa :schersonia placenta B. et Br. Maria, Bulacan Prov., Noyember 1924; no. 6566 (part), on Flemingia stro- No. 1847 (part), on Preiina, Manila, bilifei-, Paniqui, Tarlac Prov., January 1925; no. 7113 (part), on Ichno- llknown Rubiaceae, Stotsenberg, Pam- carpus volubilis, Florida Blanca, Pampanga Prov., October 1925; no. 22 119 )9 (part), on Zizyplzus talanai, Nueva (part), on Cyathea,Mt. Isiuog, Camarines Prov., colI. M. Ramon, No- l23-24; no. 2639, on Ficus, Polo, Bu- vember 1913. 346 (part), on Phyllanthus reticulatus, Aschersonia Oava Petch. No. 2584, on unknown Urticaceae, Stotsen- i, on Premna odorata BIco., loc. cU., berg, Pampanga Prov., October 192R. ia, loco cit., September 1923; no. 2972, Aschersonia Coffeae P. Henn. No. 2959, on Psidium Guajava 1., Ilgan, 3, Manila, September-October 1923; Isabela Prov., December 1923. eles, Pampanga Prov., October 1923; Torrubiella luteorostrata Zimm. Sterile stromata only. No. 609, on iis (Wedd.), Bambang, Nueva Vizcaya Streblus asper Lour., Tarlac Prov., March 1923; no. 2583 (part), on on Lepidopetaluin Perrottetii, Tarlac, Semecarpus, Stotsenberg, Pampanga Prov., October 1923. 92, on Cleinensia inacrantlza, Mt. Apo, Peziotrichum Lachnella Sacco No. 2583 (part), on Semecarpus, Stotsen- no. 6028, on Glochidium rubruni BL., berg, Pampanga Prov., October 1923; no. 4653 (part), on Streblus asper bel' 1924; no. 7102 (part), on Ficus, Lour., Porac, Pampanga Prov., March. 1923. tober 1925. Aegerita Webberi Fawcett. No. 970, on unknown Rubiaceae, Orani, Bataan Prov., May 1923; no. 2408 (part), on Ard'isia, Olongapo, Zambales (part), on Premna, Castilejos, Zam- on Premna nauseosa, Iligan, Isabela Prov., March 1924; no. 2412 (part), on Hydnocarpus subfalcata Merr., trt.), on Piper, Asingan, Pangasinan lac. cit., March 1924; no. 2472 (part), on ?Guioa, loc. cit., March 1924; r?icus, Iba, Zambales Prov., February no. 2909 (part), on Zizyphus talanai, Nueva Ecija Prov., December- auseosa, Angat, Bulacan Prov., No- January 1923-24; no. 3060 (part), on Antidesma bunius (L.), Del Norte, Manila, January-February 1924; no. 3088 (part), on Psycho 'luseosa, Paniqui, Tarlac Prov., Ja- tria luzoniensis ~ea melastomoides, Baguio, Benguet (Cham. & Schlecht.), loc. cit., January-February 1924; no. 3151 (part), on Piper, Asingan, Pangasinan Prov., November 1923; no. 6019 (part), No. 922 (red and yellow forms) on on Premna nauseosa Blco., Santa Maria, Bulacan Prov., November 1924; Bataan Prov., May 1923; no. 1847 no. 6156 (part), on Premna nauseosa, Angat, Bulacan Prov., November 1924; no. 2341 (part), on unknowii 1924; no. 6566 (part), on Flemingia strobilifera, Paniqui, Tarlac Prov., rov., October 1923; no. 2408 (part), January 1925; no. 7102 (part), on Ficus, Florida Blanca, Parnpanga Prov., T., March 1924; no. 2646 (part) on 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 species. From what experience he has had with phanerogamic , he is forced to 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 is easy 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 ends in 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 almost only hypothetical links between them, or none at alL. Every cryptogamist admits the essential truth of this broad statement, as applied to the lower cryptogams. The necessity of some kind of nomenclature, however, even for the most primitive or rudimentary organisms, obliges one to cling to the cnstomary 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. Evidently mycologists seem to have extended the conception of species to the fungi without questioning the feasibility of so doing. For two phanerogamic plants to be recognized as belonging to separate species, certain require- ments must be fulfilled. Primarily, they must differ in a certain, but not too small number of characteristics. In addition there should be no inter- mediate forms. Furthermore, hybrid forms, if any exist, should be more or less sterile. When the levelling effect of intraspecific crossfertilization is excluded, there appears the phenomenon known as J ordanism, i. e. the Linnean species resolve into numerous smaller, more or less recognizable units, which are necessarily constant, since reproduction is apogamous. How much of all this is applicable to fungi? Evidently the fathers of mycology, being fundamentally phanerogamists, having in most cases only widely distinct forms and isolated specimens 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 attention to 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 were :ion of species in mycology. described as, first, forms, or varieties, then, as species. When the ten- . Ciferri. dency to distinguish the fungi according to their host plant came into vogue, the application of the first requirement was also greatly limited, at ntention to discuss again the apparently least for certain groups of fungi. As far as we know, the third require- y constitutes a plant species. From ment for specific rank in the higher plants (surely, the crucÎal criterion in th phanerogamic plants, he is forced the of the phan:erogams), is of but doubtful application in nits called "Linnean" species, although mycology. A strict interpretation of this law for fungi is nonsense. On only too often have to depend upon the other hand, the true significance, in Nature, of heterothallism and of ize them. Nevertheless, in some groups, pleomorphism is still unknown. B ri e l' 1 e y' s point of view on the in- scarcely exist. If the conception of applicability to fungi and bacteria, of genetic concepts derived from the I to certain phanel'gamic genera, it is study of the highest organisms, is to a high degree justified. We can when it is applied to the lower crypto- admit in fungi the existence of pure lines or cross-fertiized generations, itO the realms of the microscopic orga- alone or combined, with every transitional stage between these extremes. nition of species, until at last all ends The transference of the conception of species from the phanerogamic GO deal with is not a number of well- plants to the fungi, leads to the conclusion that true Linnean species or 3,eroganiic sense, but rather groups of .Tordanian species, in a phanerogamic sense, do not exist in mycology. ~d the one with the other, with almost The last recognizable group in mycology is that of a more or less arti- :m, or none at all. Every cryptogamist ficially distinguished, small aggregate of individuals, resembling, in most oad statement, as applied to the lower cases, Jordanons rather than Linneons, to use the traditional phanero- gamic terminology. nomenclature, however, even Ïor the From this fundamental ambiguity in the interpretation and, of course, ~'anisms, obliges one to. cling to the i~ the limitation of the specific unit in mycology, comes the greatest indi- ie they ever so difficult of application. vidual variation in the selection and use of criteria for specific rank in ) phanerogams, the basic unit of taxo- creating species of fungi, polarized in two opposite extremes, the "pure" ~he species, but a kind of "species", morphologic, and the "pure" biologic, with- every intermediate shading. s among the phanerogams. Evidently The first and oldest was gradually subjected, according to the criterion of the conception of species to the fungi most mycologists, to the interference of the second, but the tendency to of so doing. For two phanerogamic multiply species according to the host plant has reached such proportions r to separate species, certain require- that a general review of the situation may well be demanded. Dr. But Ie l' Ghey must differ in a certain, but not recently ilustrated (Proc. Tnt. Congr. Plant Sc., VoL II, pp. 1590-1597; In addition there should be no inter- 1929) the confusion produced by the multiplicity of the criteria employed, forms, if any exist, should be more which has given rise to the species created during the last twenty years. ect of intraspecific crossfertilization is The critical point in. the classification in mycology is the taxonomy Lenon known as Jordanism, i. e. the of parasitic species of fungi, involving the biologic interpretation of the us smaller, more or less recognizable species, the "cultural" criteria being derived from and included in the it, since reproduction is apogamol1s. former. For these, the problem can be reduced to a fundamental question: Ie to fungi? Evidently the fathers of Must we accept a biologic interpretation in the taxonomy of the para- irogamists, having in most cases only sitic fungi? If so, can specific rank be maintained for biologic units? specimens for study, described as 1) "Habitus" is unterstood both from macroscopic and microscopic point of view. 124 R. Ciferri An exclusive morphologic application applied to obligate parasitic fungi, as defined by the group of neo-morphologists (C u n n i n g ham, e t The a. 1.), is, according to Our views, scarcely justified. One cannot return to each di¡ the primordia of mycology, and renounce the application of those prin- Ascomyi ciples in classification derived from fifty or more years of investigation of and, for the mutual relation between fungus and host plant. In the parasitic fungi, host pIa as well as in fungi with highly developed biochemical activities (such as applied. the yeasts), all phases of their life are subordinate to their biological func- of the in tions, a complex morphologic development frequently being sacrificed to Specific i a more perfect adaptation to the host. The consequence is the primitive- gical diff ness of many parasitic fungi (caused by a regressive evolution or by a nol1- on matrÙ evolution), whose uniformity of morphology is not correlated with the are cham range and exaltation of biological function. A classification of parasitic or animal fungi on purely morphological grounds takes a one-sided view, perhaps symptom: that of the least important side, compared with the synthetic picture offered of the boi

by utilizing all characteristics of the organisms of Ascom: 1). In conclusion, the writer's opinion is that no rules can be concerted than thost for, nor limitations imposed upon the use of one or another or all criteria (e. g., Ta for the definitioIt and delimitation of systematic entities in mycology. The biochemic: Thus i1 present chaotic situation in this science, as ilustrated by But Ie r, is not of what sh due to the multiplication of systematic units (this multiplication being the natural outcome of the development of Our knowledge), but to their as in thost gronp. A inadequate taxonomic expression. In the solution of the latter problem, Dr. But i. two courses are open: (1) The adoptIon of specific rank for all systematic But the units in mycology, independent of the criterion or criteria employed, but with an expressed distinction between different kinds of species; or (2) the !!''neity of : tlir dispari adoption of specific rank for morphological units only, with one or more In the j inferior ranks for units established from criteria other than morphologic. siiut fungi The adoption of one of these two solutions, sanctioned by an inter- particular ¡¡ national agreement, while'not bearing upon the actual relationship between his reason in the species, would, at least, tend to make their taxonomy less beWildering. Even thE 1) An exclusive application of morphologic characteristics to the classification nirieties, an of parasitic fungi, e. g., to the Ustilaginales, as demanded by C u n n i n g ham, the fact tha1 beginning with the aggregation of the loose smuts of barley and ,vheat, might end is by no me: in the partial reconstruction of Us t i lag 0 c a I' b 0 and a few old "Sammel- taxonomic u: species", which would include most of the smut species, and affect members of Sub-groups, : all familes, from Graminaceae to Compositae. Furthermore, if host specialization should be valueless for the classification of parasitic fungi, it is very doubtful whether any more weight should be attached to the localization of the parasite ') The m phological diff, of the plant host. As rightly expressed by Dr. But i e r: "If we refuse to accept great difficulty the one in the classification, it is difficult to justify the use of the other". Con- be based on b sequently, the classification of certain groups, e. g., Dothideales, as presented by eta 1. Morphc The i Sse nand S y d 0 w, should not be considered. 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 oldest pa as well as the most important criterion, when dealing with herbarium ph materiaL. All morphological characteristics, however, are not of the same 2. C u It importance. One can distinguish between: behav 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- teristi defined and easily recognized structural differences. artifi c 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- site oj tively small differences in shape of any formed element. with ~ 3. B i 0 met ri c c h a r act e ri s tic s, based on small, but stil stati- These stically recognizable differences in size only. A. Ph; B. Biological characteristics, including all other than morphological nat ones. The biological criteria, originally most neglected, are now widely B. Bio used in distinguishing species"). cat They are of several kinds: ran 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 ana symbiotic phenomenon between fungus and living host plant. Ac- C. Ph~ cording to the nature of the different manifestations of their pa't"a- the sitism, they may be subdivided into: resi A. Specialization characteristics, based on the range of the parasitic Geographi adaptation of each fungus. According to the nature of the sitic fungi as methods employed, we can distinguish: distribution 0 a) S p e cia i i z a t ion cap a b i e 0 f d ire c t proof from Most of tl the outcome of cross-inoculation with positive result. taxonomy of t b) S p e cia i i z at ion cap a b i e 0 fin d ire c t proof by of the twent cross-inoculation with negative result (inability to infect a morphologic ( definite host plant under experimental conditions), or by proposed on n failure to produce natural infection under natural conditions a rule, with i used, and, as (e. g., in the field). c) Sup p 0 sed s p e cia i i z a t ion, not directly or iiidirectly to that of the proved, but based on a probable or possible analogy of beha- with the matr vior with a specialized fungus of known host range. found. No Spi alone, but seVt These three criteria are arranged in order of their im- portance. liarities. Very B. Ecological characteristics, based on the study of the localization alone, and non of the fungus in or upon definite organs of the plant host. Taxonomic C. Pathographic characteristics, based upon the study of the effect much employee hut varieties h, of the parasitic fungus (mechanical, teratological, metamorphic, hystolytic, etc.) on the host plant, as well as on the reaction of teristics (i. e. the same to the parasite. This characteristic is, for the most variety TOU1'E slig'htly larger ") A classification based upon the interrelation between two organisms, and takig into consideration the complex fungus plus the host plant, is a phyto- 4) These tliri pathological classification rather than a mycological one. Of course, this remark of instruments u may affect the classification only from a philosophical point of view. physical, chemic~ cases the division 128 R. Ciferri characteristics combined (e. g., U stilago longissiina iSchlecht. J Mey., type on 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 the variety foliicola P. Henn. on the leaves of the same plant host). The sub-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 less Ii /I with the same criteria as those of the varieties, generally based on ecolo- " gical and matrical characteristics (e. g., Entyloma linariae Schr. on Linaria 'i 1Jllgaris, with the form veronicae Wint. on Veroiiica, with amphigenous /, I' sori as an additional characteristic). 11 , ,'' Only one author has used race as a taxonomic entity, based on micro- i' , ,II morphologic and matrical characteristics combined (e. g'., U1'ocystis ane- ii' mones iPers.J Wint., on Anemone neinorosa, with the slightly morpholo- i1' 8 II! i gically distinct race ranunculi-1'epentis Bub., on Ranunculus repens). ~ :11 i Nom i n a n u d a were frequently created by the earlier authors as vi ,/I I \-arieties or forms, based, as a rule, only on matrical characteristics or on ~ "ii'i I some supposed specialization. o;j .,1!, i , In conclusion, new entities have been described promiscuously as ~ Ji' / species more often than units of inferior rank, with the result that there --'J T/fl / / is a considerable amount of confusion as to the real meaning of "species"

as applied to the Ustilaginales. ~ct itl/- ~ ~, In order to ilustrate recent splitting of Some classical species, the ~ ¡' ~ ~\\ writer has arranged the elements diagrammatically, according to the o ~\ \ principal characteristics upon which they have been based. Figure 1 .- _II \ ;; 'ii \ \ shows the species derived from Ustilago violacea iPers.J Rouss., as o iii \ recently described, chiefly on the basis of experimental cross-inoculation ctbJ iii" ,\ as presented by L i l' o. Of eleven species listed, including U. violacea s. :- Iil i stricto reduced to the smut attacking the type host plant, nine were based ;:~ ,I,1\ \1

upon the results of cross-inoculations, and two 'are micro-morphologic and 1" matrical speci8s (Fig. 1). Of the six species derived from U. hypodytes 1\' iSchlecht.J Fr. (Fig. 2), three were based on morphologically differentiated '1\ 'iIi, \ characteristics and on a supposed specialization on some unstated host I, ' plant; one on pathographic characteristics combined with the variation II i in the host plant; one on ecological characteristics and host plant; one on I' a supposed host plant specialization. Of the many species derived from II \1 Ustilago striaeformis iWest.J Niess!. or the cycle thereof, today divided I II almost entirely according to the host plant parasitized, Figure 3 ilu- II strates the actual situation of ten species. Three species were based on i I morphological differences, as well as on a supposed host plant speciali- I zation; four species were isolated upon the basis of ascertained host specialization; all other species upon a supposed specialization. 801'0- sporiimi saponariae Rud. was recently sub-divided into four species, in- The criteria for definition of species in mycology. 129 'Jngissima iSchlecht.J Mey., type z Liro on Triticum repens with I U. antherarum (Fries) Liro. . " " . perforation of the leaves). , Jcological characteristics (e. g., I i its of Triticum vulgare, with the , TBmiTn:) of the same plant host). The : / U. coronariae Liro " ." ..".". m employed in the Ustilaginales, '/ Ii "e been proposed, more 01' less " ;eties, generally based on ecolo- " 'i ityloma linariae Schr. on Linaria " JI I U. diamhorum Liro . . . . . :: m Veronica, with amphigenous 'II :: ,II )!qduSoqiBd ~o 'I ~ :onomic entity, based on micro- ;,, ..'" " i '" ~ombined (e. g., Urocystis ane- '"c. '': i U. lychnidis - dioicae (D. C) Liro ~a, with the slightly morpholo- o "" '" ~ :''' u'" l., on Ranunculus repens). '" cI 'ii I o tted by the earlier authors as 'i I ~ "ii ~ 1 matrical characteristics or on :: ~/' . I" L' o o Ii' I U. niva is iro...... )!.á°10)3: QI ~ ~/' i .. ri described promiscuously as oVJ t' Ii i '" ank, with the result that there :u 'I ;: , the real meaning of "species" 2 ~ it- -- U. pallida Liro . . . . . -0 '" td;\ -; of some classical species, the .. ~i\ mmatically, according to the o ~\ \ õ .- \1 I \ E , have been based. Figure 1 uO!iBZ!TB!);id s o ;; 11 I , J: o 1)\\ 'u silenes - nutantis (D. C) Liro . . . i violacea iPers. J Rouss., as p;isoddns -0 '" bf "I \ ~ experimental cross-inoculation C' " i .. :- ',', '" isted, including U. violacea s. t: 1\ \ -0 ;: ,I, I '" pe host plant, nine wer.e based '" I" i U."silenes-inflarae (D.C) Liro '." 'ü wo are micro-morphologic and a.'" 1,1 vi es derived from U. hypodytes II' '1\ morphologically differentiated 'iI I \ . uO!iii!TB!);ids ,. zation on, some unstated host p;iU!Bii;i)SV 0. 1\ \ U. stellariae (Sow) liro . . . . . , . ~ combined with the variation \1 iristics and host plant; one on II II ie many species derived from I cycle thereof, today divided II I, it parasitized, Figure 3 ilu- i U. superba Liro ...... Three species were based on I I supposed host plant speciali- I Iie basis of ascertained host i I U. violacea (Pefs.) Rouss. s" stricto .. J!.á°ToqdJoW .pposed specialization. Soro- divided into four species, in- 9 130 R. Ciferri The critE

pUTIin:)

I U. aculeata (Ul , , . i U. agrestis Syd. , I , I I , I , . I U. agrostidis-p, , i I i .. i I i J!qdUJÉ'oqiud ¡i I, , , , I .. , I I ~ 'I , u U. airae-caespit , .. l' i , :a ..0 1:1 , u. amplexa Syd. u 'i' , I ~ ~ :' i 0 , vi ,I i .. , '" , I .. .'1 ~ Vi ,', U. alopecurivor: 1 i V" I I I I vi I I ;; V ,'. I , I -0 ,ii , vi I I 0 Z ,,, I .. II 0. ,', I ~ , I '" 0.. ,,, , , i .. V" ,'i i .. " J!É'OIOJ3: 0 Q) "" , U. brizae (Ule) .. /I ~ "iI ~ , U. athenae Maire . òO ..U II i . '" tq I " II I .. './ i ~ ,/ i .. rJ ).1' .. '" '1 i ~ ~ !l/ 'êt-~~~ U. bromina Syd 8 i. II ~( .. ~I\ I U l,' -0 iU '\\'1\ i ~ '"t.li iiii '- , 'u d rd' '" .~i. \\\,,\ \ U. phlei - praten~ o i i . . acry i 15 Maire .. ..f/ I'II Ii ' 0. I, 0 " . II S 0 " ", ~ \ ' 0 0. ,\ ' .. UOPUZ!IUpddS oJ II I I' ~ II , \ ', -0 '0 1\ U. milii (Fuck.) 0 i' pdsoddns .. f/ I' b. I , "t , , C' , .. ;: I. .. , -0 . I I '0 , '" I i Vl , I U. Iygei '(Maire) .. \ , i Maire .ü , U. Salvei B. et .. ~ , 0. I I CJ i i , , i uo!iuZ!rE!Jdds i , , i N U. scaura Liro . \ PdU!UlldJSV 1 oj \ ¡. , i U. valesiaca (SchelL.) Maire .

~

t, )~ i, _i J!É'°roqdJow -r ~:j

.: ( 'Ii, ri The criteria for definition of species in mycology. 131

pllnlin:)

piJniin:: i U. aculeata (Ule) Liro. '. I . , I I , I on U. agrostidis - paluscris Dav. . on i'i I OJ i I Z )!adiii~oalTld ~ I i )!adiiJâ'oqiiid ' , " , , ":'" .! . , v l' , U. airae-caespitosae (Lind.) Liro ~ OJ 0 1/1 'Y :av .. '" en " I .. ~ :' i 8 vi iI I ~ on I: ' 0 OJ '- -;'" U. alopecurivora (Ule) Liro . OJ '; ::,' i os -0 .~ , I I 0 ,I, , ~ ''i J '" 0. Z ,', I J!â'°IOJ3: ~ ~ ,,, I 0 .. ..'" 'I' J b/ )!~OJO)g: 0 Q) "" l U. brizae (Ule) Liro . .- os b/ ~ '',1 , ~ os ~ './,:.' " '" u ;: '" ;: 8 'Si- fl',-~~~ U. bromina Syd. -0 8 ~t ~ -0 i. i'"1\ ' .. V 'ii' .. ~ uo!iiiz!pipdds 0 .. . ~~ \\,,' \ \ .. .. II I \ u. phlei - pratensis Dav. pdsoddns 0a 0 ~ 11,,I. ~ \' " -0 6 0 OJ 0 0. ''. ' .~ UO!lTlz!JTlpdds ~ C1 I, ' - ii \ OJ -0 U. milii Liro -0 OJ 'C II (Fuck.) pdsoddns on ~ ~ I' OJ .:; ' \ '0 " ~ , . OJ -0 , \ 0. on ' , en OJ \ \ 'ü \ U. Salvei B. et Br. 0.OJ , uO!iiiz!ITlpdds (/ , '" , pdU!iiiJd)SV J , 0. uO!liiZ!ITl!)dds \ ¡E , N U, scaura Liro . PdU!Tlll;))SV oi ii

J!â'oioqdJoW

)!~OloadlOW

9* 132 R. Ciferri The (

pmUjnJ i E. ancej , I' E. arnic I' ,1 , , S. alsinearum Cif. . . . . ;, I E. arno~ , , " , i,' / , ,,' I E. as(eri / J!i:duá'oi:iud -0 ", , :: i,', , " , i: , lU '" I,,: i E. bavar I .. "i,,' / I S. dianrhi Rabenh. '" , i c '::;; i E. bolro o , / o ','" ¡ ~ , , a. i I '" ~ i I '" o 1'," in / , ~ ).~/,',' J E. catan; / " E ,1,/1, i "" / I :: r. '','' J .. I,il,' " E. Davis ii;: , 'ii J!É'OjOJ3: o a. i:;; ~I,,' ""i" i v Î/ , S. dianrhorum Cif. '" o ~ /1 , .. ~ ,",,, . ~-- ~i/,/ I' ./ o ~ ii', i' ' E. eriger vi o l~i' I C ~' 8 "" ~¡// l o t, ;: 1~11 ~Il'" -0 ~ II, ~ o .t,i / E. eupa(i rJ ~\ ',. "õ "t;',,/ \\ " ~r--- E. heleni 8 \ \ , o \ .. S. gypsophilae Cif. . . . . --f' ~ . \ E .- . , uori¡¡z!jupdds o "1.~ i,\t"", '- " E. hierac l. I I c. C ''1," o \ \ pdsoddns i \ -0 lU ~. 111\\ Vl0. I \ , i 'E; u~ II 'i I i' E. incern o I \ lU l. I \ -0 \ II o i \ co ~ i'll C/ \ \ lU 8 1111'. 'ü \ i S. silenis - inflatae (2.) Cif. a.lU o i i . E. leontc i vi __ 1\ i :: Ii \ \ .i 1\ \ I C E. lepach i rT' \ i , ~ , uopuzt¡U!Jdds ,. 'i I öO Ii E. Maire¡ I ¡i \ PdU!UiJdJSV \ \ \ , : S. saponariae S. striCto \ E. madia,

E. parthe

E. Saccan

J!É'Ojoi:diow 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. . puniinJ i' " i . I E. arnoseridis Syd. . I' " '" "'i E. asteris - alpini Syd. -c "i ". )!4duJ~04lUd :: hi, " ¡: i: E. bavaricum Syd. v ll /,': ,i )!4duJ~04lUd c: '" ,"i, .. "" .- " Cif. . -0 c ','1,1 E. boltoniae :: 0 '.'"i 0V' c. 1'''1 " 0 OJ '" .. 1" " " ~ "I' J E. catananches Cif. E 1,/1' I :: :: rn 'Ii" 1 -0 1'11; i i c .. '" )!~oIO)3: 0 ;; f'i' I J E. Davisii Cif. . c. ¡: " '" (¡I,',' i v \ Q) 0 "'1, / )!~oIO)3: " .. E. erigerontis Cif. 0 Cl ii'" i i E Cf 0 l~,/ / 0 'i Wi I 8 ;: I,11 è' E. eupatorii Cíf. c -0 oV' 'L11 1/ // "' ~ 1/ -; il _ 8 t' ' ---. E. helenii Cif. Õ f / -0 -J¡' ~ E '".:: ~,i\til", '. -; UO!iuz!lul);)ds 0 E. hieracii Syd. c. C'ii," Õ p;)soddns -0 ~ Ill"'. '" E .¡::: Ut' II'i Ii i UO!iuz!¡ul);)ds 0 '" E. incertum Cif. c. -0 I II t' ,Ii' p;)soddns -0 " \ i '" '" 8 III i E. leootodontis Syd. 'ü \ I ' ,t '" 0 '" c. - , I -0 Cf ~ IIi '" .¡ I I \ '" c: ii E. lepachydis Cif. Û '"0. .. ~ 'I Cf \1 UO!iuz!lul);)ds bJ E. Maireanum Cif. . ¡; \ P;)U!UlJ;nsy , UO!iuz!lul);)ds \ , 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 differentiated S. dianthi Rabenh.; six in all. (Fig. 4). Four species were isolated upon terion or i the basis of micro-morphologic characteristics, and on the basis of host fication. ,,: hand, this plants attacked; one on ecological characteristics and a supposed host species", i specialization. dytes, s Under the caption, Entylonia calendulae (Oud.) De Bary, European mycologists included all Entyloma living on the members of the Com- The se positae family. On the other hand, American mycologists assigned the arrange th same function to E. coinpositaruni Farl. The two species, from the morpho- species", ( logical standpoint, are identicaL. S y d 0 w sub-divided the European some scale species according to the. genus of the host plant parasitized, and the nomic valt writer the American species, chiefly upon a similar basis. The result was given the J that both species are at present represented by nearly twenty-five species, second pIai of which twenty-one are sketched in Figure 5. Besides the supposed host characteris specialization, common to all derived species, eight were based on small cies", and morphological differences; none on ascertained host specialization, nor on perts, char ecological, pathographic or cultural characteristics. A tenta The writer trusts that the ilustrations given above wil show in how Taxonomic many different senses the term "species" has been used in mycology. No Taxonomic doubt, mycologists would welcome any effort toward a standardization of meti this term, although it must be admitted that a wholly consistent nomen- Taxonomic clature, closely reflecting the true value of any taxonomic unit" is an spec unattainable goal. Taxonomic Some relief from the present confusion may be obtained by the simple Taxonomic expedient of always indicating, by means of some abbreviation, w hat Taxonomic kin d 0 f s p e c i e s one has in mind. For instance, m. would indicate a Taxonomic morphologic species; ec. an ecologic species; pa. a pathographic species; Applyiii cu. a cultural species. A double or multiple abbreviation would indicate Figure 4, tl t,hat a species has been proposed on two or more characteristics; e. g., Sora m. ma. would indicate that the species was proposed on morphologic Soro (micro- or macro-morphologic or biometric) and matrical grounds, directly Sora or indirectly ascertained, or predicated on some supposed specialization. Soro Applying this method to the species embraced in Figure 2, we should So 1'

have: So 1" Ustilago valesiaca (Schell.) Maire, ma. pa. The inc( U. dactylidis Maire, ma. m. in rank ill U. lygei Maire, ma. ec. would not J U. athenae Maire, ma. m. zoological i U. aniplexa Syd., mai m. inferior uni U. agrestis Syd., mai A third lt is clear that this suggestion has the advantage of not changing the nomenclatu status of species already described, nor the rank, leaving on one side the became gen problem of elevating all varieties and forms to the dignity of species. The yan Hall levelling of all entities to the same specific rank, independent of the cri- take rank ii The criteria for definition of species in mycology. 135 old but very little differentiated terion or criteria upon which the species is based, is but a formal uni- Four species were isolated upon fication. "Species" wil always be as heterogenous as before. On the other ristics, and on the basis of host hand, this does not consider the "mother species" from which the "minor acteristics and a supposed host species", 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). ulae (Oud.) De Bary, European The second (and considerably more revolutionary) proposal, is to 5 on the members of the Com- arrange these different kinds of "species" as in.ferior units of the "mother erican mycologists assigned the species", or the 'old morphological "S am m e Is p e c i e s", according to 'he two species, from the morpho- some scale supposed to express con v e n t ion all y their relative taxo- o w sub-divided the European nomic value. Evidently, morphological characteristics would have to be Qost plant parasitized, and the g'iven the main consideration, while matrical characteristics would take a a similar basis. The result was :I second place, and so on. Only units based on broadly different morphologic ~d by nearly twenty-five species, characteristics (macro-morphologic species), would be maintained as "spe- re 5. Besides the supposed host cies", and these should be selected in cies, eight were based on small' each group by a committee of ex- tined host specialization, nor on perts, charged also with ranging the units inferior to the species. cteristics. A tentative scheme of arrangement could be as follows: Taxonomic entities based on macro-morphologic characteristics: SPECIES given above wil show in how tias been used in mycology. No Taxonomic entities based on micro-morphologic characteristics or bio- metric characteristics: ...... SUB-SPECIES ort toward a standardization of hat a wholly consistent nomen- Taxonomic entities based on matrical characteristics and ascertained of any taxonomic unit" is an specialization:...... ' VARIETY Taxonomic entities based on ecological characteristics: . SUB- VARIETY may be obtained by the simple Taxonomic entities based on pathographic characteristics: . . .. FORM ; of some abbreviation, w hat Taxonomic entities based on cultural characteristics: . . . . SUB-FORM , instance, m. would indicate a Taxonomic entities based on some supposed host plant specialization: RACE es; pai a pathographic species; Applying these criteria to the binomial indicated in the diagram Ie abbreviation would indicate Figure 4, the new arrangement would be: or more characteristics; e. g., Sorosporium saponariae Rud. (species). was proposed on morphologic S01"sporium saponariae Rud. subsp. silenis-inflatae (Zigno). and matrical grounds, directly Sorosporium saponariae Rud. subsp. gypsophilae (Cif.). some supposed specialization. S01"sporium saponariae Rud. subsp. dianthorum (Cif.). braced in Figure 2, we should S01"sporium saponariae Rud. subsp. diantki (Rabenh.). Sorosporium saponariae Rud. var. alsinearum (Cif.). Maire, ma. pa. The inconvenience of this method would be the very numerous changes in rank involved. However, "new" combinations for these inferior units would not necessarily have to be used, and the application of the rule of zoological nomenclature for new combinations, eventually limited to these inferior units, could be accepted in a botanical nomenclature. A third and last suggestion is the adoption of the so-called trinary ,dvantage of not changing' the nomenclature, adopted by zoologists and some American botanists, which rank, leaving on one side the became generally applicable in botanical taxonomy, under the auspices of to the dignity of species. The van H a i 1. In this case all derived units inferior to the species would rank, independent of the cri- take rank in order of their fundamental macro-morphologic species, selected 136 R. Ciferri, The criteria for definition of species in mycology. as proposed in the preceding paragraph, without considering the criterion employed in the definition of these units. Applying the trinary nomenclature to the entities indicated in Figure L we should have: Ustilago violacea (Pers.) Rouss. U. violacea antherarum (Fries). U. violacea coronariae (Liro). U. violacea dianthoritm (Liro). A b bot t, E. V. U. violacea lychnidis-.dioicae (DC.). XXI, 1931, U. violacea nivalis (Liro). Abbott, E. V. U. violacea pallida (Liro). XXIV, 1932 U. violacea silenes-in(latae (DC.). Agostini, An U. violacea silenes-nutantis (DC.). Istit. Bot. L U. violacea stellariae (Sow.). Agostini, Ai U. violacea superba (Liro). barinus COl Agostini, Ai 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- onicomicosi geneity of the "species" would be maintained, the first line containing the A p pel, O. Die morphological (macro-morphologic) characteristics, but without the sacri- (Rep. Proc fice of all other taxonomic entities established on different criteria, and -401.) with a minimum of change of the present nomenclature. Arnaud, G. J sites: CalI( XVI, 1930, Arnaud,G. LE dn Muséun Mangin. P Arnaud, G. T. 1931, p. 7' Arnaud, G. E Ie blotch £1 V égét. XV Arnaud, G. e thologie V Arnaud,G.e arbres d'al 43 fig., tal Arnaud, G. e II. (Annal Aschieri, E

lop us caui voL. III, 1\ Ashworth,l ture. (Tr: tab. VlI-