MARINE BIOLOGICAL LABORATORY. Received ^^g^s-t* 1940 Accession No. 4^504 Givenby C. V« llosby Co. Place, St, Louis, Mo. *;^*I4o book OP pamphlet is to be pemoved fpom the Ijab- oratory tuithout the permission of the Trustees. MEDICAL MYCOLOGY p ii MEDICAL MYCOLOGY FUNGOUS DISEASES OF MEN AND OTHER MAMMALS BY CARROLL WILLIAM DODGE, Ph.D. MYCOLOGIST, MISSOITRI BOTANICAL GARDEN; PEOFESSOR, HENRY SHAW SCHOOL OF BOTANY, WASHINGTON UNIVERSITY ST. LOUIS ILLUSTRATED ST. LOUIS THE C. V. MOSBY COMPANY 1935 COPTEIGHT, 1935, BY THE C. V. MOSBT COMPANY (All rights reserved) Printed in U. S. A. Press of The C. Y. Mosby Company St. Louis TO MY WIFE WHOSE CONTINUED ENCOUEAGEMENT AND HELP FOE A DECADE HAS MADE THIS BOOK POSSIBLE PEEFACE This work was undertaken at the suggestion of the late Professor Roland Thaxter while the writer was a member of the faculty of Harvard University. After a preliminary survey of the literature, a course in Medical Mycology was offered in 1924 and the book has grown out of that course. The manu- script was completed in February, 1932, but unavoidable delays in publishing have prevented its earlier appearance, though it has been used by my classes and research students for several years and has frequently been revised. It contains a summary of all the literature in this field to the end of 1933 except for a few scattered references in relatively rare periodicals which are marked by asterisks in the bibliographies at the ends of the chapters. I have also included all of the references available to me up to July 1, 1934, either at the library of our own Medical School or in the Boston Medical Library, but probably important papers in this period have been missed, as many titles had not found their place in abstracting and indexing journals ; work in my own laboratory during this period has been incorporated, even though the papers resulting from the work have not yet been published. During the correction of proofs I do not intend to include any references unless new names or combinations proposed in them affect the nomenclature of species I have already recognized. For the first time in this field, a relatively complete and accurate bibli- ography of existing literature is presented. In such a large bibliography, the abbreviation of names of journals presents a serious problem. The code of the Leagaie of Nations* was received too late to be followed. If the abbreviation be too brief, it may cause ambiguity to one who is not well acquainted with the journals of a given country or of a given subject, especially when journals have similar names. For example, Arch. Derm, might refer to Archives of Derma- tology and Syphilology, Archiv fur Dermatologie und Syphilis or Archivio Italiano di Dermatologia e Sifilologia. When Ann. is confused with Arch., as is regularly done bj'' the writer of one text in this field, we have also to consider Annales de Dermatologie et de Syphiligraphie and the Anales Dermatologicas y Sifiliograficas. More rarely we have several journals with the same name, when it is desirable to add the name of the editor, especially if the journals are contemporary. For example, the abbreviation Jour. Bot. is almost mean- ingless. It might refer to the Journal fur die Botanik, Journal de Botanique (Desvaux), Journal de Botanique (Morot), Journal of Botany (Hooker) or Journal of Botany, British and Foreign (edited successively by Seemann, Triemen, Britten and Rendle, and sometimes cited as Seemann' s Jour., etc. In cases where the usual abbreviation would indicate a journal only by the •League of Nations International Institute of Intellectual Cooperation, 1930. Interna- tional Code of Abbreviations for Titles of Periodicals. 12 pp. and Supplement 18 pp. 1932. 8 PREFACE presence or absence of a diacritical mark, I have not abbreviated owing to ease with vt^hich diacritical marks are overlooked in proofreading. The num- ber of the series has been given in Roman numerals, the volume number in boldface, and where the pagination is not continuous, the number of the article or part is given in ordinary type between colons. Some care has been taken to ascertain the dates of publications where new species or new combinations are proposed, since this date is the effective one for deciding questions of priority. If it is desirable to record the first isolation of an organism as a matter of historical interest, it should not be connected with the scientific name in bibliographical citations. From bitter experience one learns to disregard the dates given by certain authors, or to add from one to ten years to the dates given. When an author indulges in this practice with reference to his own new species and uses the date of publica- tion or even misquotes the latter in the case of species proposed by others, one cannot help but suspect intentional dishonesty. Another objectionable practice of some authors is the copying of a bibli- ography without verifying the citation or reading the article quoted. For example, many authors quote Bohin 1847 when they really mean Bohin 1853. In Robin's doctoral thesis, Les vegetaux qui croissent sur les animaux vivants, viii, 120 pp., 3 pis., Paris, 1847, he uses only the name Achorion Schoenleini, although he summarizes the work of previous authors very carefully. "When the thesis was reissued in book form under the title Histoire naturelle des vegetaux parasites qui croissent sur I'homme et sur les animaux vivants, x, 704 pp., Paris, 1853, with an Atlas of 15 plates, the text was greatly expanded and the various organisms were given scientific names. Hence all names attributed to Robin should be cited 1853 not 1847. It follows that when an author cites Robin 1847 for a species name, he is copying without having read the very rare thesis of that date. Such carelessness tends to throw doubt upon an otherwise acceptable piece of work. It is recognized that errors occur in proofreading, but it is felt that by giving complete bibliographic data as to volume and year, there is little chance that both will show the same typographic errors. The author will be grateful for corrections of errors, for information of the location of refer- ences marked with an asterisk or for references to significant work which has been overlooked or to new literature, looking toward a revision. The chapter on microscopy and staining has been kindly contributed by Dr. Morris Moore and the section on hydrogen ions (pp. 34-38) by my wife. The text-figures have been redrawn from original sources, duly acknowledged in each, by Dr. Gladys Baker, Mr. Albert Heinz, Dr. Morris Moore and the late Mr. Thomas O'Brien, most of those except the yeasts by the latter. The drawings by Dr. Morris Moore are the result of his own research. While the author assumes full responsibility for the statements of this book, he is grateful to Dr. Margaret B. Church of Urbana University, to Dr. Morris Moore of the Barnard Free Skin and Cancer Hospital of St. Louis, and to Dr. Joseph Swartz of the Medical School of Harvard University for PREFACE 9 reading the text of the chapters covering their respective fields and for their constructive criticism; to the late Professor Thaxter of Harvard University and to Dr. Charles Thorn of the Bureau of Soils for the kind interest and advice they have so generously given; to Mr. James F. Ballard and Miss Lotta McCrea of the Boston Medical Library and to Miss Ella B. Lawrence of the Library of the Washington University School of Medicine for their sympathetic help in finding incorrectly cited references ; to many of my former students for helpful suggestions ; to the Chancellor of Washington University and to the Director of the Missouri Botanical Garden for a leave of absence to complete reading in the files of rarer periodicals in Boston before begin- ning my duties in their respective institutions ; and to my wife who has helped throughout in the dreary tasks of preparing the manuscript for the printer, correcting the proofs and preparing the index. — C. W. D. St. Louis. August 1, 1935. CONTENTS CHAPTER PAGE I. General Morphology of Fungi ______________ 37 " Vegetative structures, 17; Eeproductive structures, 19; Sexual organs and sexuality, 21; Classification, 24; Bibliography, 25. 11. Physiology of Fungi With Special Reference to Reproduction _ _ _ _ 31 Water, 31; Inorganic salts, 32; Carbon, 33; Nitrogen, 33; Hydrogen ion concentration, 34; Oxygen requirements, 38; Temperature require- ments, 39; Influence of light, 39; Tropisms, 40; Radium, 40; Bibliog- raphy, 40. III. Culture Media, Their Preparation and Sterilization _______ 44 Cleaning glassware, 44 ; Sterilization, chemical methods, 45 ; Physical agents, 46; Culture media, 48; Liquid media, 49; Solid media, 50; Bibliography, 54. IV. Isolation op Microorganisms _______________ 57 Transfer, 57; Isolation from skin, 57; Isolation from feces, tongue scrapings, etc., 58; Dilution, 58; Inliibitors, 58; Microcultures, 59; Giant cultures, 60; Single cell cultures, 61; Ascospore detection, 61; Fermentation, 62; Animal and human inoculations and recovery of organisms from lesions, 64; Bibliography, 65. V. Microscopy (by Morris Moore) ______________ 66 Mounting media, 66; Spore stains, 67; Stains for fungi in skin, i Stains for fungi in other tissues, 68 ; Stains for hair and scrapings, 69 ; Stains for sputum, 70; Vital staining of fungi, 70; Fixing agents, 71; Parafiin method, 71; Nitrocellulose method, 71; Bibliography, 73. VI. Botanical Nomenclature ________________ 75 Historical sketch, 75; International Rules of Botanical Nomenclature, 76. VII. Phycomycetes ___________________ 97 Mucorales ____________________ 97 Mucoraceae, morphology _______________ 98 Mucor Micheli __________________ 110 Absidia Tieghem _________________ 111 Bhisopus Ehrenberg ________________ 115 Mortierella Coemans ________________ 118 Bibliography ___________________ 119 Vin. Ascomycetes __-______-_--------- 121 IX. Endomycetales __________________ 126 Spermophthoraceae _________________ 127 Ashbyaceae ___________________ 128 Piedraia Fonseca & Area Leao; Piedra of hair ________ 131 Pichiaceae _________--__------- 136 Guilliermondella Nadson & Krassilnikov __________ 136 10 CONTENTS 11 CHAPTER PAGE Ascoideaceae ___________________ 137 Endomycetaceae __________________ 137 HanseTmla Sydovv _________________ 142 Eanseniospora Zikes ________________ 143 Dipadascaceae ___________________ 145 Actonia Dodge _________________ 145 Coccidioideaceae __________________ 147 Coccidioides Stiles; Coccidioidal granuloma _________ 147 Bhinosporidiii/ni Minchin & Famthani ___________ 151 Histoplasma Darling ________________ 152 Paracoccidioides Almeida ______________ 155 Neoffeotrichum O. Magalhaes _____________ 156 Protomycetaceae __________________ 157 Taphrinaceae _ __________________ 159 X. Eremascaceae __________________ 161 Zymonema Beurmann & Gougerot ____________ 165 Oleina Tieghem _________________ 179 Octomyces Froilano de Mello & Gonzaga Fernandez _______ 180 Bargellinia Borzi _________________ 182 Hemispoi-a Vuillemin ________________ 182 XI. Eremascaceae Imperfectae ______________ 186 Methods of study, 186; Morphology, 190; Classification, 194; Eespira- tory infections, 204; Thrush, 204; Sprue, 205; Skin infections, 205; Keys, 206. Proteomyces Moses & Vianna _____________ 208 Geotrichum Link _________________ 215 Mycoderma Persoon ________________ 223 Candida Berkhout ________________ 229 Schizohlastospoi-ion Ciferri ______________ 234 Pseudomycoderma Will _______________ 235 Parendomyces Queyrat & Laroche ________ ____ 238 Castellania Dodge _________________ 246 Parasaccharomyces Beurmann & Gougerot _________ 265 Monilia Bonorden _________________ 270 Syringospara Quinquaud _______________ 272 Blast odendrion Ota ________________ 282 Beduellia Ciferri _________________ 289 Mycotoruloides Langeron & Talice ____________ 290 Mycocandlda Langeron & Talice ____________ 293 Pseudomonilia Geiger _______________ 295 Doubtful position _________________ 297 XII. Saccharomycetaceae ________________ 300 Schizosaccharomyces Lindner _____________ 307 Debaryomyces Kloecker _______________ 308 Saccharomycopsis Schionning _____________ 316 Sacchnromyces Meyen _______________ 317 XIII. Saccharomycetaceae Imperfectae ____________ 325 Cryptococcus Kuetzing ; Torula meningitis _________ 326 Pseudosaccharomyces Laer ______________ 841 "/(.So^ 12 CONTENTS CHAPTER PAGE Atelosaccharomyces Beurmann & Gougerot _________ 342 Torulopsis Berlese _______ _________ 347 Eutorula Will _________-_____--- 354 Asporomyces Chaborski _______________ 356 Microilastosporion Ciferri ______________ 357 Trigonopsis Schachner _______________ 357 XIV. MALASSE.ZIA BaILLON; CLINICAL DISCUSSION OF SEBORRHEA AND ACNE _ 358 Bibliography of Endomycetales _____________ 371 XV. Plectascales _________-_____--- 425 Gymnoascaceae __________________ 425 Ctenomyces Eidam ________________ 429 Gymnoascihs Eidam ________________ 430 Ateleothylax Ota So Langeron _____________ 431 XVI. Trichophytoneae (Gymnoascaceae Imferfectae) _______ 425 Clinical discussion of lesions produced on epidermis, 435; Tinea im- bricata, 436; Tinea circinata, 436; Eczema marginatum, 438; Tinea unguium, 440; Lesions of hair and hair follicle, 440; Favus, 441; Tinea tonsurans, 444; Microsporum infections, 444; Trichophyton in- fections, Sabouraudia type, 445 ; Malmstenia type, 445 ; Neoendothrix type, 447; Sycosis, 448; Kerion Celsi, 448; Granuloma of Majocchi, 448; Colony characteristics, 449; Morphology of organisms, 449; Variations and mutants, 455; Pleomorphism, 456; Phylogeny, 457; Classification, 462; Geographic Distribution, 465; Physiology, 466; Therapeusis, 471; Immunology and related phenomena, 474. Pinoyella Castellani & Chalmers ____________ 476 Epidermophyton Sabouraud ______________ 477 Endodermophyto'n Perry _______________ 489 Ectotrichophyton Castellani & Chalmers __________ 493 Megatrichophyton Neveu-Lemairc ____________ 509 Favotrichophyton Neveu-Lemaire ____________ 512 Trichophyton Malmsten _______________ 527 Microsponom Gruby ________________ 537 Achorion Eemak _________________ 551 Bibliography _ __________________ 562 XVII. Aspergillaceae ___________------- 608 Aspergillus Micheli ________________ 621 Penicillvum Link _________________ 639 Paecilomyces Bainier ________________ 642 Scopulariopsis Bainier emend. Thom ___________ 643 Phaeoscopulariopsis Ota _______________ 651 Allescheria Saccardo & Sydow _____________ 652 Onygenaceae __________--------- 653 Bibliography __________--------- 653 XVIII. Fungi Imperfecti — Hyphomycetes _____________ 665 Mucedinales __________---------- 665 XIX. Toruleae. ____________------- 669 Coniosporiww. Link ___________..__-- 673 Pullularia Berkhout ____________---- 67 3 CONTENTS 13 CHAPTER PAGE Dcmatkim Persoon; Carate or pinta ___________ 676 Madurella Brumpt; Mycetoma _____________ 680 Indiella Brumpt _________________ 685 Bibliography ___________________ 687 XX. ACTINOMYCETEAE __________________ 694 Morphology, 694; Methods of study, 702; Classification, 703. ActinoTnyces Harz _________________ 705 Bibliography ___________________ 767 XXI. Sporotrichieae __________________ 786 Aleurisrna Link _________________ 786 Trichosporium Pries ________________ 791 Acremonium Link _________________ 795 Acremoniella Saccardo __________--^__ 798 Sparotrichum Link; Sporotrichosis ___________ 798 Bibliography ___________________ 811 XXII. Chalareae ___________________ 822 Chalara Corda __________________ 822 Cephalosporieae __________________ 823 Syalopus Corda _________________ 824 Cephalosporium Corda _______________ 826 Corethropsis Corda ________________ 831 Phialophoreae __________________ 833 Phialophora Thaxter ________________ 833 Gonatobotrytideae _________________ 834 Thomiella Dodge _________________ 834 Botrytideae ___________________ 835 Acladium Link _________________ 836 Monosporium Bonorden _______________ 837 Verticilleae ___________________ 841 Spicaria Harz __________________ 843 Haplographieae __________________ 843 Catenularia Grove ________________ 843 Eaplographium Berkeley & Broome ___________ 844 Eoi-7nodendron Bonorden _______________ 845 Periconieae _______________---- 850 Gomphmaria Preuss ________________ 850 Hyalodidymeae __________________ 851 DiplosporiMm Link ________________ 853 Phaeophragmieae ___________-__---- 853 Acrothecium Preuss ________________ 854 Spondylocladi'n/m Martins ______________ 855 Phaeodictyeae ____________------- 856 Altcrndria Nees ab Esenbeek _____________ 856 Stilbaceae _________-_--------- 857 Dendrostilbella Hohnel _______________ 858 Tilachlidmm Preuss __________------ 858 Tuberculariaceae _________--------- 858 Fusarvum Link __________--_---- 859 Doubtful position _______-_--------- 860 Bibliography ______------------- 861 ILLUSTRATIONS FIG. PAGE 1. Hypnospore formation ___.__________-__-- 18 2. Oidial formation ____________________ 20 3. Coremium _______________________ 21 4. Sporangia __________________----- 99 5. Showing the development of the sporangium of Sporodinia grandis ______ 100 6. Blakeslea trispora ____________________ 102 7. Syncephalastrum cinereum __________________ 103 8. Mortierella niveovelutina ______________---- 105 9. Zygospores ______________________ 106 10. Development of the zygospore of Sporodinia grandis __________ 107 11. Pyronema confluens ____________________ 122 12. Pyronema confluens ________________---- 123 13. Spore shapes in the Endomycetales __________--_-- 127 14. Spermophthora Gossypii __________________ 128 15. Piedraia Hortai _____________________ 129 16. Eremothecvwm Gossypii ____________--_---- 130 17. Nematospora Coryli _________ __________ 131 18. Endomyces Magnusii ___________________ 138 19. Endomyces decipiens ___________________ 139 20. Endomycopsis fibiiliger ___________________ 140 21. Endomycopsis Lindneri __________-___----- 141 22. Dipodascus albidn,s ____________________ 144 23. Coccidioides immitis (Pseudococcidioides Mazzai) ___________ 148 24. Coccidioides immitis _______________----- 150 25. Ehinosporidiiom Seeberi ___________________ 152 26. Mistopla-sma capsulatum ___________-__---- 153 27. Histoplasmu pyriforme ______________----- 154 28. Protomyces macrosporus ____________-__--- 158 29. Taphrina deformans _____________------- 159 30. Eremascus fertilis ______-____--------- 162 31. Eremascus albus ______________------- 162 32. Zymonema capsulatum _____-______-_----- 163 33. Zymonema dermatitidis ___________-------- 164 34. Oleina nodosa Tiegh _______________---- 165 35. Zymonema Harteri _________-_--------- 178 36. Hemispora stellata __________---------- 183 37. Hemispoi-a coremiformis ____________------ 184 38. Blastodendrion intermedvum _________________ 189 39. Types of blastospores ________----------- 192 40. Showing Shrewsbury 's cell types in Eansemda ____________ 193 41. Proteomyces infestans ________-__-------- 209 42. Proteomyces asteraides _______-____------- 211 43. Proteomyces cutaneus _______------------ 212 44. Proteomyces Balseri __________--------- 213 45. Geotrichum ________-------------- 216 46. Geotrichum versiforme ________----------- 222 47. Candida {Geotrichoides Langeron & Talice) ____________ 230 14 ILLUSTRATIONS 15 FIG. PAGE 48. Candida Krusei _____________________ 232 49. Monilia (Candida Langeron & Talice) ______________ 271 50. Syringospora {Mycotorula Langeron & Talice) ___________ 273 51. Syringospora clilamydospores _________________ 277 52. Blast odendrion intermedium _________________ 284 53. Blastodendrion Pinoyi ___________________ 286 54. Eedaellia elegans Cifeni __________________ 289 55. Mycotoriiloides _____________________ 290 56. Mycocandida ______________________ 294 57. Schizosaccharomyces octosporus ________________ 301 58. Zygosaccharomyces Chevalieri _________________ 301 59. Torulaspora Bosei ____________________ 302 60. Saccharomyces cerevisiae __________________ 302 61. Nadsonia fulvescens ____________________ 303 62. Debaryomyces Kloecheri __________________ 303 63. Saccharomycodes Ludwigii _________________ 305 64. Debaryomyces Hudeloi __________________ 310 65. Debaryomyces Leopoldi ___________________ 313 66. Saccharomycopsis guttulatus _________________ 316 67. Saccharomyces gramdatus __________________ 318 68. Saccharomyces anginae __________________ 319 69. Saccharomyces annulatus __________________ 323 70. Malassesia ovalis ______ ______________ 368 71. Amuuroascus verrucosus __________________ 426 72. Gymnoascus setostis (Eidamella spinosa) _____________ 427 73. Gymnoascus gypseus and Ctenomyces serratus _____----___ 427 74. Ctenomyces serratus ___________________ 428 75. Section through hair from a case of favus, caused by Acharion Schoenleini _ _ _ 442 76. Section through hair from a case of tinea tonsurans microsporica, caused by a species of Microsporum ___________________ 444 77. Section through hair from case of tinea tonsurans __________ 447 78. Mycelium ______________________ 449 79. Nodular organs _____________________ 450 80. Arthrospores _____________________ 451 81. Pedicellate chlamydospores _________________ 452 82. Intercalary chlamydospores _________________ 452 83. Showing closterospores and their transition to chlamydospores ______ 453 84. Aleurospores ______________________ 454 85. Endodermophyton Boquettei _________________ 492 86. Favotrichophyton camerounensis ________________ 516 87. Microsponmi ferrugineiim _________________ 546 88. Aphanoascus cinnaiarimis _________________ 609 89. Monasous ruber _____________________ 610 90. Diagrammatic radial sections of colonies _____________ 611 91. Diagrammatic radial section of a colony of Paecilomyces Varioti ______ 612 92. Thielavia basicola ____________________ 613 93. Types of phialides ____________________ 614 94. Conidial stages _____________________ 615 95. Gliocladiitm roseum ___________________ 616 96. Penicillium Brefeldianum __________________ 616 97. Aspergillus nidulans ___________________ 617 98. Penicillium Wortmanni ___________________ 618 16 ILLUSTRATIONS PIG. PAGE 99. The ascospores of AspergilMs ________________ 620 100. Aspergillus Amstelodami __________________ 631 101. Aspergillus Jeanselmei __________________ 638 102. Penicillium Bertai ___________________ 640 103. Paecilomyces Varioti ___________________ 643 104. Scopulariopsis brevicaulis (Sacci.) Bainier ____________ 644 105. Alternaria sp. _____________________ 666 106. Hormiscium stilbosporum Corda and Eormiscium pinophilum Nees _____ 672 107. Pullularia Jeanselmei ___________________ 675 108. Demati/wm articulatum Pers. _________________ 676 109. DematiAim, hispidulum __________________ 677 110. Madurella mycetomi ___________________ 680 111. Indiella Mansoni ____________________ 686 112. Actinomyces II isolated from soil _______________ 696 113. Actinomyces XVIII ___________________ 698 114. Actinomyces alius (A. griseus Krainsky?) ____________ 699 115. Actinomyces XVIII ___________________ 700 116. Actinomyces OMreaus ___________________ 701 117. Actinomyces Lavendulae __________________ 702 118. Actinomyces scahies (Thaxter) Giissow _____________704 119. Acremoniwm alternatum Link ________________ 796 120. Acremonvum Potronii Vuillemin _______________ 797 121. Bhinocladium coprogenum Saccardo & Marclial ___________ 800 122. Sporotrichum, Schenki var. Beurmanni _____________ 807 123. Sporotrichum Lesnei Vuillemin ________________ 810 124. Thielaviopsis paradoxa __________________ 823 125. Chalara fusidioides Corda _________________ 823 126. Eyalopus muscorum Corda _________________ 825 127. Cephalosporium. Acremonvum Corda ______________ 827 128. Cwethropsis paradoxa Corda ________________ 832 129. Phialophora verrucosa Thaxter in Medlar ____________ 833 130. AcladiAim conspersum Link _________________ 836 131. Acladvum Castellanii ___________________ 836 132. Monosporium spinosum Bonorden _______________ 838 133. Monospwvum apiosperrmim Saccardo ______________ 839 134. Verticillvum agaricinum, (Link) Corda _____________ 842 135. Eaplographium chlorocephalum (Fresenius) Grove __________ 844 136. Hormodendron olivaceum Corda _______________ 846 137. Hormodendron Fontoynonti _________________ 847 138. Gomphinaria Pedrosoi __________________ 851 139. Arthrohotrys superia Corda _________________ 852 140. Trichothecium roseum Link _________________ 852 141. Diplosporinim vaginae __________________ 854 142. Acrothedum nigrwm ______---_-_---- _- 855 MEDICAL MYCOLOGY CHAPTER I GENERAL MORPHOLOGY OF FUNGI The Fungi form a large heterogeneous group of plants, including all those lacking chlorophyll, which are not closely nor obviously related to other groups. In a few primitive families, the vegetative body is naked and amoeboid. In the rest of the fungi, it is surrounded by cell walls and usually appears as septate filaments, called hyphae. The vegetative hyphae are collectively known as mycelium. Under certain conditions of nutrition, as in solutions of very high or of very low osmotic pressure, the hypha grows by sprouting, when a small protuberance enlarges, rounds off and is abjointed (cut off by a septum) from the mother cell. The daughter cell, called sprout cell or blastospore, continues to increase in size and eventually separates from the original group of cells. In certain groups, as in the yeasts, no other type of vegetative body is known. When conditions for growth are unfavorable, the protoplasm contracts, rounds up and secretes a special thick wall, forming resting cells, called hypnospores or chlamydospores. (Fig. 1.) With the re- turn of favorable environmental conditions, these hypnospores again develop normal vegetative mycelium. In a few groups of fungi, the hyphal wall gives the cellulose reaction, in most others that of chitin. In fructifications and resting cells, the hyphal wall first appears as a thin, hyaline membrane which becomes thicker and may be further differentiated by secretions and deposits of minerals or resins, or colored by pigments. A relation between the fundaments of the wall and mitosis has been demonstrated in only a few cases, as in ascospore formation. In general, the wall is gradually differentiated from the cytoplasm without nuclear aid. The septum often forms by furrowing (annular thickening of the walls, like an iris diaphragm in a camera) . For the maintenance of inter- cellular communication, the septa are frequently pierced by a few openings through which pass protoplasmic threads. During rapid growth, there may be a delay in the formation of septa, later compensated by simultaneous or successive development of septa. In the Phycomycetes, the septa are wholly suppressed ; the whole mycelium is then a single, branched, multinuclear mass of filaments, becoming septate during the formation of reproductive organs, or during conditions of poor nutrition or of senescence. The individual hyphae generally are intertwined in feltlike masses. Such a group of hyphae, called the mycelium, usually absorbs food at any point 17 18 MEDICAL MYCOLOGY over its whole surface. Small mycelial branches, which serve to attach the mycelium to the substrate and to absorb the nutrients, have become special- ized in form and function in several groups. When the absorbing organs are rootlike, filamentous, finely branched and tapering, they are called rhizoids (frequent in the Chytridiales). When they are intracellular, clavate, bluntly lobed, coarsely branched or coralloid organs of parasites which do not imme- diately injure or kill the host cell, they are referred to as haustoria (frequent Fig-. 1. — Hypnospore formation in 1 ; 2, Mucor racemosus; S, M. dimorphosporus ; i, Rhizopnis arrhisus. (After Lendner 1908.) in plant parasites). When the function seems to be purely one of attachment and the mycelial branches are flattened and disciform, bluntly lobed or branched, or even coarsely filamentous, they are called holdfasts (e.g., Bhizopus of the Mucorales). Appressoria are holdfasts which are superficial or non- penetrating, although they may be intercellular (frequent in the sooty molds). In many cases, the hyphae grow together in groups, intertwine, adhere, and form a thick tissue, called plectenchyma. If the single hyphal elements are GENERAL MORPHOLOGY OF FUNGI 19 still recognizable as such, they are referred to as prosench3rma ; if the hyphae have lost their individuality so that they lie adjacent, with the cells (in sec- tions of the tissue) appearing isodiametric and continuous, they are called pseudoparenchyma since they are formed by cell division in a single plane while the parenchyma of higher plants develops from cell division in three planes. Sclerotia are small hard masses of plectenchyma with a firmer pseudo- parenchjrmatic rind and a looser prosenchymatic core. These structures serve to carry the organism over unfavorable environmental conditions and, with the return of normal conditions, germinate to the usual mycelium or to a fructification. Sclerotia are formed on drying out of the culture in many species of Aspergillus. Bulbils are small sclerotia formed of a few layers of cells and are often present in large numbers. The grains in the lesions of mycetomas (Madura foot, etc.) belong in this general category. Reproductive Structures. — In most fungi, at a definite age and imder favorable conditions of nutrition, reproductive structures develop on the mycelium. The products of the reproductive processes are chiefly spores. Spores may be defined as characteristically formed cells or groups of cells which separate from the mother plant and may grow independently to new individuals. They serve either for propagation (multiplication and dispersal) or for resisting unfavorable conditions of the environment (prolonged desic- cation, overwintering, etc.). Spores specialized for the latter function are often called hypnospores (Fig. 1). In the simplest case, hyphal cells separate from the parent hyphae and develop into new hyphae. These individual cells are called arthrospores or thallospores and are homologous with the cells of a chain of blastospores, though the latter arise by sprouting rather than by the breaking apart of the cells of a hypha (Fig. 2). From arthrospores there is a gradual transition to more typical spores with characteristic color, form, and sculpturing of the wall. In many cases they are abjointed directly from the cells of an ordinaiy hypha ; in other cases they arise on specialized sporophores. AVhere these sporophores form the spores within specialized sporogenous cells, sporangia., they are called spor- angiophores and the spores, if they are enclosed and nonmotile, sporangia- spores (e.g., in the Mucorales). Sporophores which abjoint their spores ex- ogenously at their tips are referred to as conidiophores and their spores conidia (e.g., in the Fungi Imperfecti). A chlamydo spore is any thick-walled asexual spore without further regard to its morphologic significance. In the higher fungi, the mycelium surrounding a group of conidiophores (or sexual organs) is known as a fructification. When these groups are in fascicles, they are called coremia (Fig. 3) ; if they form widespread cushions, they are called sporodochia (Tuberculariaceae) or acervuli. The tissue from which they arise is then known as their stroma. When the conidiophores develop in cavities in the stroma, the finictifications are called pycnia, and the conidia are often called pycnospores or stylospores. These different spore 20 MEDICAL MYCOLOGY forms, blastospores, arthrospores, chlamydospores, conidia, etc., usually de- velop on the liaplont (thallus of the haploid stage of the life cycle). A single species which successively produces several spore forms is called polymorphic. The spores of a second type are connected with sexuality and develop after fertilization or meiosis in the spore mother cell. These changes are connected respectively with the beginning or the end of the diploid phase. The spore forms following fertilization are recognizable morphologically since they are encysted zyg-otes (the products of a sexual act) ; biologically they usually develop as hypnospores or resting spores (e.g., the zygospores of the Mucoraeeae) . The spore forms which follow meiosis are morphologically recognizable because they form tetracytes (as daughter cells of gonotoconts, the organs in Fig. 2. — Oidial formation in i; S, Mucor racemosus ; S, Mucor Prainii. de Botanique.) (After Chodat, Principes which meiosis occurs). Apparently, since they are products of meiosis, they have become constant in number, usually 8 (in the Ascomycetes) or 4 (in the Basidiomycetes) ; biologically they are also hypnospores in the higher fungi. When the sporogenous cells which serve as gonotoconts form their spores internally through free cell formation, they are called asci and their spores ascospores (whence the term Ascomycetes). When the spores are cut off externally from the gonotoconts, they are called basidiospores and the sporogenous cells basidia (whence the term Basidiomycetes, the saprophytic mushrooms, puffballs, etc., and the parasitic rusts and smuts). As the conidiophores of the haplont, these gonotoconts are usually grouped in a superficial layer of tissue, known as a h3nnenium. This tissue generally OENERAL MORPHOLOGY OF FUNGI 21 develops on a fructification whose structure is highly differentiated. In the higher groups, the fructifications containing the gonotoconts become visible to the naked eye and are called the perfect forms. The other spore forms are referred to as imperfect or secondary spore forms. Sexual Organs and Sexuality. — The sexual function in its simplest stage involves but two processes : (a) fertilization, a fusion of two nuclei, recurring periodically in the course of development, initiating specific stimuli for fur- ther development; and (b) meiosis, a return to the single chromosome num- ber. This rotation (haplont -^ fertilization — > diplont -^ meiosis — >) comprises the changes of nuclear condition in most organisms. A few fungi seem to exist without such a change; they seem to live an unlimited number of "gen- erations" without reconstruction of their nuclei and to propagate themselves Fig-. 3. — Coremium. Hemispora coremiformis. (After M. Moore 1935.) by imperfect stages only. Such fungi with incomplete, or incompletely known life cycles are called Fungi Imperfecti. The fungi with complete life cycles are divided into homothallic (her- maphroditic) and heterothallic forms. In contrast to the higher plants and animals, the division of sexes is here limited to the haplont, i.e., the mycelium. The homothallic form is often indicated by the symbol ±, the heterothallic forms as + or -. The + and - mycelia of the latter group may be distinguished from each other only physiologically, as indicated by their sexual tendencies, or morphologically as well (e.g., in growth form, sexual dimorphism). Fertilization in fungi, as in protozoa, may occur in many ways. The simplest normal type of fertilization, when two spatially separated, not closely related sexual cells fuse to form a new entity, is called amphimixis. If the sexual cells arise as daughter cells of a differentiated mother cell and are themselves characteristically formed, they are called gametes and 22 • MEDICAL MYCOLOGY the mother cells gametangia. The copulation of two gametes of this type is called merogamy. If the gametes appear wholly of the same size and shape, the copulation is isogamous; if the gametes are differentiated, their copulation is heterogamous (anisogamous). When the female gamete is a large and non- motile egg and the male gamete is a small and motile sperm, their copulation is oogamous, a condition confined to primitive fungi, although present in most higher animals and primitive plant phyla. The more advanced fungi show various stages in the degeneration of merogamy. At a comparatively low stage, the differentiation of gametes is suppressed and the contents of the gametangia remain polyenergid. Thus the original copulation of gametes disappears, being replaced by many sec- ondary processes which compensate for the loss of the original merogamy. All these secondary processes are classified as deuterogamy. The higher algae and flowering plants have also developed these processes, while the primitive merogamy has persisted through the highest vertebrates. In deuterogamy, the gametangia assume the function of their daughter cells, the gametes, and their coenocytic content fuses without further differ- entiation. The sexual act occurs between two sexual organs instead of between two sexual cells and sexual attraction passes from the latter to the former. This type of copulation is called gametangial. It assumes close contact be- tween two gametangia and has a biologically obvious consequence that one gametangium can fertilize only one other which must be located nearby ; it has the advantage that the fusion of gametes is no longer fortuitous, since the gametangia provide that their nuclei can come into contact and fuse. Thus gametangial copulation is an efficient type, since most of the nuclei of both gametangia succeed in their activity with an increase in the number of zygote nuclei. The fate of the gametangium hangs upon the occurrence or nonoc- currence of a single sexual act, from which results one single, very strong, coenocytic zygote instead of many smaller unicellular zygotes. Also a single mature gametangium no longer depends on a definite medium (e.g., water) for the copulation of its gametes, a condition which has facilitated the transi- tion from water to land habitats and to parasitism. Whether gametangia are borne on specialized branches or whether hyphal branches as a whole complete the act of fertilization, they are called copulation branches. When sexual dimorphism is present, the male is called an antheridium and the female an ascogonium. Among the higher Ascomycetes, the fundaments of the antheridium are gradually reduced, whereby cross-fertilization generally ceases and is replaced by self-fertilization, i.e., a new group of deuterogamous processes, between daughter cells of the same mother cell or between nuclei of the same cell, which are included in the term automixis. Automixis is represented in fungi by two forms : parthenogamy and autogamy. Parthenogamy is fertilization which occurs between two female cells, i.e., in fungi usually between two cells of the ascogonium. In some groups this parthenogamic fusion of two specialized cells is replaced by the GENERAL MORPHOLOGY OF FUNGI 23 pairing of nuclei within a single multinucleate cell of the ascogonium. This automictic fertilization within a cell is called autogamy. From these forms in which the sexual organs (or in any case the female organ) are apparently typical in form but no longer functional and serving only as a site of automictic processes, there is a series of intermediate stages to the other extreme where the sexual organs disappear entirely, the sexual processes occurring in the mycelium between any two sexually differentiated cells. The latter process is called pseudomixis. Since the copulating cells are not morphologically distinguishable from other vegetative cells and since only the release of specific developmental stimuli marks this anastomosis of two vegetative cells as a sexual process, pseudogamy is often difficult to dis- tinguish from the usual pseudosexual anastomoses which are brought about by food relations. Its true character is recognizable cytologically only in the pairing of nuclei. If pseudogamy occurs between two sprout cells, tliej^ are sometimes wrongly called gametes. In order to avoid misunderstanding, this term should be reserved for merogamous gametes. The ambiguous term pedogfamy, often employed in other senses, should be used to indicate pseu- dogamy between adult and young cells. The special case of pseudogamy between mother and daughter cell is called adelphogamy. Apomixis, the entire loss of fertilization, represents the last step in this series of reduction of sexuality where growth from reproductive cells occurs vegetatively without cell or nuclear fusion, or any external stimulus of de- velopment. If the new individual (in the absence of fertilization) arises from haploid sexual cells, the process is called parthenog'enesis ; if they arise (in the absence of meiosis) from diploid sexual cells, the process is called apogamy. In the study of fungi, there is the further difficulty that the original processes of fertilization are replaced by all sorts of substitutes. Among the lower fungi, there is simple fertilization when a fusion of the cytoplasm of two sexual cells (plasmogamy) is immediately followed by a fusion of both haploid nuclei into a diploid zygote nucleus, a syncaryon (caryog-amy). In most fungi, however, caryogamy is delayed and is only completed just before meiosis. Thus the sexual haploid nuclei, while remaining spatially separate, unite only to form a dicaryon, where the paired nuclei divide synchronously (conjugately) while retaining the same ability to activate somatic develop- ment as after complete caryog*amy. This phenomenon is analogous to that in Cyclops, in which the parent chromosomes remain distinct up to the time of egg formation (synapsis) although they are surrounded by the same nuclear membrane, whereas in the fungi they remain Avithin their original nuclear membranes. In this retardation of caryogamy, the binucleate "zygote" continues its growth without completing nuclear fusion, developing a new mycelium whose cells, morphologically virtually diploid, contain two sexually differentiated haploid nuclei. This new phase, intruded between plasmogamy and caryog- amy, is called the binucleate phase. In the Ascomycetes, this phase is mostly limited to the ascogenous hyphae and the hymenium of the fructification, but in the Basidiomycetes, it is usually the most conspicuous phase of the life cycle. 24 MEDICAL MYCOLOGY In spite of this removal and retardation, caryogamy always occurs in definite organs. The organs in which the fertilization processes are completed and the dicaryon ends are called zeugites. As caryogamy is delayed until the necessity for meiosis appears, the zeugites also frequently function as gonotoconts. These two processes, the transformation of the cells which com- plete the sexual act and the division of the sexual act itself into plasmogamy and caryogamy, separated in time and space, are both fundamental and very useful in the study of phylogeny and classification. In the present state of our knowledge, the various groups of fungi are apparently polyphyletic and unrelated. Some members seem more closely related to other groups of plants than to other groups of fungi; e.g., the Chlamydobacteriales and perhaps the Thiobacteriales seem more closely re- lated to the Myxophyceae than they are to other groups of bacteria or of fungi. The following list of classes of fungi illustrates the main subdivisions while not necessarily assuming that all the fungi of these larger groups are monophyletic. In time some of these groups will probably be further divided, e.g., the Phycomycetes seem quite heterogeneous. For our purposes, how- ever, only one order of these, the Mucorales, has been shown to have members attacking man and other mammals, and needs consideration here. Schizomycetes : bacteria in the larger sense. Myxomycetes: slime molds, having many resemblances to Protozoa in some stages of their life cycle. Phycomycetes: coenocytic fungi of varied origin and relationship. Ascomycetes : a large poljonorphic group with a common method of spore formation in asci. Basidiomycetes : a very large group with spores borne on a specialized gonotocont, the basidium, furnishing most of the conspicuous fungi. Fungi Imperfecti : a heterogeneous group whose life cycles are unknown in full, or which have degenerated until sexuality has been lost, provisionally grouped together until they are better known. Schizomycetes. — Bacteria. While this group is by far the most important from the standpoint of the physician and the surgeon, it is also the best known to the medical profession and will not be given further consideration here. The bacteria seem almost wholly unrelated to the other groups of fungi, and some of the higher forms are suggestive of Myxophyceae which have lost their chlorophyll. Myxomycetes. — The slime molds are a very interesting group with many stages suggestive of similar conditions found in the Protozoa, especially the Rhizopoda, while other stages are analogous to those in other groups of fungi. The stages commonly observed being plantlike, they have been studied mostly by botanists and only in the last decade has any long continued or thorough attempt been made to follow the life cycle in detail or to study the cytology. At present the group is known to cause plant diseases but has not been suggested in relation to animal disease. Any related organisms attacking man have undoubtedly been studied and classified as Protozoa. GENERAL MORPHOLOGY OP FUNGI 25 Phycomycetes. — This group in its more primitive members seems distantly related to the Flagellata of the Protozoa. Most of the members have a vegeta- tive body of mycelium and in all but one order reproduce by a flagellated body. Most of the group are saprophytes or parasites on plants except a few facultative parasites of fish or invertebrates. The only group at present known to produce human parasites is the Mucorales which will be treated later in the appropriate chapter. Ascomycetes. — The more primitive members show distant relationships to the higher Phycomycetes on the one hand and to the Ehodophyceae (Red Algae) on the other. After a half century of bitter controversy on this ques- tion, there is still little agreement. Here the vegetative body is a typically uninucleate, septate mycelium and sexual reproduction occurs by means of cells produced within an ascus, following meiosis in all forms where this has been carefully studied. Of the twenty orders into which this group is usually divided, members of only two, the most aberrant and primitive (or degener- ate), have been shown to cause human disease. The bulk of the group are saprophytes or parasites of the leaves and bark of plants, while the most highly specialized (Laboulbeniales) are at present known only as parasites of living insects. Basidiomycetes. — While the life cycle shows a certain parallelism to that of the higher Phycomycetes and the Ascomycetes, it is difficult to assign to these any very close relationship with other groups. The vegetative body again consists of uninucleate or binucleate mycelium, forming a conspicuous fructification on which the reproductive structures are borne. Reproduction occurs by means of basidiospores. So far as is known the group is saprophytic on decaying vegetable matter in the conspicuous species, such as the mush- rooms, punks, puffballs, etc., or parasitic, usually on leaves, in the Uredinales (rusts) and Ustilaginales (smuts). Fungi Imperfecti. — These are a large group, artificially classified together while we await more knowledge of their life history. The greater part, whose life history has been discovered, has been found to be Ascomycetes, but it is never safe to predict that several members of a given genus of Fungi Im- perfecti will necessarily belong to the same genus of Ascomycetes or even will be Ascomycetes; e.g., when Oedocephalum was carefully studied, one species was found to be a Phycomyeete, another an Ascomycete, and a third a Basidiomycete. Various subdivisions have been proposed, but none has proved altogether satisfactory. Further considerations may well be delayed to a subsequent chapter (p. 665). In the following chapters only those orders will be discussed in which mammalian pathogens have been found, and the reader is referred to Gau- mann and Dodge (1928) for information on other groups. BIBLIOGRAPHY The following bibliography includes references to early work in which it is difficult or impossible to be sure of the group of fungi involved, also gen- 26 MEDICAL MYCOLOGY eral review articles, textbooks, etc., covering most of the groups of human pathogenic fungi. These have not been repeated in the bibliographies of the individual groups, even though they often contain as much information about a single group as do the references given for that group. Asterisks (*) de- note that I have not read the reference in question, but it is quoted from a fairly reliable author or abstracting journal. I have not tried to quote any references from certain writers whose references are notoriously incorrect, often showing errors in over half the titles of a short bibliography. Agostini, Angela. 1934. Miceti della Somalia, Atti 1st. Bot. B. Univ. Pavia TV, 4: 191-201, 4 fig. Bail. 1867. Ueber kraukheiteneizeugende Pilze, Wiener Med. Woch. 17: col. 977-979; 993- 996. Baillon, H. 1889. Traite de botanique medicale cryptogainique, Paris, 376 pp. [see pp. 246- 249]. Baumgarten, Paul Clemens. 1884. Ueber patliogene pflanzliclie Mikroorganismen. I, Deutsch. Med. Zeitschr. 1884i: 149, 150; 163-165; 175-178; II. Die pathogenen Schizomyceten, Ibid. 18842: 169-173; 181-185; 193-197; 205-208. — . 1890. Lehrbuch der pathologischen Mykologie. Vorlesungen fiir Arzte und Studierende. Harald Brulm; Braunschweig, IX, 973 pp., 1 pi, 101 figs. Beurmann, Lucien de. 1912. Les nouvelles mycoses, exoascoses (ex-blastomy coses) oidio- mycoses, sporotrichoses, botrytimycose, oosporose, hemisporose, Paris, Masson & Cie. 165 pp. Beurmann, L. de & H. Gougorot. 1909. Les Exascoses, endomycoses, et parendomycoses (Mu- guet) saccharomycoses (mycose de Busse-Buschke) et parasaccliaromycoses. Zymone- matoses (Mycose de Gilchrist) Envision et demembrement de I'ancien groupe des Blastomycoses, Bull. Mem. Soc. Med. Hop. Paris III, 28: 222-246; 250-263. — . 1912. L'etat actuel de la question des mycoses, Biol. Med. 10: 133-166; 187-222, 18 figs. Bianchini, Giuseppe & G. Paolo Manfrini. 1924. La micologia del cadavere umano nei rispetti della cronologia della morte e delle trasformazioni tanatologiche, Atti B. Accad. Fisiocrit. Siena IX, 16: 335-351, S figs. Blanchard, Raphael. 1896. Parasites vegetaux a 1 'exclusion des bacteries, in Bouchard, Charles. Traite de pathologic generale 2: 811-926. — . 1899. Sur une affection causee par les spores d'un champignon parasite du roseau ou canne de Provence (Arundo donax L.), Arch, de Parasitol. 1: 503-512. Bodin, E. 1902. Les champignons parasites de I'homme, Paris, Masson et Cie, 208 pp. Brocq Eousseu. 1921. Les recherches mycologiques en medecine veterinaire, Bull. Soc. Myc. France 37: 99-103. Brooke, Gilbert E. 1908. Aids to tropical medicine. New York, William E. Wood, 163 pp. Brumpt, E. 1927. Precis de parasitologic. 4 ed. Paris, Masson & Cie, viii, 1452 pp., 5 pis. Castellani, Aldo. 1912. Note on the importance of Hyphomycetes and other fungi in tropical pathology, Brit. Med. Jour. 2: 1208-1212. — . 1917. Notes on tropical diseases met with in the Balcanic and Adriatic zones, Jov/r. Trop. Med. Hyg. 20: 157-164; 170-174; 181-186; 198-202; 209-214; 219-223. — . 1918. Alcune osservazioni sulla malaria e su altre malattie tropicali della zona Balcanico-Adriatico, Ann. Med. Nav. 24i: 169-213. — . 1920. Higher fungi and human pathology, Lancet, 198: 847-852, 895-901, 943-946 [also reprinted in Jo^tr. Trop. Med. Hyg. 23: 101-110, 117-125, 132-138]. — . 1923. Medical mycology, Brit. Med. Jour. 2: 937-1041 [also reprinted Jour. Trop. Med. Hyg. 27: 49-53, 61-65, 1924]. GENERAL MORPHOLOGY OF FUNGI 27 — . 1924. Tropical Dermatology, Int. Conf. Health Problems of Trop. Amer. [United Fruit Co.] 485-499. — . 1925. Observations on some diseases of Central America, Jour. Trop. Med. Hyg. 28: 1-14, 31 figs. — . 1926. Chronic bronchites with hemorrhagic sputum of nontubercular origin, N. Orleans Med. Surg. Jour. 79: 20-42. — . 1927. Notes on certain bronchomycoses which may simulate pulmonary tuberculosis, Am. Bev. T^tiberculosis 16: 541-574. — . 1927-1928. Fungi and fungous diseases. Arch. Derm. Syphilol. 16: 383-425; 571-604; 714-740, 1927; 17: 61-97; 194-220; 354-379; 1928 [reprinted Chicago, Amer. Med. Assn. 203 pp., 44 figs. 1928]. Castellani, Aldo & Albert J. Chalmers. 1910-1919. Manual of tropical medicine, New York, WilUam Wood & Co., XXV + 1243 pp.; ed. 2, XXXII -f 1747 pp., 16 pis., 909 figs., 1913; ed. 3, X -F 2436 pp., 16 pis., 909 figs., 1919. Castellani, Aldo, MacKenzie Douglas, & J. Thomson. 1923. Further observations on tonsil- lomycoses, Jou,r. Trop. Med. Hyg. 26: 19-24. Castellani, Aldo, & Henri Tejera. 1923. Notes on the etiology of ''cute," Jour. Trop. Med. Hyg. 26: 183, 184. Cavara, V. 1928. Le micosi oculari, Siena, 494 pp. Chiurco, Giorgio Alberto. 1922. Simbiosi tra ifomieeti patogeni e schizomiceti, Biv. Biol. 4: 684-688. * — . 1922. Simbiosi tra ifomieeti patogeni e schizomiceti, in rapporto al loro potere patogeno, 1st. Bot. B. Univ. Siena, 75 pp. — . 1923. Micosi chirurgiche sperimentali, Pathologica. 15: 487, 488. — . 1925. Esperience nelle cavita articolari tessuto muscolaree cutaneo dei ratti e delle cavie in rapporto alia simbiosi ifoschizomicetica, II, Atti B. Accad. Fisiocrit. Siena IX, 16: 171-207, Pis. 1, 2. — . 1927. Experimentell hervorgerufene mykotische Arthritiden und Myositiden, Derm. Woch. 84: 228-232. Coupin, Henri. 1909. Atlas des champignons pathogenes de I'homme et des animaux. 58 planches renfermant 1000 dessins reproduits d'apres les travaux originaux, Paris, Octave Doin et Fils. Dubreuilh, William. 1891. Des moisissures parasitaires de I'homrae et des animaux superieurs, Arch. Med. Exp. Anat. Path. 3: 428-447, 566-592. Durante, G. 1927. Les mycoses meconnues. Bull. Mem. Soc. Med. Hop. Paris [51] : 1513- 1516. Fairman, Ch. E. 1920. The ascomycetous fungi of human excreta, Lyndonville, N. Y., 11 pp., S figs, 1 table. Fleisher, M. S., So M. Wachowiak. 1924. The relation of fungi imperfecti to diarrhoeal con- ditions. Am. Jour. Med. Sci. 168: 371-380. Fonseca filho, Olympio Oliveira Eibeiro da. 1928. Algumas consideragoes de ordem geral sobre as dermatomycoses, Sciencia Med. 6: 565-569. — . 1929. As mycoses na clinica dermatologica e syphilographica da faculdade de Medicina do Eio de Janeiro, Bev. Med. Cirurg. Brasil. 37: 126-128. — . 1929. Notas sobre os exames de laboratorio na pesquiza e diagnostico das mycoses, Bev. Med. Cirurg. Brasil. 37: 169-179. Frescoln, L. D. 1916. Mycology as a part of practical dermatology, Internat. Clin. 2: 170- 172, 1 col. pi. Frey, Ludwig. 1888. II. Fungus articulat. talo-calcaneae. Injection con Jodoformather Heilung und voUkommene Wiederherstellung der Functionsfahigkeit, Wiener Med. Presse. 29: col. 1325, 1326. Furbringer, Paul. 1876. Beobachtungen iiber Lungenmykose beim Menschen, Arch. Path. Anat. Physiol. [Virchow] 66: 330-365, PI. 15. 28 MEDICAL MYCOLOGY Giaumann, Ernst A., & Carroll William Dodge. 1928. Comparative morphology of fungi, New York, McGraw Hill Book Co., 701 pp., 398 figs. Gautier, L. 1907. Eecherches biologiques sur quelques champignons parasites de I'homme el des animaux (diastases & toxines), Brest, P. Gadreau, 149 pp. Gedoelst, L. 1902. Les champignons parasites de I'homme et des animaux domestiques, Bruxelles, 199 pp., 1£4 figs. Grazia, Francesco de. 1903. I microorganismi dei polmoni dei Cardiaci, Bif. Med. 19: 709- 713. Greco, Nicolas V. 1916. Origine des tumeurs (etiologie du cancre) et observations de mycoses (blastomycoses, etc.) Argentines, La Semana Medica: Buenos Ayres. VI + 853 pp., 29 pis., 492 figs. Greenwood, Arthur M. & Joseph H. Swartz. 1927. [Fungous diseases of the skin] a review of the literature [of the last two years]. Arch. Derm. Syphilol. 15: 404-414. Greig, E. D. W. & G. C. Maitra. 1918. Observations on the occurrence of hyphomycetic elements m ulcers of the skin in 19 cases, Indian Jour. Med. Res. 5: 481-490, 1 fig. Grohe. 1870. [Experimente . , . iiber die Injection von Pilzsporen], Berlin. Klin. Woch. 7: 8, 9. Griitz, O. 1927. Die Hyphenpilze oder Eumyceten, Hand. Path. Mikroorg. [Kolle So Wasser- mann, ed. 3] 5: 133-320, 10 pis., 66 figs. Gueguen, Fernand. 1904. Les champignons parasites de I'homme et des animaux. Generalites, classification, biologie, teclinique. 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Precis de pathologie exotique (maladies des pays chauds et des pays froids), ed. 4, Paris, Octave Doin, v. 1, 942 pp.; v. 2, 1232 pp. *L6wenberg. 1880. Des champignons parasites de I'oreille humaine, Paris. GENERAL MORPHOLOGY OF FUNGI 29 Magalhaes, Octavio de So Aroeira Neves. 1926. Ensaios de mycologia. Contribuigao para o estudo dos cogumelos em Bello Horizonte, Janeiro 1912-Juiiho 1920, Mem. Inst. Oswaldo Cruz 19: 245-283, Pis. 56-91 [translated: Essays on mycology, contribution to the study of fungi in Bello Horizonte. Jan. 1912- June 1920, Ibid. 19: 285-322, 1926]. Maillard, J. P. 1933. Les mycoses du conduit auditif externe, These Fac. Med. Univ. Paris 537: 1-73. Marchoux, E. 1922. Mycose pulmonaire, BiM. Soc. Path. Exot. 15: 919-920. Mattlet, J. 1926. Mycoses dans I'Urundi, Ann. Soc. Beige Med. Trop. 6: 1-41, 18 figs. Mayer, A. C. 1815. Verchimmelung (Mucedo) in lebenden Korper, Deutsch. Arch. Physiol. [Meckel's] 1: 310-312. Mendelson, Ealph W. 1921. Tropical bronchopulmonary mycosis, Jour. Am. Med. Assn. 77: 110-112, 4 figs. — . 1925. Some interesting diseases observed in the clinic of the Bangkok Central Hospital, Jour. Trop. Med. Hyg. 28: 393-405, SI figs. Mohler, J. 1904. Mycotic stomatites of cattle, U. S. Dept. Agr. Burewu Animal Industry, Circ. 51: 1-6. Nannizzi, Arturo. 1934. Repertorio sistematico dei miceti dell'uomo e degli animali, Siena, S. A. Poligrafica Meini xii + 557 pp., 224 figs. [Received in 1935 after this book was iA galley proof, too late to be considered except for changes of nomenclature of species already included.] Neumann, L. G. 1892. Traite des maladies parasitaires non microbiennes des animaux domestiques, ed. 2, Paris, XVI + 768 pp., 364 figs. Neveu-Lemaire, Maurice. 1911. Parasitologie des animaux domestiques. Maladies para- sitaires non bacteriennes, Paris, J. Lamarre & Cie, 1257 pp. — . 1921. Precis de parasitologie humaine, ed. 5, Paris, J. Lamarre, VI -l- 466 pp. Niethe, Ulrich. 1926. Die Bedeutung der gewohnliche Schimmelpilze fiir die menschliehe Haut, Arch. Derm. Syphilis 152: 358-364. Ota, Masao. 1928. Champignons parasites de I'homme (Etudes morphologiques et sys- tematiques), Jap. Jour. Derm. Urol. 28: [202 pp. in Japanese, 7 pp. of French summary, 5 pp. of bibliography]. Perazzi, Piero. 1926. T miceti dimoranti nello- regione genitale della donna, Atti B. Accad. Fisiocrit. Siena 17: 361-410, Pis. 1-S. Perin, Arrigo. 1925. Le micosi pulmonari e generalita sui miceti patogeni. Siena, Libreria Editrice Senese, viii -|- 294 pp. Pinoy, E. 1903. Les champignons pathogenes; leur classification d'apres les caracteres botaniques. Bull. Inst. Pasteur. 1: 761-774. Plant, Hugo C. 1903, 1912. Die Hyphenpilze oder Eumyceten, Handb. path. Mihroorganis- men [Kolle & Wassermann] 1: 526-660; ed. 2, 5: 1-154, 7 pis. Eamsbottom, John, & Arthur Whitfield. 1931. Fungi pathogenic to man, Syst. Bact. in Relation to Medicine [Med. Res. Council] 8: 11-70. Raymond, Victor & Jacques Parisot. 1916. £tiologie, prophylaxie et therapeutique de I'afEection dite gelure des pieds, C. B. Acad. Sci. 162: 694-696. RedaeUi, P. 1931. Tecnica micologica medica, Bologna, L. Capelli. 227 pp., 58 figs. Robin, Charles. 1847. Les vegetaux qui croissent sur les animaux vivants, These Fac. Sci. Paris, VIII + 120 pp., 3 pis. — . 1853. Historie naturelle des vegetaux parasites qui croissent sur I'homme et sur les animaux vivants, Paris, x + 704 pp. Atlas 15 pis. Rockwell, George E. & John H. Highberger. 1927. The necessity of carbon dioxide for the growth of bacteria, yeasts, and moulds. Jour. Inf. Dis. 27: 438-446. Rockwood, Ethel M, 1930. A study of fungus infected nails. Arch. Derm. Syphilol. 22:. 395-400, 7 figs. Eoeckl, G. 1884. tJber Pneumonomykose. Deutsch. Zeitsrhr. Thiermed. Vergleich. Path. 10: 122-132, PI. 12. 30 MEDICAL MYCOLOGY Eouyer, E. & J. Pellissier. 1915. Contribution a 1 'etude de certaines mycoses de blessures de guerre, Ann. Inst. Pasteur 29: 551-555. Eudolph, Max. 1914. tJber die brasilianische Figueira, Arch. Schiffs-Tropenhyff. 18: 498. Sartory, A. 1920-1923. Champignons parasites de I'homme et des animaux, 14 pts. 1 suppl. 1-86, 1920; 91-484, 1921; 487-721, 1922; 727-895 and index 1-47, 1923; suppl. 1-78, 1923. Schmidt, Hans. 1921. ijber Beziehung zvvischen Tuberkelbazillen und Schimmelpilzen, Ztschr. Klin. Tu'berlculose. 46: 456-459. Schmidt, P. W. 1933. Dermatomykosen, Derm. Zeitschr. 66: 241-261. See, Pierre. 1922. Les mycoses. Gas. Sop. Civ. Milit. 95: 469-476. Senator, H. 1891. [Pneumaturie und Diabetes], Intern. Beitr. Wiss. Med. 3: -319-332 [also cited Festschrift Kudolf Virchow]. Shear, Cornelius Lett. 1927. Mycology in relation to human pathology. Am. Nat. 61: 151-159. Siebenmann, Friedrich. 1889. Die Schimmelmykosen des mensclilichen Ohres. ed. 2, Wies- baden, 118 pp., 4 pis. Sorokin, N. 1882. Rastytel'nye parazity cheloveka i zhivotnykh kak prichina zaraznykh boleznei dlia naturalistov, vrachei, studentov i veterinarov. 4 vol. and atlas. S. Peterburg. Souls, Ferdinand Xavier Felix. 1891. Contribution a I'etude des otomycoses, These Fac. Med. Pharm. Univ. Bordeaux 9: 1-48, 2 -figs. Stein, E. O. 1914. Die Fadenpilzerkrankungen der Menschen. Miinchen, 1914, 99 pp., S2 pis. [also cited Lehmanns Med. Atlanten. 12: 1-99, 3^ pis., 1914]; ed. 2, 128 pp., 32 pis., 1930. Strong, Eichard P. 1930. The African republic of Liberia and the Belgian Congo. Based on observations made and material collected during the Harvard African expedition, 1926-1927. Cambridge [Mass.], Harvard University Press, v. 1, xxvi + 568 pp., 6 pis. Strong, Eichard P. & George C. Shattuck. 1930. Medical and pathological investigations in Liberia and the Belgian Congo, Contr. Dept. Trap. Med., Inst. Trop. Biol. & Med. 55: 210-461. Talice, Eodolfo V. 1930. Le concept actuel des mycoses medicales de I'appareil respiratoire, Bev. Stid-Amer. Med. Chir. 1: 181-188. Thibierge, G. 1925. Les enseignements dermatologiques de la guerre 1914-1918, Ann. Derm. Syphiligr. 6: 481-514; 641-661; 705-724. [III. Mycoses cutanees, pp. 641-653.] Uhthoff, W. 1883. Beitrage zur pathologischen Anatomic des Auges. III. Partielle Necrose der mensclilichen Hornhaut durch Einwanderung von Schimmelpilzen, Arch. Ophthal- mol. 29: 3: 178-181. Virchow, Eudolf. 1856. Beitrage zur Lehre von den beim Menschen Vorkommenden pflanz- lichen Parasiten, Arch. Path. Anat. Physiol. [Virchow 's] 9: 557-593. Vuillemin, Paul. 1931. Les champignons parasites et les mycoses de I'homme. Paris, Paul Lechevalier & Fils, 291 pp. Weidman, Fred D. 1929. Light from the botanic field on medical mycologic problems, Arch. Derm. Syphilol. 19: 867-877. Wright, James Homer. 1913. Non-bacterial fungus infections — the mycoses, in Osier, William & Thomas McCrae, Modern medicine in theory and practice — ed. 2, 1: 1033-1060. Xavier, A. A. 1926. Apontamentos sobre micologia (Notas de um curso), Sciencia Med. (Eio de Janeiro) 4: 60-68. Young, James. 1921. Organism cultivated from carcinomata, Brit. Med. Jour. 1: 933. Ziirn, Friedrich Anton & Hugo C. Plaut. 1887-1889. II. Die pflanzliche Parasiten in Ziirn, F. A., Die Schmarotzer auf und in dem Korper unserer Haussaugetiere sowie die durch erstere veranlassten Krankheiten, deren Behandlung und Verhiitung. Weimar, 837 pp., 4 pis. CHAPTER II PHYSIOLOGY OF FUNGI WITH SPECIAL REFERENCE TO REPRODUCTION The life of a fungus may be divided into three periods: (1) vegetative growth, (2) multiplication by asexual means, and (3) the development of sex- ual organs usually resulting in a spore able to resist unfavorable factors of the environment. Since studies of classification and evolution of the various groups of fungi are usually based on this last stage which presents the great- est diversity of phenomena, much of laboratory work in the last half century has been directed toward securing the production of sexual stages. After Bary, Brefeld, Hansen and Zopf had developed the technic of cultivation, Klebs in the years from 1896 to 1904 laid the foundations of much of our pres- ent knowledge by detailed studies of the conditions necessary for vegetative growth and reproduction in a few organisms. Subsequent work of C. H. Kauffman and his students has done much to extend Klebs' generalizations to further groups of fungi and to confirm his well-known dicta. These may be summarized as follows : 1. Among all organisms, growth and reproduction are life processes, Avhich depend upon different conditions : among lower organisms, probably environment determines whether growth or reproduction will occur. 2. As long as the environment is favorable for growth, reproduction will not occur. An environment favoring the latter is usually more or less un- favorable to further growth. 3. The limits of each factor of the environment are narrower for reproduc- tion than for growth. Therefore growth can still take place, although repro- duction is limited by a too weak or too strong influence of some factor. 4. Before reproduction can occur growth must have been sufficient to have stored the products necessary for the reproductive processes. As a consequence of these dicta, most specialized methods for securing sexual or even asexual reproduction in a given group consist essentially in growing the organism for a time as nearly at optimum conditions as possible, then suddenly changing one or more factors of the environment to a condition less favorable for growth but still within the limit for reproduction. Exam- ples of this will be given in the following analysis of some of the chief factors of the environment. Water. — The water requirement and transpiration have been little studied in fungi. While spores often have mechanisms for resisting desiccation over long periods and some very complex specialized structures are developed among the higher fungi for preventing the reproductive organs from drying out, in general the fungi both grow and reproduce in relatively high humidi- 31 32 MEDICAL MYCOLOGY ties. The optimum humidity seems very closely related to oxygen supply and respiration. Many of the lower fungi grow well, or even normally are found, immersed in water. Many others will form a thick hyphal mat over the sur- face of a liquid culture medium, although in this case much more oxygen is required for normal development than in purely aquatic fungi and the repro- ductive structures are usually produced in the aerial mycelium. Still other groups will grow well and reproduce only on solid substrata, and a few grow best in comparatively low humidities. Growth is usually much slower at low humidities.* Some species have a narrow range while others will tolerate a very wide range. Transpiration rates and humidity are often important in initiating repro- duction. Very frequently reproductive structures are produced only when vegetative growth is severely checked by the drying out of the media. This, however, is usually associated with partial exhaustion of nutrients and the accumulation of toxic products of metabolism (staling) so that it is often difficult to evaluate the influence of the various factors. (Klebs 1896, Coons 1916.) Nutrition. — The most emphasis has been placed upon this factor and a great body of literature on culture media and their effects has been produced. The complexity of the vast majority in use does not lend itself to an analysis of the separate factors involved. These may be considered under the head- ings of inorganic salt requirements, carbon and nitrogen supply, and relations to concentration of hydrogen ions, although these are closely interdependent and also related to the physical factors of the environment. Inorganic Salts. — The requirement of these seems much the same as for the higher plants, traces of potassium, magnesium, iron, and calcium being necessary to secure good growth and reproduction. Calcium, however, is much less important for fungi than for the flowering plants. Of the nonmetal- lic elements, phosphorus, sulphur, carbon, and nitrogen are important. Other elements are often present and may influence growth and reproduction, but they are relatively unimportant in comparison with the above (e.g., McHargue & Calfee 1931). Phosphorus is usually supplied as a phosphate, where it is often useful in regulating the concentration of hydrogen ions in the solution. In many media it is supplied as nucleoproteins, nucleic acid, and lecithin, and seems equally available in these forms. Sulphur. — This element is usually supplied as a sulphate, but is also utilized from sulphocyanates, thiosulphates and the sulphur-containing amino acids and their compounds. The literature on this subject was well sum- marized by Armstrong (1921) and will not be covered here. In various or- ganic media probably a large portion of the necessary sulphur is available from the proteins since practically all of them contain some cystine or cysteine. Where large amounts of sulphur are available, particles of sulphur may be ♦This fact may b© utilized in stock cultures by transferring to a 4-5% agar instead of l%-3% commonly employed. Since growth is slow and, water loss is slower from the harder agar, one safely can allow a much longer time to elapse between transfers. PHYSIOLOGY OF FUNGI 33 deposited in the cells or hydrogen sulphide may be produced, although these phenomena are rare under ordinary cultural conditions. Carbon. — The possible sources of carbon are almost infinite, since there are many possible combinations from which it may be utilized in dilute solu- tion. For practical purposes the carbohydrates and organic acids are em- ployed most. Of these, glucose has the widest use although the other hexoses, especially mannose and fructose, are easily utilized, and also sucrose (cane sugar) by organisms which secrete invertase. The other sugars and aldehydes, celluloses, starches, etc., are variable in effect, those with low molecular weights generallj^ being toxic. The lower alcohols are also generally toxic, although some of higher molecular weight, such as glycerol (glycerin) and mannitol (mannite), are very readily assimilated. In general, substances hav- ing a three- or six-membered carbon chain are the most assimilable. Among the organic acids, the amino acids and those of the aliphatic series having a low hydrogen ion concentration, are readily utilized. Acids of the cyclic (benzene) and heterocyclic series are rarely metabolized unless there is a side chain of at least three carbon atoms, as in the amino acids, phenylalanine, tyrosine, and tryptophane. There are also scattered observations on utiliza- tion of depsides, tannins, alkaloids, amines, etc. The proteins and their de- composition products often furnish a suitable carbon source, although they are primarily regarded as a nitrogen source. In many of the compounds assimilability depends upon the ability of the organism in question to secrete some enzyme which will hydrolyze or so break down the higher compound that one or more of the resulting products will be utilizable while none will be toxic, e.g., in depsides and tannins, usually the hexose is utilized and the galloyl compounds are not attacked. Many of the older authors attempted to arrange carbon compounds in the order of decreasing availability, but this is rather futile, as the exact order varies Avith the organism in question and with other environmental factors. (Pasteur 1860, 1862 ; Naegeli 1879, 1882 ; Reinke 1882 ; Duclaux 1885, 1889 ; Linossier & Roux 1890; Went 1901; Ekman 1911. For action of cyclic carbon compounds, see Waterman 1913, Bokorny 1920.) Nitrogen. — While from time to time atmospheric nitrogen has been indi- cated as a source of nitrogen for fungi, the inadequacy of methods of deter- mination of total nitrogen has tended to discredit this source for most fungi. Duggar and Davis (1916) have carefully summarized the literature and dis- cussed the methods employed. Klotz (1923) has adequately summarized the literature for other sources. Older authors attempted to group fungi as nitrate, ammonium, amino acid, peptone or protein organisms, but in view of the large number of variables involved and the difficulties of adopting a suit- able criterion for growth these groupings are quite inadequate. While some fungi are capable of utilizing nitrates or ammonium salts, in the groups in which we are most interested here, they are so few that they need not be con- sidered. Czapek (1902) made a very extensive study of substances as nitrogen 34 MEDICAL MYCOLOGY sources and concluded that amino acids and compounds nearly resembling them, are most fully and easily utilized, since the principal use of nitrogen in nutrition is in the production of proteins. Lutz (1905) extended and con- firmed these observations. Ritter (1909, 1912, 1914) studied the effect of the acid produced when ammonium salts were used. Kossowicz (1912, 1914) ex- tended his studies to purines and related compounds. Brenner (1911) extended his observations to include many compounds which proved toxic. While these and many other later papers have much interest for the prob- lem of the mechanism of protein synthesis in fungi, they have little direct bearing on the problems of cultivation. Few studies have been made on the vitamin requirements of fungi, and the evidence is not very conclusive at present. (See excellent review of litera- ture by Sergent 1928, Peskett 1933.) However, the possibility of such fac- tors should be borne in mind in nutritional studies (see Freedman & Funk 1922; Robertson & Davis 1923). The problem of nutrition of fungi is closely bound up with the varying degrees of parasitism. The division of fungi into saprophytes and parasites is rather artificial since there are so many intergrading forms. However, it is often convenient to speak of facultative parasites, species which normally are saprophytic, but under especially favorable conditions may develop in the tissues of other organisms, usually as secondary invaders. Other fungi may develop during part of their life cycle as parasites and reproduce sexually only on the dead tissues of the host, or, vice versa, with, a reproductive cycle in the living host and saprophytic vegetative existence outside, as in Cocci- dioides immitis. A few, such as the plant rusts and the Laboulbeniales on living insects, complete their whole life cycle on the living host, and have not been successfully cultivated apart from their hosts. Even more artificial and confusing are the terms employed for symbiosis and related phenomena to express varying degrees of the interaction of two organisms which may vary all the way from parasitism at one extreme to epiphytism at the other. Hydrogen Ion Concentration. — Although biologists have ceased attempt- ing to explain nearly all phenomena by hydrogen ion concentrations, yet these do play an important part in metabolism and should be considered. At this point it might be well to review briefly the underlying concepts and the methods of determining this concentration of hydrogen ions. In a given ionizing solvent, of which water is the only one which can concern us here, solutions of various substances are observed to conduct an electric current to a greater or lesser degree. Pure, distilled water is for aU practical purposes a nonconductor. Solu- tions of sugar for instance, are quite as unaffected by electric current as pure water. Acetic acid in solution conducts a given current slightly, while substances such as HCl, NaCl, and NaOH are strong conductors when dissolved. It is quite reasonable, then, to assume that in conducting solutions there are present conducting units and that the quantity of current conducted is in some way proportional to the numbers of these units. Now, since most of these substances that are good conductors PHYSIOLOGY OF FUNGI 35 when dissolved are nonconductors in the absence of moisture, we must postuUite some change in the solute. Without going any further into the proofs for electrolytic dissociation, we will state here very simply the long accepted fact that if a substance when dissolved in water is found to be a conductor of electric current, it dissociates from the uncharged form into ions capable of carrying current and yet balancing one another in electric charges. The graphic representation of this fact may be made as follows: XY :^ X+ + Y". Arrows are used to indicate that this is an equilibrium, not a completed reaction, that must adjust itself to whatever other equilibria other solutes may necessitate. Let us take, now, the hypothetical acid HA. It dissolves in water and dissociates accord- ing to the following representation: HA ^ H^ + A- At the instant of solution we may visualize the undissociated acid breaking up into H* and A- ions and the dissociated ions recombining to form HA at such rates that the equilibrium for the given acid will be established. Thereafter the rates of dissociation and reassociation must be equal. If, now, we add to the solution, some salt BA, then the concentration of the A" ions will be materially increased. If by any chance we have doubled the concentration of A" we have doubled the possibility of collisions between H* and A" and doubled tlie velocity of the reaction from right to left. If now, we add another acid so that the concentration of H* will be doubled, we have, obviously, quadrupled the possibility of collision. The velocity, then, in a given direction is proportional to the products of the concentrations of the reacting groups. Eepresenting concentrations by square brackets, we may express this as follows: Velocity from right to left kj [H+] [A~] where k^ is the proportionality factor. At the same time, however, we are having HA redissociating at a rate proportional, more or less, to the concentration. Velocity from left to right k^ [HA]. Since, as we have already stated, equilibrium is established, the velocity in one direction must be equal to the velocity in the other. Equating these two expressions, we get: k, [H^] [A-] = k, [HA] or [Hi [A-] ^^^;g- [HA] k, which will be a determinable quantity, characteristic of the solute for which it is determined and referred to as equilibrium or ionizing or dissociation constant. Similarly, a hypothetical base BOH, dissociating as follows: BOH ^ B* + OH- will have a characteristic ionization constant expressed by the equation: Kb = [B-] [0H-] [BOH] And the salt BA: BA ^ B^A- Kba = [Bi [A-] [BA] Assume, now, that we mix equal quantities of two solutions, one of the acid and one of the base, in equivalent concentrations. We will have then in solution the four ions: H+, A", B*, OH". These will adjust themselves so that the constants, Ka, Kb, Kba, 'wiU be satisfied and, in addition, another constant which will express the equilibrium between the H+ and OH" ions as follows : ^ _ [Hi [0H-] 36 MEDICAL MYCOLOGY Since HOH is water and since the quantity formed in a given solution by reassociation of ions is negligible in comparison with the sum total of molecules present, we may for all useful purposes write the equation: Kw ■= [H-] [0H-] The above is a relationship that holds in all aqueous solutions. However concentrated the hydrogen ions, there still must always be enough hydroxyl ions present to satisfy the constant ; however concentrated the hydroxyl, there must still be enough hydrogen ions. Thus, for instance, we might express the acidity of solutions in terms of the hydroxyl ion concentra- tion. As a matter of convention, but not necessity, we express acidity and alkalinity in terms of hydrogen ion concentration. Kw has been determined with considerable care to be practically or lO-i*. Thus 1014 [H+] [0H-] = 10-14 always holds, whatever the solute or solutes. In chemically pure, distilled water, which is of course entirely neutral, [H+] c= [0H-] = 10-7. A neutral solu- tion, then, is one in which this equation holds. For higher concentrations of hydrogen ions, those from [H+] = IQi up to [H+] = 10"^ (remember the exponent is negative) solutions give acid reactions. For lower concentrations, between [H+] = 10"^ and [H+] = lO'i*, solu- tions are alkaline. As a convenience in manipulation, the reciprocal of the hydrogen ion concentration is generally used. This obviates the use of the negative exponent. Thus, in alkaline solutions, varies between 10^ and IQi*, another way of expressing the facts expressed above, but somewhat simpler. It is still simpler, and quite permissible, to use the logarithm of the reciprocal _ _ = 10" logj, = x logjJO. Since tlie logarithm of ten is unity, [H+J H+ log = X. This X, the logarithm of the reciprocal of the hydrogen ion concentration, is H* what is referred to as the pH of a solution. It follows, then, that from pH 1 to pH 7 a solu- tion is acid and from 7 to 14, a solution is alkaline. There are two common methods of measuring hydrogen ion concentration. The most direct method is by measuring the potential of the so-called hydro- gen electrode in a portion of the solution to be determined. It has been found that with this quantity measured, the concentration may be deduced very simply according to the following formula : Potential , 1 = log Numerical factor [H"^] With the factor once determined and the potential directly measured, there should be no further difficulty. The apparatus, however, is expensive and while apparently simple to manipulate is actually unreliable except in the hands of an expert who appreciates the various possible sources of error and is ever on the alert for them. There are many types of potentiometer on the market, but it is safe to say that none of them is fool-proof. The indirect method, the colorimetric, depends upon the fact that certain series of organic compounds exhibit great changes in color with varying pH. Phenolphthalein, for instance, is colorless in acid solution, but becomes ma- genta at pH 9. Litmus is violet in alkaline solutions, red in acid, changing at approximately pH 7. Suitable indicators may be found for almost any desired PHYSIOLOGY OF FUNGI 37 pPI or range of pH's, notable among which are a long series of sulphon- plithaleins and another of azo compounds. There is not space here to go into the theory of indicators or even to list them. Thorough and satisfactory discussions may be found elsewhere. While apparently crude, this method, in addition to the obvious advantages of being cheap and quick, may, in the hands of a normal, intelligent manipulator, be developed to quite a high degree of accuracy. It must be remembered that the color perception of some individuals is very imperfect so that no amount of training will give them satisfactory re- sults Avith a colorimetric method in biochemistry or bacteriology. The ex- tremely simple method of matching colors Avith a colored chart, such as fur- nished by Clark (1920, 1922), is only a very rough approximation and is practically useless Avitli even slightly clouded solutions. A more reliable method, in Avhich the unknown + indicator is compared Avith a standard + indicator as seen through an equal depth of solution Avithout indicator, is quite commonly used and is satisfactory if the unknoAvn solution is not highly colored. It is important that the vessel be of the same size, shape, material, color, and thickness, and that the light intensity be equal. Simple devices may be purchased quite cheaply. By carefully eliminating sources of error by use of a colorimeter of the Duboscq type and varying the light intensity, the Avriter (Dodge 1919, Duggar & Dodge 1919, Duggar 1919) Avas able to secure as great accuracy as is usually attained by the potentiometric method, and extend the range of indicators in both directions so that only half as many need be used as in the usual series proposed by Clark & Lubs (1917). It should also be kept in mind that the alkali sloAvly dissolves from glass, in- creasing the alkalinity of solutions so that standards sealed in glass should not be used indefinitely. The safest Avay is to prepare carefully one's OAvn stand- ards and store them in paraffin lined bottles, for use as occasion demands. These standards, usually mixtures of buffers, are solutions of Aveak acids and their salts, which can maintain their pH almost unaltered on the addition of considerable quantities of strong acids or bases. Any substance capable of remoAdng hydrogen or hydroxyl ions from the solution either physically or chemically Avill act as a buffer; e.g., Bovie (1915) has shoAvn that charcoal has a buffer action. We have, however, to deal only with chemical buffers, a simple explanation of Avhich might not be amiss. Acetic acid (we will indicate the acetate group, CH3COO-, here by the simplified Ac-) is a weak acid. That is, though in a normal solution there is one gram atom of ionizable hydrogen per liter, most of this remains as un-ionized HAc and the reaction of the solution is only slightly acid, the pH being only a little below 7. Sodium acetate, being a salt, is almost completely ionized. NaAc :;±: Na+ + Ac- The Ac- ion will immediately start to take up the H* ion of the water until the dissocia- tion constant for HAc is satisfied. This will liberate an excess of OH- ions which will remain virtually uncombined because NaOH is a very highly ionized base. Thus a normal solution of pure NaAc in pure water will give a definitely alkaline reaction. 38 MEDICAL MYCOLOGY Assume now that we have in a given solution a mixture of solutions of HAc and NaAc in such proportions that the resulting reaction of the mixture is acid. We have, then, present large quantities of Na*, 0H-, H* and Ac- ions and of un-ionized HAc, practically no un- ionized NaOH. Let us now visualize what would happen if to such a solution we add a solution of a strong acid, such as HCl, almost completely ionized to H+ and C1-. The reaction does not, as one might hastily assume, turn sharply acid. Before the pH can be materially lowered, the free H+ added must first adjust its equilibrium with the free Ac- and OH- in the solution. This will be accomplished by association into almost un-ionized HAc and HOH and until this is accomplished, addition of HCl will not materially affect the reaction of the solution. By suitably selecting the acid and its salt, we may secure satisfactory buffers for almost any region of the whole pH scale. Boric acid, H3BO3, and phosphoric acid, HjPO^, are among those most commonly used. These have the double advantage of being weak acids and of ionizing in three stages: e.g. H3PO, ^ H* + H„PO,- H,PO,- ^ H+ -1- HPO,- HPO,= ;^ H+ -f-PO/ The dissociation constants of these three stages are successively smaller, the possible use in buffering correspondingly greater. The actual composition of the buffer solutions we will not go into here, save to state briefly that boric acid is usually used in conjunction with borax, while, instead of using phosphoric acid itself, there is used a mixture of the two salts KH.PO, and Na.HPO, (or, rather, its hydrate). Clark (1922) gives a series of possible buffers, with directions for their preparation and tables showing their pH range. For a comprehensive discussion of the subject one should consult such works as those of Clark (1922) and Michaelis (1926). This matter of buffers is very important in the preparation of media and in interpreting the results of older authors, who measured the total acidity, usually using phenolphthalein as an indicator. It was customary to titrate a sample medium and add the calculated amount of acid or alkali necessary for a given total acidity. If the medium was made up of phosphates, etc., as some of the early media were, this added acid had little real effect, while if few buffers were present it might change the reaction very much. While generalizations are unwarranted, a large number of observations indicate that most bacteria grow best betwen pH 7 and 9 while most fungi grow best between pH 5 and 7 with some, especially members of the Asper- gillaeeae, able to grow slowly at much higher acidities. This fact is sometimes utilized in restraining the growth of bacteria in isolations by using media to which a small quantity of an organic acid, such as acetic or lactic acid, has been added (see p. 59). Klebs' dicta hold in this case as well as in other factors of the environment, since growth is possible at a much wider range than is reproduction. Talice (1930) records optima, maxima, and minima of about thirty common human pathogens on three usual media. (See also notes of Mallinckrodt-Haupt 1929, 1932; Kadisch 1929.) Oxygen Requirements. — Since the fundamental processes of respiration are the same for plants and animals and are already known as to conditions and end products, no attempt will be made to discuss aerobic and anaerobic PHYSIOLOGY OF FUNGI 39 respiration here. Such works as that of Kostyehev and Lyon (1927) and cur- rent textbooks on general physiology may be consulted for further informa- tion. However, one should clearly distinguish between oxidation and fermen- tation phenomena. Space will not permit of a full discussion of this problem here, but some practical suggestions will be given in connection with the methods for the study of fermentation (see p. 62). In general, little atten- tion need be given to a consideration of oxygen supply under the ordinary conditions of culture (Kadisch 1933). It sometimes happens, however, that in small, tightly plugged test tubes there may not be a sufficient amount of oxy- gen present to support sexual reproduction. In some cases a sudden change in oxygen pressure may stimulate reproduction, especially if applied along with other changes in the environment. Temperatiu-e Requirements. — These vary greatly with the species, some of which grow well only at body temperature while others will make good growth at all temperatures between room temperature (20° C.) and body tempera- ture (37.5° C). The earlier authors spent much time in search for an opti- mum temperature, little realizing that a temperature may be optimum for an organism under one set of conditions while far from optimum under another set of conditions (Blackman 1906). Later studies considered the effect of temperature on rate of growth and found that growth roughly follows van 't Hoff's law of doubling the rate for every ten degrees of tem- perature in the middle part of the temperature range, but with a rapid falling off of rate after a certain critical point is reached. The limits of growth are usually much wider than those of reproduction as Klebs postulated. In gen- eral, spores are adapted to withstand higher and lower temperatures than vegetative structures, and ordinarily, thick-walled spores are much more re- sistant than thin-walled spores, while spores resulting from the sexual act are more resistant than those of asexual origin. Influence of Light. — That light has strong morphogenic influence has long been recognized from observations in nature (well summarized by Elfving 1890). Since some organisms develop reproductive organs only in response to light stimuli, light may be of considerable importance in cultures. On the other hand, many fungi seem to develop as well in the dark as in the light. Elfving suggested that light acts as an inhibitor of organic synthesis and that the closer the food available is to the constituents of the protoplasm, the less action the light has. In most plants light, especially of shorter wave lengths, tends to restrict vegetative growth. Many subsequent investigators have ex- tended these early observations. Neidhart (1924) reports lethal action of x-rays and radium in Sporotrichum and Ectotrichophyton gypseum. Nadson & Phillippov (1925) report interesting effects of x-rays on sexuality in Muco- raceae. Dome & White (1931) report differential action of x-rays in different groups of fungi, while using the x-rays for therapeutic purposes. (See also Liebesny, Wertheim & Scholz 1933.) 40 MEDICAL MYCOLOGY Chemotropism. — The strongest chemotropic reaction of liyphae is usually negative ; the hyphae grow away from regions which have been staled by the products of their own metabolism (Clark 1902, Fulton 1906, Balls 1908, Graves 1916, Brown 1922, 1923, 1925, Pratt 1924). A simple example of this reaction is found in the circular growth of mycelia. In so far as a clear field is avail- able, the hyphae tend to grow equally in all directions from the point of in- fection. The same factor may account for the alternate dense and sparse zones which characterize many fungal colonies. Energetic growth results in the deposition of catabolic substances, and growth is accordingly reduced until a few hyphae pass beyond the inhibiting zone and give rise to a new ring", or frequently the germination of fresh spores outside this zone produces a similar effect. Ammonia and potassium bicar- bonate are often among the substances producing staling, as this phenomenon is called. Incubation at higher temperatures hastens staling. Hydrotropism may occur but is difficult to prove. Phototropism. — Many reproductive structures are very sensitive to light and by means of this reaction adjust themselves in a position favorable to the distribution of their spores, since the direction of light is usually that of the direction of open spaces (BuUer 1909-1931, Jolivette 1914, Parr 1918, and Blaauw 1914). It is probable, however, that there is little positive phototropism among the human pathogens. Radium. — Little work has been done on the effect of radium on patho- genic fungi. Sartory & Meyer (1926) report that in Aspergillus fumigatus on media containing salt, exposure to 3-7.2 millicuries, either discontinuous or continuous, produced an increase of conidiophores, with a tendency to de- crease the size of the head and approach conditions found in Penicillium. On media nearly free from salts, there was a tendency to form large-celled oidia rich in oils, or large, thick-walled spores, 3-8/a in diameter, singly or in pairs, and large pseudosporangia, 30/i, in diameter, with echinulate walls but no spores observed within them. It was noted at the same time that in dissociated media reducing power was lowered and pH was increased. (Sucrose 5 gm., gelatin 7.5 gm., NaCl 1 gm., carrot juice q. s. for 100 c.c.) In undissociated media, reducing power was increased and pH was decreased. "With higher dosage (10.2 millicuries per sq. cm.), hard, fusiform sclerotia were produced in submersed mycelium. These contained perithecia. BIBLIOGRAPHY Armstrong, G. M. 1921. Sulphur nutrition; the use of thiosulfate as influenced by hydrogen ion concentration, Ann. Missouri Bot. Gard. 8: 237-281. Balls, W. Lawrence. 1908. Temperature and growth, Ann. Bot. 22: 557-591, 11 figs. Blaauw, A. H. 1914. Licht und Wachstum I, Zeitschr. Bot. 6: 611-703. Blackman, F. P. 1906. Optima and limiting factors, Ann. Bot. 19: 281-295, ^ fiffs. Bokorny, Th. 1917. Benzolverbindungen als Nahrsubstanzen, Zentralbl. Physiol. 32: 55-63. Bovie, William T. 1915. A direct reading potentiometer for measuring and recording both the actual and the total reactions of solutions, Jour. Med. Bes. 33: 295-322, 14 figs. PHYSIOLOGY OF FUNGI 41 Brenner, W. 1911. Untersuchungen iiber die Stickstoffernalirung des Aspergillus niger und deren Verwertung, Ber. Deutsch. Bot. Ges. 29: 479-483. Brown, William. 1922. On the germination and growth of fungi at various temperatures and concentrations of oxygen and carbon dioxide, Ann. Bot. 36: 257-283, 4 figs. — . 1923. Experiments on the growth of fungi in culture media, Ann. Bot. 37: 105-129, 7 figs. — . 1925. Studies in the genus Fusarium. II. An analysis of factors which determine certain microscopic features of Fusarium strains, Ann. Bot. 39: 373-40S. BuUer, Arthur Henry Reginald. 1909-1934. Researches on Fungi, vols. 1-6. Longmans Green & Co. London. Cerutti, Pietro. 1933. Conccntrazione idrogenionica e sviluppo degli ifomiceti patogeni: ricerche sperimentale e cliniche, Patkologica 25: 32-37. Clark, Judson F. 1902. On the toxic properties of some copper compounds with special refer- ence to Bordeaux mixture, Bot. Gas. 33: 26-48, 7 figs. Clark, William Mansfield. 1920, 1922. The determination of hydrogen ions, Baltimore, Williams & Wilkins Co., 317 pp., 1920; ed. 2, 480 pp., 1922. Clark, William Mansfield So H. A. Lubs. 1917. The colorimetric determination of hydrogen ions and its applications in bacteriology, J. Bad. 2: 1-109. Coons, George H. 1916. Factors involved in the growth and the pycnidium formation of Plenodomus fuscomaculans. Jour. Agr. Bes. 5: 713-769. Czapek, Karl. 1902, 1903. Untersuchungen iiber die Stickstoffgewinnung und Eiweissbildung der Schimmelpilze, Beitr. Chem. Physiol, u. Path. 1: 538-560, 1902; 3: 47-66, 1903. Dodge, Carroll William. 1919. [The fate of] Tyrosine in fungi: chemistry & methods of studying the tyrosinase reaction, Ann. Missouri Bot. Gard. 6: 71-92. Dome, Maurice & Cleveland Wliite. 1931. Treatment of superficial infections with the long wave length Roentgen rays (Grenz rays), Arch. Derm. Syphilol. 24: 409-416. Duclaux, E. 1885. Sur la valeur alimentaire de diverges substances pour I'Aspergillus niger, C. B. Soc. Biol. 37: 91-94. — . 1889. Sur la nutrition intracellulaire, Ann. Inst. Pasteur. 3: 97-112, 413-428. Duggar, Benjamin Minge. 1919. The microcolorimeter in the indicator method of hydrogen ion determination, Ann. Missouri Bot. Gard. 6: 179-181. Duggar, Benjamin Minge & A. R. Davis. 1916. Nitrogen fixation, Ann. Missouri Bot. Gard. 3: 413-437. Duggar, Benjamin Minge & Carroll William Dodge. 1919. The use of the colorimeter in the indicator method of hydrogen ion determination, Ann. Missouri Bot. Gard. 6: 61-71. Ekman, Gunnar. 1911. Studien iiber den Nahrwert einiger KohlenstoffqueUen fiir Aspergillus niger van Tiegh, ofversight af FinsTca Vetensh Soc. Forh. 53A: 16: 1-43. Freedman, Louis & Casimir Funk: Nutritional factors in the growth of yeasts and bacteria, I. Vitamines, /. Metai. Res. 1: 457-468. II. Protein hydrolysates, IMd. 1: 469-480. Fulton, Harry R. 1906. Chemotropism of fungi, Bot. Gas. 41: 81-108. Graves, Arthur Harraount. 1916. Chemotropism in Rhizopus nigricans, Bot. Gaz. 62: 337- 369, 4 figs. Janke, Alexander & Stephan Kropacsy. 1929. Die Bestimmung des Wasserexponenten mittels des Stufenphotometers. I, Biochem. Zeitschr. 213: 154-169. Jolivette, Hally D. M. 1914. Studies in the reactions of Pilobolus to light stimuli, Bot. Gaz. 57: 89-121, 1£ figs. Kadisch, E. 1929. tjber die Bedeutung der Nahrbodenalkalitat in der Mykologie, Derm. Zeitschr. 55: 385-396. — . 1933. Der Einfluss von Temperatur und Sauerstoif auf die Lokalisation der Infektionen. Wiigende Untersuchungen an Fadenpilzen. Untersuchungen mit dem Pulfrich Photo- meter an Hefen. Modifikation des Pulfrich Photometers, Arch. Derm. Syphilis 168: 438-475, 10 figs. 42 MEDICAL MYCOLOGY Klebs, Georg. 1896. Die Bedingungen der Fortpflanzung bei einigen Algen und Pilzen, Jena, G. Fischer, 543 pp., Spls. Klotz, L. J. 1923. Some aspects of nitrogen metabolism in fungi, Ann. Missmiri Bot. Gard. 10: 299-368. Kossowicz, A. 1912. Die Zersetsung von Harnstoff, Harnsaure Hippursaure und Glykokoll durch ScMmmelpilze, Zeitsch. f. Garungs-Physiol. 1: 60-62; 2: 51-54. Nitritassimila- tion. Ibid. 2: 55-58. tJber das Verhalten einiger Schimmelpilze zu Kalkstickstoff, Ibid. 2: 124, 125, 1912. — . 1914. Zur Kenntnis der assimilation von Kolilenstoff- und Stickstoff-verbindungen durch Schimmelpilze, Bioch. Zeitschr. 67: 391-399. tJber das Verhalten von Hefen und Schimmelpilze zu Nitraten, Ibid. 67: 400-420. Kostychev, S. & C. J. Lyon. 1927. Plant respiration, Philadelphia, P. Blakiston's Son So Co., 163 pp. Liebesny, P., H. Wertheim & H. Scholz. 1933. tJber Beeinflussung des Wachstums von Mikro- organismen durch Kurzwellenbestrahlung I, Klin. Woch. 12: 141-145. Lutz, L. 1905. Sur 1 'assimilabilite comparee des sels ammoniacaux, des amines, des amides et des nitriles, C. B. Acad. Sci. Paris 140: 665-667. McHargue, J. S. & R. K. Calfee. 1931. Effect of manganese, copper, and zinc on growth and metabolism of Aspergillus flavus and Ehizopus nigricans, Bot. Gaz. 91: 182-193. Mallinckrodt-Haupt, Asta St. von. 1929. pH Messungen bei Pilzkulturen, Derm. Zeitschr. 55: 374-384. — . 1932. Der Wert der pH Messung bei Pilzkulturen, ZentralU. BaU. I. 125: 368-374. Meyer, Jacques. 1927. Contribution a 1 'etude de 1 'Aspergillus fumigatus Fresenius. ifitude experimentale de 1 'influence du radium et des milieux sur la production de phenomenes sexu^s, These Fac. Pharm. Univ. Strasbourg 31: 1-112, SO figs. Michaelis, L. 1926. Hydrogen ion concentration, its significance in the biological sciences and methods for its determinations. I. Principles of the theory. Authorized trans- lation by William A. Perlzweig, Baltimore, Maryland. Williams & Wilkins Company. Nadson, G. A. & G. S. Philippov. 1925. Influence des rayons X sur la sexualite et la forma- tion des mutantes chez les champignons inferieurs (Mucorinees), C. B. Soc. Biol. 93: 473-475. Naegeli, Carl. 1880. Ernahrung der niederen Pilze, Bot. Mitt. 3: 395-483. K. Ahad. Wiss. Milnchen, Math. Nat. CI., Sitsungsber. 10: 277-367. — . 1882. Ernahrung der niederen Pilze durch Kolilenstoff- und Stickstoffverbindungen. Un- tersuchungen uber niederen Pilze aus dem, Pflanzenphysiologischen Institut in Miinchen. 1-75. Neidhart, Leo. 1924. Beitrag zur Strahlenempfindlichkeit pathogener Hautpilze (Sporo- trichum Beurmanni und Trichophyton gypseum), Inaug. Diss. Med. FaTc. Univ. Zurich, 29 pp. Parr, Rosalie. 1918. The response of Pilobolus to light, Ann. Bot. 32: 177-205. Pasteur, Louis. 1860. Memoire sur la fermentation alcoolique, Ann. Chim. et Phys. Ill 58: 323-426. — . 1862. Memoire sur les corpuscules organises qui existent dans 1 'atmosphere, examen de la doctrine des generations spontanees, Ann. Chim. et Phys. Ill 64: 1-110. [IX. Sur le mode de nutrition des ferments proprement dits, des mucedinees et des vibrioniens, pp. 100-110.] Peskett, Geoffrey Lewis. 1933. Growth factors of lower organisms, Biol. Rev. 8: 1-45. Pratt, Clara A. 1924. The staling of fungal cultures. II. The alkaline metabolic products and their effect on the growth of fungal spores, Ann. Bot. 38: 599-615, 1 fig. *Eeinke, J. 1883. Unters. Bot. Lab. Gottingen 3: 13. Hitter, G. E. 1909, 1912. Ammoniak und Nitrate als Stickst off quelle fiir Schimmelpilze, Ber. Deutsch. Bot. Ges. 27: 582-588, 1909; 29: 570-577, 1912. PHYSIOLOGY OF FUNGI 43 — . 3914. Ammonnitrat und freie Saltpetersiiure als Stickstoff quelle fiir Schimmelpilze, Biochem. Zeitsch. 60: 370-377. Eobertson, R. C. & D. J. Davis. 1923. Food accessory factors (vitamins) in bacterial growth, observations on the ultimate source of accessory growth substances for yeast, Jour. Infect. Bis. 32: 153-158. Sartory, Antoine, E. Sartory, & Jacques Meyer. 1926. Etude de Paction du radium sur 1 'Aspergillus fumicatus Tresenius en culture sur milieux dissocies et non dissocies, C. B. Acad. Sci. 183: 77-79. — . 1926. La formation des peritheces chez 1 'Aspergillus fumigatus Fresenius sous 1 'influence du radium, C. B. Acad. Sci. 183: 1360-1362. Sergent, A. L. 1928. Les facteurs de croissance des microbes sur milieux artificiels, These Doct. Med. Paris, 182 pp. Talice, Eodolfo V. 1930. Le facteur pH en mycologie. Son influence sur la culture de certaines especes de champignons parasites de I'homme, Ann. Parasitol. Hum. Cornp. 8: 183-188, 2 talles. Waterman, H. J. 1913. Over eenige factoren, die de ontwikkeling van Penicillium glaucum beinvloeden Proefschr. Delft. [tJber einige Faktoren, welche die Entwicklung von Penicillium glaucum beeinflussen. Beitrag zur Kenntnis der Antiseptica und der Narkose. Centralbl. Baht. II 42: 639-688.] Went, F. A. F. C. 1901. Uber den Einfluss der Nahrung auf die Enzymbildung durch Monilia sitophila (Mont.) Sacc, Jahrb. Wiss. Bot. [Pringsheim] 36: 611-664. CHAPTER III CULTURE MEDIA, THEIR PREPARATION AND STERILIZATION Probably the oldest methods of cultivating fungi were developed in grow- ing the common mushroom of commerce {Psalliota campestris and related species), but these methods had little, if any, influence on the scientific study and cultivation of fungi. Undoubtedly many mycologists of the nineteenth century brought into their laboratories young fructifications attached to their substrate, and watched their development, also making many important ob- servations on material of early stages collected in the field. It remained, how- ever, for Pasteur and Koch in the decade 1873-1883 to develop methods of sterilization, isolation, and pure culture of bacteria to pave the way for our present technique. Earlier authors in several instances had anticipated these methods more or less completely, but their accounts had either been forgotten, buried under the debris of the theory of spontaneous generation, or lost in a little-known periodical of very limited circulation ; e.g., the work of Bizio on Serratia marcescens (Bacillus prodigiosus) which was published in 1823 and was only generally known among scientists after its translation and publication in 1924. For a more complete account of the historj^ of bacteriologic methods the reader is referred to the excellent short account in Conn & Conn's Bac- teriology and, for formulae of the media used to Desgardes (1921) and to Levine and Schoenlein (1930). In all work with pure cultures it is essential that everything used should be clean and sterile. This statement seems so axiomatic to the well-trained medical man that its emphasis here may appear out of place, but in this gen- eration when so much routine work is left to technicians and even humbler laboratory folk, it may not be out of place. Also in my classes I frequently find students with so little previous training in bacteriology that in the fol- lowing pages I shall not assume any previous experience in handling micro- organisms in pure culture. The cleaning of glassware, while one of the drudgeries of the laboratory, is very important.* Needless to say the glassware should be washed with soap and water until clean tap water will drain freely from it, and not hang in drops over the sides of test tubes, etc. This stage may be called physically clean. We must then be sure that it is chemically clean, i.e., that it does not contain any substances, minute traces of which may inhibit or cause abnormal growth of the organism to be cultivated. ♦If the glassware has been used for cultures, it should be sterilized by autoclaving (see pp. 47, 48) before proceeding' to wash it. Pautrier and Rietmann (1924) report infection of a laboratory worker with Trichophyton granulosum (ordinarily confined to horses) while clean- ing tubes containing year-old cultures ! 44 CULTURE MEDIA 45 Some new glassware when first received into the laboratory may still contain enough alkali to change considerably the hydrogen ion concentration of the medium to be employed. It may be necessary to heat this glassware with strong acids before using it for careful work, although it may still be used for many of the more tolerant organisms without further treatment. In many laboratories all glassware is treated with a strong oxidizing agent, such as an acid solution of sodium or potassium bichromate, commonly called cleaning solution. This tends to neutralize any free alkali in the glass as well as to destroy any life or organic material. This is especially necessary if dry sterilization of the glassware is expected, as failure to remove all organic material will result in a deposit of carbon which reduces visibility and is almost impossible to remove later, although it probably does not interfere with the growth of the organism. After treatment with cleaning solution (if necessary), the glassware should be thoroughly rinsed with distilled water. If dry sterilization takes place after hard tap water has been used for rinsing, a deposit of salt may be baked in, causing decreased visibility and, rarely, toxicity. The alternative method of rinsing with the solution to be used is not recommended in ordi- nary practice, a^ some liquid may adhere to the top of the tube of the neck of a flask and cause trouble in later procedures. Sterilization. — Many methods have been developed in the last half cen- tury, but none is equally good for everything, consequently the principles underlying each should be thoroughly understood and the proper method selected for the material in hand, with a clear understanding of its limitations. Chemical Methods (Disinfectants, Antiseptics). — These methods as com- monly applied in the laboratory consist in treating the material with a toxic chemical substance for a sufficient time to destroy all life, then removing the chemical substance and preserving the material from further contact with microorganisms until it is used. The methods are difficult and little used when other methods will serve. The chemical nature of the disinfectant must al- ways be considered, as well as its fungicidal power. Disinfectants may be grouped as halogens, salts of heavy metals, and organic compounds. Halogfens. — Fluorine and bromine are rarely used under present condi- tions. All of the halogens corrode metals and are difficult to handle. They may prove toxic to living tissue, although they are very valuable in some pro- cedures. Chlorine usually is applied as a solution of a hypochlorite which slowly gives off the chlorine in intimate contact with the material to be dis- infected. Improvements of this technic have been found useful as antiseptic dressings, disinfection of seeds in laboratory practice, etc. Calcium hypo- chlorite (bleaching powder, chloride of lime, sodium or potassium hypochlorite) is also used as a deodorant and disinfectant in outhouses and for sterilizing white clothing where other methods are not easily available. Iodine is usually used in alcoholic or potassium iodide solution or in the organic compound iodoform. This is one of the common disinfectants for 46 MEDICAL MYCOLOGY superficial lesions but causes unpleasant discolorations and in some individuals severe poisoning. It is rarely used about the laboratory as a disinfectant. The alcoholic solution is occasionally used as a counterirritant. Salts of Heavy Metals. — In general, salts of heavy metals are toxic to or- ganisms roughly in the order of the magnitude of their atomic weights, al- though the salt radical has some less clearly defined influence on toxicity. Copper compounds are frequently used in agricultural practice as a fungicide but rarely about the laboratory. Compounds of mercury have had great vogue, especially in some laboratories, but they have some severe drawbacks and probably many may be well replaced by other disinfectants. The oldest and most widely used is mercuric chloride (bichloride of mercury, corrosive sublimate). Toxic in extreme dilution, the solution is apt to dry, leaving tiny crystals which blow about the laboratory and occasionally prevent growth when all other conditions are favorable. Glassware which has been used for mercuric chloride solutions should never be used for cultures again, for if growth takes place at all, it is apt to be abnormal and stunted. In some per- sons it also causes severe dermatoses extending over several years. The recently developed mercurochrome has eliminated several of these objections. Silver nitrate (lunar caustic) has long been in use and recently some of the organic silver salts, especially nucleinates (argyrol and similar com- pounds) have been widely used about the mouth, ej'es, and genitalia. Organic Compounds. — Of the innumerable organic compounds only the lower aliphatic alcohols and aldehydes and the simpler compounds of the aromatic series, such as phenols and salicylic compounds, have found wide use. Methyl alcohol is quite efficient, but is rarely employed on account of the injury of the optic nerve by the vapors. Ethyl alcohol is perhaps the most widely used of the group. Absolute ethyl alcohol has little value, but the intermediate dilutions with water are good, especially for sterilizing cutting instruments which cannot be subjected to heat or to the stronger, but more corrosive, disinfectants. The higher alcohols are little used. Formaldehyde had a great vogue at one time as a gaseous disinfectant, but now is seldom used about the laboratory, except in aqueous solution (formalin) in seed disinfection and as a cheap preservative for class material. Phenol (often knoAvn in its 4% aqueous solution as carbolic acid) has stood the test of half a century, although its popularity has varied. For many years it was taken as a standard for comparison of the efficiency of antiseptics. Salicylic compounds belong rather to medicine, although salicylic acid itself is used externally in dermatology as a keratolytic agent. Volatile oils have been found useful with pathogens of the skin (Kingery et al. 1928, 1929). Physical Ag-ents. — Heat and light are the only sterilizing agents in gen- eral use at present, although it is probable that radiations of still shorter wave lengths would be effective if not so expensive. Light, especially the ultra- CUIiTURE MEDIA 47 violet end of the spectrum, at present belongs rather to the realm of thera- peutics than to laboratory practice and need not be considered here, although it may play a great part in hygiene. Hot dry air is one of the oldest methods employed and is still used for glassware, such as Petri dishes. The clean material, usually heat resistant glass, is put into a gas oven (rarely electric) heated to 150°-180° C. for from a half hour to an hour or more, depending upon the material to be sterilized and the length of time necessary to kill spore-forming organisms which may be found in a given laboratory. Passing material through a flame for a varying period is also a very old practice, now mostly used for sterilizing inoculating tools, such as needles, platinum spatulas, etc. The metal is ordinarily heated to redness in the flame, then allowed to cool to room temperature without touching any object which might contaminate it until used. The former surgical practice of cautery with red hot iron probably owed its success in part to this method of sterilization. It is obvious that this method is not available for tempered tools, such as scalpels, etc. A variant of this practice is to dip in alcohol and ignite, prob- ably less efficient but adequate in many cases. Steam. — None of the above-mentioned methods are useful for most cul- ture media, and it was only with the use of steam that culture media in the modern sense began to develop rapidly. Steam may be applied at atmospheric pressure in which case the medium reaches 100° C. and is held at that tem- perature for a period. This is usually carried out in an Arnold steam steri- lizer, a simple apparatus for generating steam with a minimum loss of water during the process. When spores were discovered in bacteria, Tyndall ap- plied this knowledge by proposing discontinuous sterilization, by heating the medium for a definite period, sufficiently long to kill all the organisms in the active vegetative state, then incubating sufficiently long to allow the spores to develop the vegetative stage, and heating again. This process must be repeated until the medium remains sterile on incubation. Under ordinary conditions the usual procedure is to heat in steam at 100° C. for an hour each day for three successive days. If the period between sterilizations is too long, the spores may have germinated and formed new spores, and if too short they may not all have germinated. The long time necessary to prepare media by this method, as well as several more theoretical objections, has tended to eliminate this method from the laboratory. However, some biologic products used as media are profoundly altered by higher temperatures, and it is neces- sarj^ to employ this method for these substances. Superheated Steam. — In this method the steam is confined in an autoclave up to any pressure desired, instead of being allowed to escape as in the ordi- nary steam sterilizer. A good steam pressure gauge on the autoclave is requisite, and a thermometer is not only desirable but also an additional safe- guard. The temperature ordinarily employed is 115°-125° C. or about 10-20 pounds pressure per square inch. An exposure to about 120° C. (or 15 lb.) for 15-20 minutes will sterilize most media, unless some very heat resistant 48 MEDICAL MYCOLOGY organism is present. The period of sterilization must of course be measured from the time the desired temperature is attained and it may require 10-15 minutes, even with a strong heating unit, to reach this temperature. These temperatures, especially if they are prolonged beyond the minimum necessary to secure complete sterilization, may injure or transform the more labile organic compounds, especially if the acidity or alkalinity of the medium is such as to favor hydrolysis of the higher carbohydrates to hexoses, etc. The autoclave may be heated by electricity or gas or connected with a steam supply pipe if there is sufficient pressure maintained. Care should be taken to see that sufficient water is present and that the lid is wiped free from dust and dirt before each sterilization. After the autoclave is full of steam and the thermometer registers 100° C, the vent is closed. It is not advisable to leave the autoclave without observation, although if the safety valve is properly set, steam will escape after the desired pressure is reached. As soon as this occurs one may safely cut down on the heat supply, since a rapid escape of steam soon exhausts the small supply of water, and often dislocates the cotton plugs or causes the medium to boil up and wet the plugs. When the necessary time for sterilization has elapsed, the gas is turned off, and the autoclave is allowed to cool until the temperature reaches 100° C. before open- ing. An experienced operator may open the cock and allow the steam to escape slowly before the pressure is wholly down, but this procedure is not advised for a beginner. Under field conditions, I have found one of the various aluminium pres- sure cookers now on the market very useful, although only a comparatively small amount of media may be sterilized at one time. Filtration. — The passage of the medium through a filter with pores smaller than bacteria is a possible though tedious process which may have to be used in cases where biologic products are so altered by heat that it is impossible otherwise to retain them in condition to use as media. Asepsis. — The careful excision of bits of plant tissue, under condition of asepsis approaching that obtaining in the operating room of a hospital, with thoroughly sterilized instruments, usually in a special room (culture chamber) where the air is kept relatively free from dust or microorganisms, is sometimes practiced. Such tissues should be incubated for a sufficient time to insure that they have not been accidentally contaminated during their preparation. This method in its simpler form is one of the oldest ways of securing culture media, but is not much used today. CULTURE MEDIA Media may be roughly classified as liquid and solid. The former were much used by the earlier workers, but at present are little used except in certain physiologic studies where solids interfere with chemical procedures and in a few other special cases. They are still employed in studies of spore srermination. CULTURE MEDIA 49 Liquid Media. — The earlier media were mostly naturally occurring liquids, such as tap water, milk, urine, etc., often with one or more other substances added. Physiologic studies then developed a series of solutions of known chemi- cal composition usually consisting of a basal solution containing all the metallic elements needed for growth and one or more organic compounds containing the requisite sources of carbon and nitrogen. Perhaps the solution known as Czapek's or Dox's solution is more frequently used by mycologists. Czapek's basal solution: Distilled water 1000. c.c. K,HPO, 1. gm KCl 0.5 gm MgS0,.7H,0 0.5 gm FeSO, 0.01 gm For other useful formulae see Levine and Schoenlein (1930). Besides a host of formulae of liquid media of definite chemical composition which have been developed in connection with physiologic studies on com- paratively narrow ranges of organisms, there are many, both solid and liquid, in common use, in which one or more of the principal constituents are aqueous extracts. These extracts may be classified as infusions and decoctions. Infu- sions are prepared by allowing the material to be extracted, more or less finely divided, to remain in contact with water, either cold or lukewarm, for a con- siderable length of time. Hay infusion was one of the very early media of this class but has practically disappeared. Meat infusions are still greatly used in the cultivation of bacteria, although much less important in the cul- tivation of most groups of fungi. The following directions may be taken as a sample of this type : IMacerate 1 part finely chopped lean meat with 2 parts distilled water in an ice box for 18 hours, stirring occasionally. Strain while cold through a fine cloth. Add 1.0% peptone and 0.5% NaCl to the filtrate. Heat until solu- tion is complete. Add NaOH until the reaction is slightly alkaline (practically neutral to phenolphthalein). Heat on a water-bath for 30 minutes and boil for 5 minutes over a free flame. Filter while hot through paper or cotton and cloth. Add 1.0% of desired nutrients. Adjust reaction as necessary (see p. 38). For the many variants of this method, consult Levine and Schoenlein. Most of the commonly employed media of this type may now be obtained in dehydrated form. In the preparation of these media the directions furnished by the manufacturer should be followed. Decoctions are usually employed with vegetable substances, as the process is more rapid and rarely are there suflScient proteins to cause trouble. Duggar (1909) proposes that for every 1000 c.c. of water in the decoction the equiva- lent of 50 gm. dry weight should be used. The plant product is washed, peeled if necessary, thinly sliced, and the necessary water added. It is boiled in a steam sterilizer for 2 hours or placed in the autoclave at 115° C. for 20 min- utes, or may be boiled over a free flame for a corresponding period, care being 50 MEDICAL MYCOLOGY taken that the vegetable does not cook on the bottom of the container and that water lost by evaporation is replaced. The decoction is then strained to remove the larger particles of vegetable and may be filtered through paper. Potato decoctions are very difficult to filter, as further precipitation may follow sterilization. Solid Media.^Some of the first media of this class were slices or plugs of vegetables, either raw or cooked (during the sterilization process). Some of these substances, such as string beans, prunes, squash, potato, and root crops, are still used in the study of plant pathogens and similar vegetable products have been used occasionally by isolated workers in the tropics with- out easy access to the more usual types of media. These are frequently very good for producing abundant vegetative growth but less satisfactory for secur- ing reproductive organs. If the vegetable is to be used raw, it must be carefully washed, the out- side thoroughly sterilized by alcohol which is allowed to evaporate or by repeated washing with sterile water in an atmosphere relatively free from dust or spores. Cylinders are then cut out with a sterile cork borer, slightly smaller than the diameter of the test tube to be used, and sliced diagonally. These pieces are then placed either in a special sterile test tube or in a test tube containing one or two glass beads and a small amount of water. If there is no objection to having the vegetable slant cooked, it may be prepared under clean but not necessarily sterile conditions and the whole sterilized together. Glycerol is sometimes added instead of, or in addition to, water in order to insure the surface of the slant remaining moist. Of course the glycerol should also be considered as a possible additional source of carbon. Rarely bits of meat or fish have been sterilized and used directly as media (Sawyer 1930, Rewb ridge. Dodge and Ayers 1929). The other solid media are all colloidal gels, either silicates, proteins, or carbohydrates. Silicates are somewhat difficult to prepare and are at present principally used where it is essential to know definitely the chemical con- stituents of the medium in physiological studies or in the rare cases in which the organism is capable of attacking and digesting the medium. Egg albumen and gelatin are the principal protein media used, mostly in the study of bacteria. Historically, gelatin has been used for a very long time and many of the important early methods and results were obtained with this medium. The directions of Dalmau (1929, 1930) for its preparation are useful, especially for workers in tropical countries. To 900 c.c. nutrient broth add 200 gm. of gelatin and heat in a water-bath until dissolved. The Bacto Gelatin and Pfanstiehl's brand are highly acid, about pH 3 or 4. Adjust reaction to about 6.5 or any desirable pH, let cool to 50° C, add whites of 2 eggs shaken in 100 c.c. nutrient broth, and bring it rapidly to a boil at 100° C. for 10 minutes in a double boiler with saturated salt solution in the lower compartment. The coagulum should clear the solu- tion. Filter through cotton, or cotton and gauze if necessary. Distribute in CULTURE MEDIA 51 tubes and sterilize at 100° C. for 15 minutes on each of three successive days. Cool rapidly after withdrawal from the sterilizer to obtain a hard gelatin, which will not liquefy at tropical room temperatures (28° C). At present it is little used except in studies of the ability of organisms to attack and digest protein. Sawyer (1930) has recently used egg yolk with very good results in his work on the Entomophthoraceae, fungous parasites of insects. The carbohydrate media are chiefly starches and agar. Commercial starches are used, either corn, potato (laundry) or inulin. Ten per cent starch is com- monly used and a colloidal solution obtained by short boiling. The medium is tubed and sterilized with or without the addition of salts or other nutrients. The hydrogen ion concentration of the added material should be carefully considered, as any considerable acidity or alkalinity will hydrolyze the starch, defeating the purpose of the medium by preventing solidification and provid- ing sugar. These media have not been widely used on this account. A variant of this medium, in which com meal mush is prepared, has been employed, al- though perhaps less frequently than the related corn meal agar. The container is partially filled with corn meal which is then thoroughly moistened with hot water (it is difficult to wet it thoroughly with cold water) and sterilized in the usual manner. This medium is never clear as the starch media may be, but it is easy to prepare, has enough of the inorganic compounds and; sources of nitrogen to support growth without further addition, and provides a medium rich in starch. The agars and similar compounds are complex substances which in part hydrolyze to galactose. They are produced during the metabolism of the marine algae, especially the Rhodophyceae (red algae), and comparatively little is known of their chemical structures. The agar of commerce is largely the product of various species of the Gelidiaceae found on the coast of Japan. It solidifies at a comparatively low temperature (about 40° C.) and melts at a very high one (about 95° C). The modern sources of supply have improved the quality very much, so that it seldom contains undesirable salts or nitrog- enous substances. Naturally, for very careful physiologic work, this point would be checked up before using it. Agar may usually be had in shreds, chopped shreds, or in powdered form. The variant methods of preparing agar found in various laboratories give about equally satisfactory final products. Since the medium carries practically no available nutrient, this is added in the form of a solution, infusion or decoction, or a combination of these. The formulae for these are almost infinite, Levine and Schoenlein recording 803 formulae, not counting the numerous variants in proportions and procedures. Of these, one of the simplest and perhaps the most widely used in culti- vating human pathogens is Sabouraud's conservation agar: Water 1000 c.c. Peptone 10 gm. Agar 18 gm. 52 MEDICAL MYCOLOGY Growth is slow on this medinm since the peptone furnishes both carbon and nitrogen, but pleomorphism of the dermatopliytes is greatly delayed. Per- sonally, following a suggestion of Thaxter who had long experience with other groups of fungi in culture, I prefer to add 40 gm. of agar instead of 18 gm. to media to be used for stock cultures. Growth is extremely slow and the cultures do not dry out so rapidly, both of which conditions are advanta- geous with routine stock cultures, as the labor of preparation of media and of transfer is greatly reduced. For isolation and study of the organisms, Sabouraud's test agar (1908), commonly called Sabouraud agar, contains 40 gm. of crude maltose. Sabour- aud has been severely criticized for using the crude sugar, as samples vary greatly, one sample used by Sabouraud on analysis yielding mostly glucose (Hodges 1928). For most organisms glucose gives equally good growth, in fact pure maltose is generally poorer. There seems little difference whether 10 or 40 gm. per liter are added. The firm from which Sal)ouraud secured his maltose ceased business during the World War (1914-1918) and since then, much research has been expended to secure a substitute product which will produce the same giant colonies as those figured by Sabouraud (1910). Sab- ouraud himself (1925) advocates the use of 8% honey, but again introduces a source of error, since honey varies very much according to the species of bee producing it and the flowers from which it is made; e.g., Berde (1926) failed to secure characteristic colonies on honey made from flowers of Rohinm or Stachys annua. Weidman «& Macmillan (1921) and Weidman (1928) in very extensive studies found that crude glucose gave equally good results, Fairchild's peptone being used instead of Chassaing. This medium is often referred to as the Pennsylvania medium. Goldschmidt (1924) proposed the following for English laboratories: Glucose 40 gm. Agar 20 gm. Peptone (Fairchild) 10 gm. Lemco (ordinary not laboratory) 5 gm. NaCl 5 gm. Tap water 1000 c.c. Lemco is an acid meat extract. Ingredients digested in a steamer for 1 hour, adjusted by "soda" to pH 6, sterilized 20 minutes each on 3 successive days. Gruetz (1923) proposed the following for German laboratories: Peptone (Knoll) 5 gm. Nervina Malz from Christiansen, Flensburg 80 gm. Agar IS gm. Water 1000 c.c. Pollacci (1922) proposed the following for Italian laboratories: 500 gm. ground beef in 1000 c.c. water; cook and filter; add 100 gm. Witte peptone, 5 gm, sodium chloride ; heat, filter, warm, and neutralize ; heat for half an CULTURE MEDIA 53 hour, filter, and add 70 gm. glucose and agar. Bruhns (1928) finds this medium superior to that of Gruetz. The medium proposed by Macleod (1928) for the cultivation of Malassezia furfur is quite similar: Agar 1.5 gm. Peptone (Cliassaing) 2 gm. Glycerol 2 c.c. Glacial acetic acid 1 m. Distilled water 100 c.c. Grigorakis (1931) obtained interesting results with 40 gm. of glycerol added to Sabouraud's conservation agar. Benedek advocates 8% crude glucose and peptone (both Merck products). Farley (1920) adds 3-5% human blood heated to 55° C. for 30 minutes to Sabouraud's test agar when cultivating Epider- mophyton. Gentian violet 1 :500,000 is a useful addition to restrain bacterial growth during the isolation of these organisms. I have found the prepared medium of the Digestive Ferments Company satisfactory as Avell as media prepared from ingredients furnished by them. Their peptone is too near neutral to reproduce exactly the giant colonies figured by Sabouraud (1910). On the other hand, there is the advantage of greater uniformity of different batches, something greatly to be desired when comparing results secured at different times. Recently Langeron & ]\Iilochevitch (1930) and others have returned to the cereal and dung media, of variable composition, but have secured inter- esting morphology not produced on the classic Sabouraud medium and its numerous variants. Nannizzi (1926), on the other hand, turned to equally variable animal products, such as bits of skin, nail clippings, horn, feathers, hair, and bone. In this direction the work of Karrenberg (1933) seems to be much the most promising. Using the brain medium of Hibler which was originally developed for anaerobic bacteria, he has maintained stock cultures over very long periods without pleomorphism or loss of virulence. While I have had no personal experience with this medium, it seems so promising that the details of its preparation may be given: Brains of recently slaughtered animals (within 24 hours) are freed from the pia mater, ground in a meat chopper and weighed. Two parts water to one of brains are added, rubbed through a sieve, and cooked for two hours in a steam sterilizer. The next day the medium is tubed and sterilized in the autoclave. Hach and Karrenberg both suggest 0.85% sodium chloride solution instead of tap water and Karrenberg found the medium .satisfactory without rubbing through a sieve. Independently Grigorakis (1933) has proposed a similar medium based on calf spleen. Pulp of calf spleen 500 gin. Peptone 10 gm. Agar 18 gm. Water 1000 c.c. 54 MEDICAL MYCOLOGY Many workers have reported excellent results with various fungi on car- rot agar. The following formula of Falchi may be considered typical: Carrots 500 gm. Water 1000 c.c. Agar 20 gm. Peptone (Rostock) 10 gm. Among the formulae for synthetic media, those of Currie (1917) and Fulmer & Grimes (1923) are widely used. Currie : Ammonium nitrate 2.5 gm. Potassium dihydrogen phosphate 1.0 gm. Magnesium sulphate 0.25 gm. Carbohydrate 10.0 gm. Water 1000.0 c.c. Agar q.s. Fulmer & Grimes: Ammonium chloride 1.88 gm. Dipotassium hydrogen phosphate 1.0 gm. Calcium chloride 1.0 gm. Sucrose 50.0 gm. Agar 15.0 gm. Water 1000.0 c.c. Both formulae have been used in studies of yeasts and fermentation. Acton & McGuire (1931) report very good results with Actinomyces from the medium described by Norris (1929). It consists of: Soluble starch Dipotassium hydrogen phosphate Calcium chloride Ferric chloride Sodium nitrate Asparagin Agar Water The medium is adjusted to pH 7.4. BIBLIOGRAPHY Biltris, R. 1929. Sur la variabilite des caracteres de I'espece chez les dermatophytes, Ann. Inst. Pasteur. 43: 281-358, 15 figs. *Bizio, Bartolomeo. 1823. Lettera di Bartolomeo Bizio al chiarissimo canonico Angelo Bellani sopra il fenomeno della polenta porporina, BiMioteca Ital. o sia Giornale di Letteratura Scienze e Arti Appendice 30: 275-295 [anno 8] [translated by C. P. Merlino, Jour. Bad. 9: 527-543, 1924]. Bruhns, C. 1928. Einige Bemerkungen iiber verschiedene Pilzarten und Pilznahrboden (Griitz agar, PoUacci agar), Dermatol. Zeitsch. 53: 104-112, 4 figs. Castellani, Aide. 1933. The advisability of using in laboratory work sugars tested by micro- biological methods. Jour. Trap. Med. Hyg. 36: 185, 186. 2.0 gm, 0.2 gm. 0.05 gm. 0.01 gm, 0.06 gm. 0.05 gm. 20.0 gm. )00.0 c.c. CULTURE MEDIA 55 Coim, H. W. & Harold J. Conn. 1923. Bacteriology, a study of microorganisma and their relation to human welfare, discussing the history of bacteriology, the nature of microorganisms, and their significance in connection with pathology, hygiene, agricul- ture and the industries, Williams & Wilkins Company, Baltimore, 441 pp. Currie, James N. 1917. The citric acid fermentation of Aspergillus niger. Jour. Biol. Chem. 31: 15-37, Pis. 1, 2. Dalmau, Luz Maria. 1929, 1930. Observations on mycologic technique with particular refer- ence to pathogenic fungi, Porto Rico Jour. Public Health Trop. Med. 5: 302-311, 1930 [tr. Maurice Langeron, Eemarques sur la technique mycologique, Ann. Parasitol. Hum. Comp. 7: 536-545, 1929]. Descouraux, J. M. C. M. 1926. Contribution a 1 'etude de nouveaux milieux de culture pour les dermatophytes, These. Fac. Pharm.. Univ. Strasbourg 24: 72 pp., 2 pis. Desgardes, DeribSre. 1921. Formulaire des milieux de culture en microbiologie, Paris, Le Francois, 98 pp. Dessy, G. 1933. La chimiotherapie des mycoses. III. Mucormycose 1, Experiences in vitro, Soc. Internaz. Microbiol. Boll. Sez. Ital. 5: 95-107; 2, Experiences in vivo. Ibid. 5: 201-206. Duggar, Benjamin Minge. 1909. Fungous diseases of plants, with chapters on physiology, culture methods and technique, Boston, Ginn & Company, 508 pp. Egyedi, Henrik. 1922. Zur Eeinkultivierung der pathogenen Schimmelpilze, Centralbl. BaTct. I, 87: 562-564. Farley, David If. 1920. The use of gentian violet as a restrainer in the isolation of patho- genic molds. Arch. Derm. Syphilol. 2: 459-465, £ figs. Fulmer, E. I. & M. Grimes. 1923. The growth of yeasts on synthetic agar media, Jour. Bad. 8: 585-588. Goldschmidt, W. N. 1924. A new medium for the growth and differentiation of the dermato- phytes, Brit. Jour. Derm. 36: 204-206. Grigorakis, L. 1931. L 'action des milieux glycerines sur le mycelium degrade par le pl6o- morphisme, C. R. Soc. Biol. 108: 94-96. — . 1933. Sur un nouveau milieu de conservation des dermatophytes (plfiomorphisme, car- actfire acquis, specificite tissulaire, C. R. Acad. Sci. 196: 60-62. GriitE, O. 1923. Beitrage zur Kultur der Dermatophyten und ihrer Artunterscheidung mittels deutscher Pilznahrboden, Derm. Wochenschr. 76: 568-573. Hedges, JB. S. , 1928. Cultures of ringworm fungi on Sabouraud's proof mediums and on mediums prepared with American peptones and sugars, Arch. Derm,. Syphilol. 18: 852-856. Karrenberg, C. It. 1933. Untersuchungen iiber Wachstum und Konservierung pathogener Hautpilze auf Hirnbrei nach v. Hibler, Arch. Derm. Syphilis 168: 438-475, 10 figs. Kingery, Lyle B. 1929. Thymol and cinnamon oil in the treatment of ringworm of the scalp, Arch. Derm. Syphilol. 20: 797-805. Kingery, Lyle B. & Alva Adkisson. 1928. Certain volatile oils and stearoptens as fungicides, Arch. Dermatol. 17: 499-511, 9 figs. Langeron, Maurice & S. Milochevitch. 1930. Morphologie des dermatophytes sur les milieu naturels et milieux a base de polysaccharides (Note preliminaire), Ann. Parasitol. Hum. Comp. 8: 422-436. Levine, Max & H. W. Schoenlein. 1930. A compilation of culture media for the cultivation of microorganisms, Monographs on Systematic Bacteriology 2: xvi + 969 pp. Macleod, J. M. H. 1928. An experimental study of the Pityrosporon of Malassez: its morphology, cultivation and pathogenicity, Brit. Jour. Derm. Syphilis 40: 139-148. Nannizzi, Arturo. 1926. Eicerche sui rapporti morfologici e biologici tra gvTiinoascacee e dermatomiceti, Ann. Myc. 24: 85-129, 6 pis., 12 figs. 56 MEDICAL MYCOLOGY Pautrier, L. M. & B. Eietmann. 1924. Trichopliytie cutanee a forme d 'epidermophytie dues au Trichophyton granulosum, et provoquee par une infection de laboratoire, Bull. Soc. Franc. Derm. Syphiligr. 31: Eeunion Strasbourg, 29-31. Pollacci, Gino &) Arturo Nannizzi. 1922. I niiceti patogeni dell 'uomo e degli animali descritti, delineati e preparati per I'osservazione al luicroscopio con notizie sopra i rimedi per combatterli. 1: prefazione. Siena. Eewbridge, Allan G., Carroll William Dodge & Theodore T. Ayres. 1929. A case of menin- gitis due to Endomyces capsulatus (new species), Am. Jour. Path. 5: 349-364, Pis. 71-73. Sabouraud, Kaimond. 1908. Milieux de culture des champignons dermatophytes (Technique de fabrication des geloses sucrees dites: Milieux d'epreuve), Ann. Derm. Syphiligr. IV, 9: 99-101. — . 1925. Note sur le remplacement de tout sucre dans les milieux de culture des dermato- phytes par le miel d'abeilles, Ann. Derm. Syphiligr. "VI, 6: 515-517. Sawyer, William H. 1929. Observations on some entomogenous members of the Entomo- phthoraceae in artificial culture, Amer. Jour. Bot. 16: 87-120, 4 pis. Weidman, Fred D. 1928. Comparison of ringworm culture ingredients. II. The nitrogen factor, Arch. Derm. Syphilol. 18: 829-837. Weidman, Fred D. & Thomas M. Macmillan. 1921. A comparison of ingredients of ring- worm culture mediums with special reference to American and French crude maltose, Arch. Derm. Syphilol. 4: 451-468, 16 figs. Weidman, Fred D. & Dorothy Spring. 1928. Comparison of ringworm culture ingredients. m, Griitz, Goldschmitt, Sabouraud (honey modification), Sabouraud 's glycerin and certain synthetic mediums, Arch. Derm. Syphilol. 18: 837-850. CHAPTER IV ISOLATION OF MICROORGANISMS Transfer. — This is perhaps the simplest process connected with the hand- ling- of organisms in culture and should be practiced by the beginner until the operations can be performed quickly and entirely without contaminations before more complicated procedures are attempted. Essentially the process consists in taking a small amount of mycelium or spores of fungi or a few bacterial cells and moving them from one receptacle of medium to another. The test tube containing the culture of the organism to be transferred and a tube of medium are held about half an inch apart between the thumb and fingers of the left hand with the thumb above. The plugs of both tubes are loosened in turn Avith a rotary motion, but not withdrawn. The transfer needle, loop, or platinum spatula, as the case may be, is grasped between the thumb and first finger of the right hand, the platinum is heated to redness in a blue flame and allowed to cool while still in the hand and without the heated por- tion touching anything. The cotton plug of the tube containing the organism to be transferred is grasped between the second and third finger of the right hand and gently withdrawn, care being taken to create as few air currents as possible. A needle or loop is inserted without touching the walls of the tube and touched to the spores or to the vegetative material if it is slimy, or a platinum spatula is used to cut away a little mycelium which should adhere to it. The needle is then gently withdrawn, the plug replaced and the other plug withdrawn, and the needle stroked along the surface of the agar (or the mycelium dislodged from the spatula). The needle is then gently withdrawn, the second plug replaced, and the needle (or spatula) flamed before laying it down, to destroy any organisms still adhering to it. The plugs may then be pushed in more firmly if necessary. In some laboratories it is customary to flame and quickly extinguish the cotton plugs. Isolation. — The simplest procedure is similar to simple transfer in which the needle or spatula is touched to the infective material and then transferred to the surface of agar in a test tube or Petri dish. This method is only occa- sionally successful, usuallj^ in those cases where the infective material is small and uncontaminated by other organisms than the one which it is desired to study. If contamination is only slight, a carefully executed transfer from the colony Avliich is desired may result in securing a pure culture. Otherwise, resort must be had to dilution and plating out. Isolations From Skin Lesions. — After cleansing the skin with 95% alcohol followed by ether, the lesion is scraped with a sharp, full-bellied, sterile scalpel or shaved with a sterile safety razor to the point where bloody serum just begins to ooze. A sterile Petri dish is placed beneath to catch the scrapings, 57 58 MEDICAL MYCOLOGY or, if they do not fall away readily, they are scraped off the blade into the Petri dish by another blade. On reaching the laboratory, a few scales are removed for microscopic examination after maceration in 40% KOH or some other similar treatment. Then a very thin film of Sabouraud maltose or glu- cose agar is poured into the plate containing the scales and detritus. Thus, the latter are caught by the nutrient medium and held. To reduce chances of bacterial contamination the scales may be moistened with alcohol for a short time, but this does not prevent the growth of some of the hardy saprophytes and may inhibit the growth of the more delicate pathogen. As soon as the colonies are visible to the naked eye, the suspicious ones are marked with a glass-writing pencil, each being given a serial number. The marked colonies are then transferred to Sabouraud 's sugar medium on slants and to the conservation medium (without sugar). This transfer is made early to prevent overrunning by the rapidly growing contaminants, such as Aspergillus and Penicillium, which may be present. Isolations From Feces, Tongue Scrapings, etc, — Poured plates are allowed to harden and then 25 points of contact are made in each with a platinum loop repeatedly soiled with the infective material. After about four days, when the colonies have developed, these are fished. In this way the percentage of points of contact of material with the medium which contains similar colonies gives a rough idea of the abundance of colonization in the material. (Dalmau 1930.) In connection with their studies with sprue, Weiss & Landron (1928) sug- gest as follows: Emulsify a small quantity of feces in a test tube of sterile distilled water and also in another tube containing whole ox bile in which has been incorporated 20% concentration of glycerol. By means of a glass rod bent at an angle of 30° a drop of each fecal suspension is spread over the surfaces of the plates of Sabouraud agar (pH 6.3) containing 4% glucose and 20% glycerol ; incubate at 35° C. Dilution. — About three tubes of agar (or other liquefiable medium) are heated gently on a water-bath (porcelain or glass beaker) until the agar melts. It is then allowed to cool until the end of the tube containing the agar can be held against the back of the hand without causing pain, i.e., until the solidifying point is almost reached. The tube is inoculated by the method suggested in simple transfer. After laying down the needle, the tube is rapidly rolled between the palms, while it is in a vertical position, to mix the contents thoroughly and scatter the inoculum. The loop is used to transfer a tiny drop of the molten medium to the second tube two or three times. This in turn is rotated, etc. The contents of the three tubes are then poured successively into three Petri dishes lying on a level surface. If the agar fails to wet the surface of the dish completely, the dish is tipped slightly to allow the agar to flow over the whole surface of the bottom. It is then allowed to solidify and is incubated bottom side up until growth is evident. Then individual colonies are transferred to slants. Sometimes it is necessary to repeat this process. In the above-mentioned processes it is desirable to work gently in ISOLATION OF MICROORGANISMS 59 order to set up as few air currents as possible, never to lift the cover of the Petri dish higher than necessary to insert the test tube for pouring, and to keep the test tubes plugged as much as possible. As soon as the agar is poured from the tubes, they should be lowered into a dish of water and boiled as soon as possible, both to kill the organism which may have adhered to the agar still in the test tube and to clean them before the agar has a chance to dry on. Inhibitors. — To keep back the rapidly growing organisms and allow the slower growing ones to develop, various substances may be added to the medium first used in isolation. Advantage is often taken of the fact that some groups of fungi grow at different hydrogen ion concentration from others. In these methods, varying small amounts of lactic or other organic acids are added to the medium to inhibit the growth of bacteria. Sometimes dyes are also used for this purpose, e.g., "gentian violet" (probably methyl violet) 1:500,000 (Farley 1920) or to indicate the presence of a small colony before it has developed sufficiently to be seen otherwise in order that it may be fished before it has been overgrown by a more rapidly growing organism. Indicators are very useful in this way if the organism one desires to isolate produces acid or alkali in the medium. Usually these special methods have been developed in connection Avith an intensive study of a single organism and are rarely useful unless the presence of a given organism is strongly suspected. Slight amounts of organic acids are useful, however, in keeping down rapid bacterial growth while waiting for the more slowly growing fungi to develop. Similarly, the choice of suitable media may do much to favor selectively the development of one organism while retarding another. No general rules can be given for these choices since they are largely the result of wide experi- ence and a shrewd guess as to the probable organism to be isolated. A thor- ough knowledge of the physiology of the various groups of fungi will be helpful, but in the present state of our knowledge generalization is very difficult. Microcultures. — In the study of the life cycle of many organisms it be- comes desirable to have a given spore or bit of mycelium under more or less continuous observation with the microscope. One of the early methods which has yielded much useful information is the hanging drop culture. A.n early and inexpensive form, usually referred to as a Van Tieghem cell, consists of a glass ring cemented to a microscopic slide with wax (made by melting to- gether pure beeswax and vaseline). The top of the ring is coated with vase- line. A drop of the culture medium or water is placed in the bottom of the cell thus formed. Another smaller drop is placed upon a clean cover slip of sufficient diameter to cover the ring. The inoculum is then placed in the center of this drop, and the whole seized by forceps and quickly inverted, care being taken that the drop does not spread too near the edge of the cover dur- ing the process. The cover (with the drop of medium or water hanging from it) is then lowered to the glass ring and pressed down gently until the soft vaseline (petrolatum) seals it to the ring. Thus we have a small moist chamber with the organism suspended in a drop from the cover. The drop placed in 60 MEDICAL MYCOLOGY the bottom of the cell prevents drymg out, since its vapor pressure is prac- tically the same as that of the hanging drop. The cell may be placed on the stage of a microscope and studied from time to time until the nutrient material in the drop is exhausted. One should be careful not to have the drop too large, since it will be difficult to focus to the bottom of it if the spore under observation should not be thoroughly wetted and lies in the surface layer of the drop, or is so heavy that it falls to the bottom of the drop. To avoid this, sometimes a thin film of agar is used instead of a liquid drop, in which case distilled water is usually placed in the bottom of the cell to prevent drying out. Sometimes a very young colony with a small amount of the surround- ing agar is cut out and placed on the cover glass, making a hanging block culture described by various authors including Dalmau (1929-1930). Studies made by these methods are especially valuable in following the early stages in the development of a spore or in the evolution of the life cycle. Various types of hollow-ground slides, etc., have been developed, but for convenience and general use, they have little advantage over an ordinary Van Tieghem cell. Giant Cultures. — At the other extreme from the microculture, we may use giant cultures. These colonies which are allowed to develop over a long period on an abundant supply of substrate, are very useful in giving gross morphology on various media and often are strikingly characteristic in ap- pearance for different species. They have had little favor in most labora- tories, as they must be allowed to grow from one month to two or three before this character can be ascertained. However, they have been utilized with very excellent results by Sabouraud and others in the dermatophytes and by Lindner in the yeasts. Since, under ordinary conditions a Petri dish is too shallow to hold sufficient medium, and allows it to dry out too readily and also to be subject to contamination, various specialized culture dishes have been devised. Perhaps the Roux flask, of which there are several types on the market, is the best known, although somewhat expensive. Ordinary cultures in Erlenmeyer flasks are satisfactory for most purposes, but they do not admit of either microscopic examination or photography without breaking the flask, which is frequently difficult to do without injury to the colony. To obviate this for photographic purposes, Shrewsbury (1931) suggests growing them in green glass medicine bottles of 10-12-ounce capacity (with flat sides if possible). The bottle may be evenly broken by application of a red hot nail along the sides. These bottles are also useful for growing cultures under diminished pressure since the glass is strong enough to resist almost complete exhaustion. He also suggests their use in exposing the inverted colony to fumes of various antiseptics, such as ethyl iodide. Karrenberg has suggested two ingenious devices which permit giant colonies to be photographed or studied with the low power of the microscope with a minimum chance for contamination. In 1926 he suggested an inner test tube carrying an agar slant from which the side had been removed. This was stored in a somewhat larger test tube plugged with cotton. When it is desired to photograph or to examine the colony, the inner tube may be easily ISOLATION OF MICROORGANISMS 61 removed and if care is taken to work in a relatively dust-free atmosphere, often several examinations may be made before the colony becomes contami- nated. The size of the colony is rather limited in this device. In 1927 he proposed a more elaborate flask. It is essentially a flask of the Erlenmeyer type, made in two pieces, a lower portion to hold the medium and a cover fitting over it rather more closely than the usual Petri dish cover. The pieces are held tog-ether by a metal plate underneath and a ring around the neck of the flask connected by three springs which hook into the upper ring to provide sufficient compression to prevent contamination. The flask is manipulated as an ordinary flask, the neck being plugged with cotton. After the colony is grown, the springs are unliooked from the ring and the top is lifted off. If care is taken in the examination, little contamination results. So far as I am aware, this type of flask has not been placed on the market. Sing^le Cell (Spore) Cultures.— Finally, there comes a time in the study of many fungi when the results of a study of cultures made by the above- mentioned methods are ambiguous, and it becomes desirable to work with mycelium and spores produced by a single spore in order that problems of sexuality or homothallism and heterothallism may be investigated, or that the relationships of apparent stages in a life cycle may be studied and verified. The problem has been variously met by different investigators, depending somewhat on the size and nature of the spore to be isolated and the instru- ments available for the work. If the spores are large and the hand is very skillful, it may be possible to pick up a single spore on a needle under the low power of the microscope and transfer it to a sterile tube or a hanging drop. Various mechanical devices have been developed to aid in this work, e.g., the old Barber spore picker or pipette or some of the modern micro- manipulators used in microdissection studies. In these devices, motion is secured by means of micrometer screws, and the spore is usually sucked into the end of a tiny pipette made by drawing out a piece of glass tubing to an inside diameter only slightly larger than the spore or cell to be isolated. This is then discharged into a hanging drop or a sterile culture tube. For detailed directions, those accompanying the instruments should be consulted. Ascospore Detection. — Since classification is based primarily upon the spore forms resulting from the sexual act (caryogamy), every effort to secure the sexual or perfect stage should be made before relegating an organism to the large heterogeneous group known as the Fungi Imperfecta There is no single method which is equally successful for all organisms, nor even for mem- bers of a single group. Perhaps patience is the first requisite. Frequently one finds evidence of sexuality by a diligent search of a colony 3-4 months old, after the agar has begun to dr5\ This seems very useful in the filamentous yeasts whose im- perfect stage is usually placed in the genus Monilia. Some media seem better than others, but usually a careful search will show traces of sexuality on most media upon which the organism has made a good growth. This method is very tedious, especially among pathogens when the physician wishes a prompt 62 MEDICAL MYCOLOGY diagnosis, at least to a genus name, and seems puzzled if the mycologist promptly reports that he has a species of Monilia and 2 or 3 months later changes his report to Zymonema. Among the true yeasts, several methods are in vogue, none of which is uniformly successful, but all of which may sometimes produce results. All should be tried before placing an unknown organism in the large and poorly defined genus Cryptococcus. Stelling-Dekker (1931) has summarized these methods and in most cases traced the method to the original author. The oldest method is that of Engel (1872), popularized by Hansen (1883), where the cells from an actively growing colony are scraped off and placed on a sterile block of plaster of Paris (gypsum) which is kept moist by sterile water, or malt extract (Klocker 1924) or by mannitol-phosphate solution (18 c.c. 2% mannitol + 2 c.c. 5% dipotassium phosphate) (Saito 1923). Gorodkova (1908) reported the use of an agar rich in nitrogen and poor in carbohydrate which usually bears her name. Distilled water 1000 c.c, agar 10 gm.,* peptone 10 gm.,* beef extract 10 gm., sodium chloride 5 gm., and glucose 2.5 gm. Various French authors, notably Guilliermond, have advocated the use of slices of carrot or potato. Beijerinck(1898) advocated the use of plain agar to which no nutrient had been added and which had been thoroughly washed to remove impurities which might possibly be a source of food. Wagner (1928) has made a more thorough study of conditions initiating ascospore formation. He emphasizes the importance of the sugars previously used in cultivating the organism and the hydrogen ion concentration of the medium. Kufferath (1928) attributes the success of his medium to its alka- linity. He prepares it as follows : Malt meal is hydrolyzed with sulphuric acid, the acid neutralized with calcium carbonate, and the agar added. It is then brought to the desired alkalinity with sodium hydroxide. In 1930 he studied the matter further. He found that in general the usual concentration of gelatin (15%) is as successful as higher concentrations. He studied the effect of alkalinity and found it rather more successful than acidity in producing ascospores, but occasionally the reverse is true. Before one can be certain that spores are not formed, one should try all the methods. Fennentation. — Another character to which some authors have attached much importance in some groups is ability to ferment or to utilize certain sugars. The term "fermentation" is used very loosely by various writers. Some, as Stelling-Dekker, would practically restrict it to the production of alcohol and carbon dioxide from a hexose, while Castellani would include all cases in which acid or gas appears in a carbohydrate-containing medium on which an organism has developed. It is probable that this different use of the term has occasioned much difference of opinion in regard to the fermenta- tive ability of a species. A further source of error is almost inherent in the methods in ordinary use, each of which indicates an equilibrium of several possible reactions, hence it is important to state clearly what method was used •Maneval (1924) recommends the omission of peptone, addition of 15 or 20 gm. agar, and reduces Liebig's meat extract to 3 gm. ISOLATION OF MICROORGANISMS 63 in determining fermentation if this character is to have meaning in the separa- tion of species. In the following discussion of methods, an attempt will be made to point out some of the sources of error and objections raised to each method. As in many other organic reactions, details of method often pro- foundly influence the point of equilibrium. Lindner's Microfermentation Method. — A hollow-ground microscopic slide is flamed and filled with sterile water. A relatively large number of yeast cells is suspended in the water and a pinch of the sugar to be tested is added. A cover glass is carefully lowered so as to exclude any air bubbles and sealed in place with lanolin or vaseline. This is placed in an incubator for 24 hours at the optimum temperature and the presence of bubbles (supposedly of car- bon dioxide) noted. In this method it is assumed that there will be com- paratively little growth and consequent production of CO2, due to absence of oxygen and the lack of nutrients. The amount of gas should be roughly proportional to the amount of the inoculum. If the organism ferments very slowly and a larger quantity of inoculum is used, the amount of glycogen trans- ferred with the yeast ceUs and that diffusing out of the dead cells may be sufficient to give gas production. Since the sugar is not sterilized, there is always a possibility of introducing some fermenting bacterium. Also there is a possibility that the seal is not tight and that subsequent evaporation may give the appearance of gas production or may allow the gas to escape. Changes of temperature or of atmospheric pressure may also give erroneous results. Guilliermond modified the method by filling a Van Tieghem cell with a rela- tively large volume of liquid, and dissolving the sugar in a definite concentrar tion in a yeast decoction. This modification presupposes that the yeast may grow, and consequently the COg may be partly the result of respiration rather than fermentation in the strict sense of the word, especially as there may be dissolved oxygen in the liquid. The Fermentation Tube Method. — In this method, bent tubes of varying pattern, perhaps originally used by Einhorn in 1885 but introduced into mi- crobiology by Theobald Smith in 1890, are filled with a sugar solution and sterilized. The amount of liquid should be sufficient to slightly more than fill the long arm but not so much as to wet the plug. This is inoculated with the organism and set aside in an incubator until the organism has grown long enough to develop gas. If the long arm is graduated, the volume of gas pro- duced may be noted. Since the surface of the liquid in the long arm is not exposed to free oxygen, strict anaerobic conditions are maintained. Also, if the organism is strictly aerobic, it grows only in the short arm, and there is not enough growth in the long arm to show fermentation, even if the organ- ism is capable of producing it. On the other hand, since the carbon dioxide is quite soluble in the solution, small amounts are dissolved and do not show up. There is also the problem as to the source of the gas, whether it is from fermentation or respiration. Aichelburg (1932) reports that gas production in fructose shows on the third day while glucose shows on the first day. 64 MEDICAL MYCOLOGY The ideal method would be one in which the reaction was studied quan- titatively in special apparatus, a condition not attained except in purely physiologic researches (cf. Kluyver 1914). Since comparatively few organisms ferment hexoses to carbon dioxide and alcohol, in the majority of cases we are really interested rather in the ability of the organism to attack and utilize sugars than in their fermentative ability in the narrow sense of the term. Hence, a quantitative titration of the sugar medium for the presence of accumulated acid is equally useful. In fact, in the majority of "fermentations" mentioned for Monilia by Castellani, the production of acid rather than of alcohol is meant. Probably this has been done very roughly, since the author rarely mentions the indicator used and never the relative amount of sugar converted to acid in a unit of time by a definite number of organisms per cubic centimeter. Stelling-Dekker sug- gests that more quantitative methods are useful, especially in the case of a trisaccharide in which several organisms secrete raffinase which by hydrolysis splits the rafSnose to mellibiose and fructose and is able to ferment the fructose so produced but not to hydrolyze the mellibiose. Aichelburg (1932) reports that slight acidity shows after one week with inulin but only after two weeks with starch. For further consideration of the problems involved, especially of the chemical reactions, the reader should consult some of the standard works on biochemistry, or special monographs on alcoholic fermentation, such as that by Harden. Maltose, fructose, and glucose are quite regularly fer- mented by many species of yeasts while galactose, sucrose, and dextrin are more variable. Inulin, raffinose, and mannite are often attacked by certain species and should be tried in placing a strange Monilia. Lactose is attacked by comparatively few fungi, but when it is attacked the amount of fermenta- tion is apt to be large. Most of the other sugars are too rarely fermented and too expensive to be used in most routine work. Ashford has shown that ultraviolet light may destroy the normal fermenting power of Syringospora psilosis (Monilia psilosis). Sometimes characteristic deposits may be evident upon cultivation in connection with fermentation studies. Ashford 's laboratory usually tests fer- mentation on 1-4% concentration of the sugar in peptone water, although nutrient bouillon may be used. The pH is adjusted to about the neutral point, and changes of acidity or alkalinity are noted. Animal and Human Inoculations and Recovery of Organisms From Lesions. — The methods used for inoculations, both of animals and human volunteers, are too well known by the medical profession to need discussion here, and have little value in the hands of an experimenter without a good medical training. The necessity of a thorough sterilization of the skin cannot be too strongly emphasized, for mold spores from dust or clothing may be picked up as a contaminant and, in some cases, even be considered as the etio- logic agent. Perhaps the advice of Erwin F. Smith needs especial emphasis in this connection. "I now endeavor to repeat all my own experiments several times over and in the end I have a rounded out and better view than the one ISOLATION OF MICROORGANISMS 65 series only could possibly give me. Incidentally, I usually succeed in eliminat- ing some errors or half truths which appertained to the first experiment." BIBLIOGRAPHY Beijerinck, M. W. 1898. tJber Eegeneration der Sporenbiklung bei Alkoholhefen wo diese Funktion im Verschwinden begriffen ist, Centralbl. BoM. II 4: 657-663, 721-730, 1 pi. Benedek, Tibor. 1926. tJber directe mikrospische Beobachtung voa Eeagensglas Pilzkulturen und ihre Verwendungs mogliclikeiten, nebst Angabe von neuen Mikroskop Objekt Tisch-Klemmen fiir Mykologische Arbeiten, Derm. Woch. 82: 1-5. Dalmau, Luz Maria. 1929, 1930. Observations on mycologic technique with particular refer- ence to pathogenic fungi, Porto Eico Jour. Public Health & Trop. Med. 5: 302-311 [tr. M. Langeron, Remarques sur la technique mycologique, Ann. Parasitol. Hum. Corny. 7: 536-545, 1929]. Dekker, N. M. Stelling. 1931. Die Hefesammlung des " Centraalbureau voor Schimmelcul- tures. " Beitrage zu einer Monographie der Hefcarten. I. Die Sporogenen Hefen, Verhandel. K. Akad. Wetensch. Amsterdam. Afdeel. NatuurT:. 28:1; 1-547, illus. Engel, L. 1872. Les ferments alcooHques, These Paris 336: 1-62, 1 pi. Farley, David L. 1920. The use of gentian violet as a restrainer in the isolation of pathogenic molds. Arch. Derm. Syphilol. 2: 459-465, 2 figs. Gilchrist, T. Caspar & William Eoyal Stokes. 1898. A case of pseudolupus vulgaris caused by Blastomyces, Jotir. Exp. Med. 3: 53-78, Fls. 4-8. Gorodkova, A. A. 1908. O bystrom poluchenii spor u drozhzhevykh gribov, Bull. Jard. Imp. St. Peterslourg 8: 165-170. Grigorakis, Leonidas. 1933. Sur un nouveau milieu de conservation des dermatophytes pleomorphisme, caractere acquis, specificite tissulaire, C. E. Acad. Sci. 196: 60-62. Hansen, Emil Christian. 1883. Unders0gelser over alkoholgjaersvampenes fysiologi og morfologi, Meddelelser Carlsherg Laboratoriet 2: 29-102; 152-210; 220-256 [in French 13-59; 92-136, 142-157]. Ges theor. Abhandl. u. Garungsorganismen, 125- 169, 1911. Karrenberg, Karl Ludwig. 1926. Ein neues Rohrchen der Ziichtung von Dermatophyten und Bakterien, Derm. Zeitschr. 49: 248-252. — . 1927. Ein neuer Kolben zur Ziichtung und Aufbewahrung von Pilzkulturen, Derm. Woch. 85: 1706-1708. — . 1927. Die Eignung der von Griitz angegebenen Nahrboden zur Bestimmung und Weiter- ziichtung der Dermatophyten, Derm. Woch. 84: 434-439. — . 1933. Untersuchungen iiber Wachstum und Konservierung pathogener Hautpilze auf Hirnbrei nach v. Hibler (Vorlaufige Mitteilung, Arch. Derm. Syphilis 168: 438-475, 10 figs. *Klocker, A. 1924. Die Garungsorganismen ed. 3. *Kluyver, A. J. 1914. Biochemische Suikerbepalingen. Delft. *Kufferath, H. 1928. Ann. Soc. Zymologie 1: 214. * — . 1930. Ann. Soc. Zymologie 2: 33. Mallinckrodt-Haupt, Asta St. v. 1926. Vitalfarbung niit Indicatorfarben bei Hyphomyceten, I, Derm. Zeitschr. 46: 293-305. Maneval, W. E. 1924. A method of securing spores of yeasts, Bot. Gaz. 78: 122, 123. Saito, Kendo. 1923. Beschreibung von zwei neuen Hefearten nebst Bemerkungen iiber die Sporenbildung bei Torulaspora, Delbriicki Lindner, Bot. Mag. Tolcyo 37: 63-66, 2 figs. Shrewsbury, J. F. D. 1931. Giant colony culture, Jour. Path. Bact. 34: 283-285, PI. 15. Wagner, Felix. 1928. Der Einfluss der Zuckerarten und der Wasserstoffsionenkonzentration auf die Sporulation der Saccharomyceten, Centralbl. BaM. II 75: 4-24, 5 figs. Weiss, Charles & Francisco Landron. 1928, 1929. Immunological investigations of tropical sprue in Porto Eico, 1-3. Am. Jour. Trop. Med. 9: 83-95, 1929. 4. Biology of Monilia psilosis in relation to sprue, Jour. Infect. Dis. 43: 557-564, 1928. CHAPTER V MICROSCOPY BY MORRIS MOORE To one acquainted with the cultural characteristics of various groups of fungi, it is easy to recognize the larger groups, but for accurate diagnosis a microscopic study of the morpholog}' of the organisms is necessary. The fungus must be allowed to grow for a sufficient length of time to permit im- portant morphologic characters to develop. If the organism is pathogenic, all the precautions mentioned for transfer must be taken to insure that spores of dangerous microbes are not detached to float in the air and perhaps inoculate some one. The slide and cover should be thoroughly cleaned with acid alcohol and passed through a flame to remove any traces of fat. In dislodging the material to be studied, great care should be taken not to entangle unneces- sarily the mycelium. With yeastlike fungi this latter step is simplified, since it is necessary only to take a loopful of the culture without fear of entangling the mycelium as may occur with filamentous forms. The mounting of the material should be done carefully. A drop of dis- tilled water, alcohol, Amann's lactophenol preparation, or glycerin, either with or without a stain, is placed in the center of the slide. The fungus is then lifted carefully from the tube with a platinum or nichrome needle or spatula, dislodged into the drop, or pushed oft* by another needle if not pulled off by the surface tension of the mounting medium. Platinum is preferable to nichrome because of its rapidity in cooling, but nichrome is a harder metal and is better for thick, hard growths which resist elevation by the needle. The material is spread out as gently as possible in the mounting medium and a cover glass is lowered carefully on the preparation, avoiding the inclusion of bubbles of air. Alcohol has the advantage of killing the organism, and, hav- ing a low surface tension, does not dislodge spores badl3^ It also has a tend- ency to form fewer air bubbles, but it soon dries out and must be replaced quite promptly by Avater or water and glycerol (2 parts water and 1 part glycerol). This is done by placing a drop on the slide next the cover glass and allowing it to be drawn in under as the alcohol evaporates. Care must be taken, if glycerol is used, that it does not wet the top of the cover glass, as it will be difficult to remove later. The disadvantage in the use of glycerol alone lies in the fact that yeastlike or even filamentous forms may be cleared to such an extent that it is \ery difficult to make out the morphology of the organism. To avoid this, various dyes, such as methylene blue, crystal violet, or eosin, are incorporated either as an aqueous or alcoholic solution (usually about 1%) in amount sufficient to produce the desired intensity. Water does not evaporate rapidly, but, owing to its high surface tension, tends to tear 66 MICROSCOPY 67 away spores from their attachments and, in general, is rather unsatisfactory for filamentous fung-i, although it is very satisfactory as a mounting medium for most 3^east and yeastlike organisms. Some of the various formulae of laetophenol give good results, as does also lactic acid alone. The latter has the disadvantage of preventing the use of many dyes in staining. So far as I am aware, laetophenol was developed in French laboratories, the formula of Amann (1896) being phenol crystals 20 gm., lactic acid 20 gm., glycerol 40 gm., and distilled water 20 gm. These are dissolved with gentle warming and then is added anilin blue (a mixture of the tri-sulphonates of tri-phenyl pararosanilin [C.I. 706] and of di-phenyl rosanilin) otherwise known as cotton blue (C. B. Poirier). Other compounds, such as methyl blue, are also called cotton blue, but are said to be distinctly inferior for this purpose. Sartory (1924) recommends 0.5% dye to his laetophenol. Linder (1929) ad- vocates the same formula while Henrici (1930) adds only 0.05% of the dye. I have found that the dye added directly to the laetophenol gives a blue back- ground to the preparation. Consequently, I use a 1% aqueous solution of cotton blue, place a drop on a clean slide, inoculate with the fungus, lower a cover glass on the mount, and then allow a drop of laetophenol to be drawn in under as shown previously for glycerol. By this method, the excess dye is pushed to the edge of the cover slip and the laetophenol forms a clear back- ground for the blue fungus. Weston (1929) recommends the addition of a small quantity of nigrosin, water soluble, either aqueous, or the picric acid solution described by Curtis and Colley (1915) in order to stain nuclei as well. Since the dye varies in different samples, at present Weston has found no other way than to add some of the dye, try it, and then add more dye or more laetophenol until satisfac- tory results are obtained. Sartory also recommends a mixture of Sudan III 1 part, and lactic acid 1000 parts by weight. This is ground in a mortar with slow additions of small amounts of the lactic acid. The mixture is then heated in a flask on a water-bath until it is completely dissolved, cooled and filtered. One part of anilin blue is added, also 1-2 drops of tincture of iodine for each 10 c.c. of solution. This triple stain colors fatty bodies, amyloid compounds, and the fungus protoplasm. Before adding the cover slip the mount should be heated gently until vapors are given off. Spore Stains. — Maneval (1924) suggests the following stain for yeast spores. Spread a film of cells in a drop of water on a slide and dry in air, fix by passing through a flame 12-15 times; stain with hot carbolfuchsin 1-3 minutes; wash with water; destain with 5% sulphuric acid 2-3 seconds; wash with water and stain with methylene blue 3 seconds, Avash with water. In 1929, he suggested the following procedure : stain with carbolfuchsin or carbol-methylene blue; destain with 5% acetic acid; treat with 5% tannin for 2 minutes; wash and counterstain with methylene blue (after carbolfuchsin) or with safranin (after carbol-methylene blue). Old spores should be heated in a small amount of sterile water for 10 minutes on a hot water-bath before 68 MEDICAL MYCOLOGY making the smear. Hufschmitt, Sartory and Meyer (1931) advocate the Moeller method which is slightly different from Maneval's procedure. Treat smear from 10 seconds to 5 minutes in 1% sulphuric acid; wash, stain with carbolfuchsin, heating for 1 minute; differentiate with 5% sulphuric acid for 5 seconds ; wash, counterstain with aqueous methylene blue for 3 minutes. Buschke and Harry (1923) recommend the Schumacher method. Fix, stain for 1 minute in carbol-methylene blue, rinse with distilled water, stain for 11/2 minutes, while slowly moving the slide, with 1% phosphin (diamido- phenylacridin). Spores also stain with Ziehl-Neelsen acid-fast procedure if the sulphuric acid is replaced by 1% nitric acid alcohol. Maneval (1929) suggests the following modification of Gutstein's pro- cedure for staining vegetative cells. Fix smear with heat, stain with 5% tannin for 2 minutes, then with safranin or 1% methylene blue, or stain with carbol-methylene blue (5% carbolic acid plus 1% methylene blue) or methy- lene blue; treat with 5% tannin for 2 minutes, wash, and counterstain with safranin. It is often very easy to find spores just by making mounts in glycerin and using some dye, such as crystal violet, or lactophenol preparations. Most of the dye preparations will stain vegetative mycelium. Stains for Fungi in Skin. — Unna, Jr. (1929) advises the following modi- fication of the Pappenheim-Unna, Sr. method for staining fungi in skin. Fix in absolute alcohol, then run through the alcohols to xylol, and embed in paraffin. Cut sections about 10/a thick; stain with pyronine-methyl green (pyronine 9 parts, methyl green 1 part, 96% alcohol 90 parts, glycerol 100 c.c, 0.5% phenol to make 1000 c.c.) for 5-10 seconds; rinse in water; dry with absolute alcohol, and mount in balsam. The fungi will be rubin red, leuco- cytes green to blue green. (N.B. The cells of the basal homy layer of the epi- dermis will have red nuclei by this method.) Fungi in tissue can be stained easily by the usual iron-alum hematoxylin and eosin procedure. The fungus elements take the hematoxylin stain rather nicely, although some difficulty may be encountered in distinguishing spheri- cal cells or spores from tissue elements. The Gram method of staining for bacteria has been used with a measurable amount of success since fungi are, in general, gram-positive. The formula of Malcolm Morris (Mallory and Wright 1924, p. 175) for staining various parasites of the skin avoids the use of hydrate of potash. The skin is placed in ether or in a mixture of alcohol and ether, equal parts, stained for 5-30 minutes in a solution of 5% gentian violet in 70% alcohol; iodine solution, 1 minute ; anilin, or anilin plus 2-4 drops of nitric acid ; anilin ; xylol ; xylol and balsam. Stains for Fungi in Other Tissues. — A number of methods listed in Mal- lory and Wright (1924) for staining bacteria in tissue, as well as various types of cells, have been used successfully with fungi. Mallory 's anilin blue stain (p. 118) has been used very nicely for Cryptococcus histolyticus (Torula histolytica) in brain tissue. The Gram-Weigert staining method (p. 288) is in MICROSCOPY 69 general use. On p. 414, the authors list two methods for staining Actinomyces in sections, although alum-hematoxylin followed by a strong eosin solution will give good results, as will also the Gram method for paraffin sections, which is as follows : Stain in anilin-methyl violet for 5-20 minutes ; wash in normal salt solution or water; iodine solution (1:2:300) 1 minute; wash in water; absolute alcohol, several changes, until no more color is given off and the section is ap- parently decolorized; xylol; xylol and balsam. The so-called "clubs" do not stain with Gram's stain while the central portion of the granule, the thin filaments, do. Stains for Hair and Scrapings. — Adamson (1895) recommended clearing with 5-10% KOH and staining by the Gram method. Chalmers and Marshall (1914) suggest soaking scales in 40% KOH for some hours in a w^atch glass in an incubator at 40° C. Transfer specimens to watch glass containing 15% alcohol for 30 minutes, remove to slide, allow alcohol to evaporate, and dry over flame; stain with anilin-gentian violet for 30 minutes. Treat with Gram's iodine solution for 3 minutes ; decolorize with anilin oil for 30 minutes ; stain in concentrated alcoholic eosin for 1 minute ; wash off eosin with anilin oil or clove oil ; treat with xylol, and mount in balsam. Priestley (1917) recommends lactophenol (lactic acid 1 part, phenol 1 part, glycerol 2 parts, water 1 part) for clearing instead of 40% KOH; or chloral hydrate crystals 2 parts, lactic acid 1 part, phenol crystals 1 part, may be used. For staining he recommends treatment with chloroform to remove the fat; boil for 2-3 minutes with formic acid; wash for a few minutes in water, stain with Sahli 's methylene blue ; wash ; differentiate with alcohol, if necessary; dehydrate, and mount in balsam. Bachman (1920) recommends the following procedure : Place scrapings in a drop of water on a cover slip, tease thoroughly with a dissecting needle, dry over a flame but do not scorch. Stain for 2 minutes; decolorize in 95% alcohol for 15-30 seconds; immerse in distilled water 15-30 seconds; pour off excess, dry by heat, and mount in balsam. The spores and mycelium will be blue, the scrapings yellow. His dye is made as follows: saturated alcoholic gentian violet 2.5 parts, distilled water 17.5 parts, orange G solution 9 parts, acetic acid 1 part, 95% alcohol 5 parts. His orange G solution is orange G 2 parts, 95% alcohol 20 parts, water 80 parts. Decolorize with 10-20% KOH. The host is not stained, the fungus appears yellowish red. Unna's method is to rub the scales of the epidermis in a little glacial acetic acid between two slides. These are drawn apart and quickly dried over a flame. The fat is removed by means of alcohol and ether, and the preparations are stained in borax-methylene-blue. Instead of these slightly complicated methods, I have found that infected hairs can be cleared sufficiently to show spores and mycelium by mounting in a hydroxide solution, sodium or potassium, 10-30%. Sodium hydroxide is not quite as satisfactory as potassium hydroxide and a 20% solution works with sufficient rapidity to give good results without seriously macerating the hair. The hair may be immersed in ether to dissolve off' oil or fat, and slight heating 70 MEDICAL MYCOLOGY of the preparation may speed up the reaction. Skin scrapings are also cleared satisfactorily by this simple method. Henrici advocates a 25% solution of antiformin, such as is used in digesting- tuberculous sputum, in place of the sodium hydroxide solution. Stains for Sputum, Pus, etc. — Fungi may be found in sputum, pus, or exudates by making mounts in 20% potassium hydroxide, or by smears. The latter is not very satisfactory except for indicating the presence of mycelium or cells, since smearing tends to disturb the arrangement of the cells. There is usually a great amount of contamination unless one is able to open a fresh lesion which shows no fistulae or draining sinuses. Where the latter are present, biopsies are necessary to find the suspected fungi. Since in exudates the or- ganisms may be few in number, it may require several examinations to locate the parasite. The hydroxide tends to dissolve most of the tissue elements and leave the fungi as refractile bodies. In potassium hydroxide preparations of scales from inflammatory lesions Becker and Ritchie (1930) have indicated artefacts which rather closely re- semble yeast cells. These bodies vary in shape from spherical to ellipsoidal and have a highly refractive wall of varying thickness. Even appearances of sprouting and septal formation occur. These may be removed by treating material progressively with absolute alcohol, ether, absolute and 95% alcohol. The appearances known as mosaic fungus, reported by Greenwood and Rock- wood (1930) to be degenerate hyphal cells, is suggested by Becker and Ritchie (1930) to be a result of inflammatory changes in the tissues. Vital Staining of Fungi. — Dalmau (1929, 1930) reports a technic of vital staining which, while not original with her, deserves much wider use than it receives at present. It is essentially similar to some of the blood stains. The slides should be new, free from blemishes, and should be freed from fat by the use of cleaning solution. After neutralizing, they should be cleaned with a fat-free cloth. Immediately prior to use, all dust must be removed with a new camel's hair brush, washed with ether, and dried. On one slide place one or two drops of Janus green, neutral red or Scharlach R solutions (1 :2500) in ethyl alcohol and let them extend over the entire slide, which they Avill do if the latter is fat-free. Dry in the air. Place a drop of the liquid medium containing the fungus upon a cover slip and invert upon the slide containing the stain. Let settle and then rim the edges with vaseline. Examine at intervals. Dalmau (1930) reports successful staining of fixed material with the com- mon blood stains (Wright's, Giemsa, and Leishman). With a platinum loop mix some of the colony with a drop of clear blood serum, placed at the end of the slide. Before it dries spread it gently with another slide, thus produc- ing a thin film. Fix with methyl alcohol from 1 to 3 minutes. Stain with the above-mentioned blood stains and use neutral water for washing or diluting the stain. The stain should remain from 5 to 15 minutes. Differentiate for a few seconds only with acetic acid (1:1000). Wash wftll. The addition of serum prevents undue shrinkage of the cells. MICROSCOPY 71 Fixing Agents. — It is at times desirable to kill and fix material to prepare the fungus for staining and clearing. A number of fixatives are used, but most authors recommend either Plemming's weak killing agent which con- sists of two solutions which are mixed when ready to fix the material (A. 1% chromic acid 25 c.c, 1% acetic acid 10 c.c, water 55 c.c. ; B. 1% osmic acid 10 c.c.) or Flemming's strong agent (A. 1% chromic acid 45 c.c, glacial acetic 3 C.C; B. 2% osmic acid 12 c.c). Ninety-five per cent alcohol is used, but it is not so suitable for fine work since the material is frequently plasmolyzed. There are various chromo-acetic acid formulae used, but the reader is referred to the standard books on methods in histology for these. One of these for- mulae, Benda's fluid, has been used favorably with yeastlike fungi. It is a modification of Flemming's strong agent and works well for chromatin inves- tigations. One per cent chromic acid 16 c.c, 2% osmic acid 4 c.c, and glacial acetic acid 2 drops. One of the best, yet most expensive fixing agents I have found for celloidin sections of pathogenic fungi is Hermann's fluid : 1% platinic chloride 15 parts, glacial acetic acid 1 part, 2% osmic acid 2 parts. Fix for 6-12 hours; wash overnight. MerkeFs fluid gives good results in organs filled with reserves of food materials. ParaJSn Method. — There are two general methods for embedding material for cutting sections : the paraffin and the celloidin technic Recently, modifi- cations have been reported, but they have not been sufficiently tested to report here. The paraffin method is familiar to practically all technicians and is in general use, although not very satisfactory for agar cultures. The diffi- culty seems to lie in the fact that it may cause shrinkage of the material, and it does not penetrate the agar sufficiently for good embedding. Masses of mycelium scraped from the surface of the agar and embedded in paraffin may give favorable results, but it is often desirable to see the characteristics de- veloped in the substrate. Nitrocellulose Method. — The outstanding method of embedding agar cul- tures is the second procedure. Although its particular disadvantage lies in the fact that it is difficult to obtain very thin sections as with paraffin and also more difficult to make serial sections, although possible, its advantages surpass those of paraffin. Material if fixed properly will show no shrinkage. The agar substrate is penetrated much better, so that celloidin blocks can be made easily and sections, even if slightly thicker than paraffin sections, clear sufficiently to permit study of the internal structure which is usually broken up or distorted by paraffin. There are various products of nitrocellulose on the market, e.g., celloidin and collodion. They are sold as shredded or granulated products and are in- flammable, but not explosive. Schering's celloidin is in general use. Du- pont's parlodion or the product of Mallinckrodt is a purified pyroxylin which gives very good results in embedding and can be obtained as small strips sold in 1-ounce jars. Celloidin may also be obtained in tablet form with directions for making the dilutions accompanying the tablets. 72 MEDICAL MYCOLOGY Because of the particular advantages celloiclin has over paraffin in mak- ing sections of agar cultures for cji:ologic purposes, I shall list steps that I follow in my work. The general procedure is that improved by Jeffrey (1928) and recently reviewed by Wetmore (1932), but with some slight changes as are better adapted to this type of growth. When the organism is sufficiently developed on a suitable medium, the fixing agent, Hermann's fluid, is poured slowly down the side of the tube. After fixation for from 6-12 hours, depend- ing on the type of organism and growth (a surface growth requiring less time than a deep growth), the fixed culture is washed overnight in slowly running tap water. Care should be taken not to let the water run strongly or the sur- face mycelium and yeast cells will be washed away. The agar slant is now taken from the test tube, which is broken, and cut up into pieces or blocks approximately 1 sq. cm. or if large tubes are used, approximately 1 cm. by the diameter of the tube. These blocks are next dehydrated in the following alcohols, 2 hours in each: 15%, 25%, 35%, 50%, 70%, 85%, 95%, 100%. Celloidin may be used warm or cold. For best results it should be used warm. An oven is kept regulated at 45° C. The material is next transferred to a bottle with a collar, containing a solution of ether and absolute alcohol in equal proportions. The bottle is corked and secured by passing a wire around the collar and over the cork as shown by Wetmore. This is placed in the oven and allowed to lie on its side for 24 hours. The preparation is now ready to be run up in celloidin, which has been washed, thoroughly dried, and made up in 2, 4, 6, 8, and 10 per cent solutions in ether-alcohol. Higher percentages may be made, but are not necessary. The material is transferred to each suc- cessive dilution every 24 hours, tightly corked, and placed in the 45° C. oven. After the 10% solution, blocks are poured as with paraffin, care being taken not to form bubbles. The blocks are arranged carefully and then set in chloroform to harden for about 12 hours or overnight. When found to be sufficiently hard, the material is placed in 70% alcohol for a few hours to allow for some softening and then it may be stored in glycerol alcohol (95% alcohol in- definitely). The blocks may be cut with a razor blade to get rid of excess celloidin and to make a uniform block. In order to make sections, the preparations are mounted on small wooden blocks as described by Wetmore. Sections are then cut, as thin as possible, placed in 95% alcohol and then run down to dis- tilled water: 95%, 85%, 70% alcohol, distilled water, 5 minutes in each change, or else run down to the percentage alcohol of the stain used. The sections are now ready to be stained. There are various stains used, but I have found it best to use a 4% mordanting solution of iron-alum (ferric am- monium sulphate) for 10 minutes, washing 4 times with water so that the excess of mordant may be removed. Then 2 drops of Haidenhain's or Ehr- lich's hematoxylin are added to a watch glass of sections in water and allowed to stand overnight. The sections are then examined to see whether the mate- rial is sufficiently stained. If heavily overstained, they can be decolored with dilute iron alum. They should be slightly overstained, however, because the MICROSCOPY 73 higher percentage alcohols usually take out some of the dye. The sections are then again dehydrated, but a few drops of chloroform should be added to the absolute alcohol so that the celloidin will not be dissolved. Two changes in absolute alcohol and chloroform and then benzol to clear completely. The sections are then mounted in balsam, making sure that they are completely flattened out. A lead weight is placed on the cover slip and the slide placed in a warm oven for 2 or 3 days. Thus the excess balsam may be pressed out to allow the preparation to be examined with oil immersion. BIBLIOGRAPHY Adamson, H. G. 1895. Observations on the parasite of ringworm, Brit. Jour. Dermatol. 7: 201-211, 237-244. [Trans. Internat. Cong. Dermatol. 3: 555, 1897.] Amann, Jules. 1896. Conservierungsflussigkeiten und Einschlussmedien fiir Moose, Chloro- und Cyanophyceen, Zeitschr. Wiss. MiJcr. 13: 18-21. Bachmann, Kowland W. 1920. Spore identification in scrapings. Arch. Derm. Syphilol. 1: 50-54. Becker, S. William So Earl B. Kitchie. 1930. The role of yeasts in the production of super- ficial dermatitis, Arch. Derm. Syi)hilol. 22: 790-802. Bigot, A. 1924. Differents procedes de coloration des cryptocoques pathogenes en medecine veterinaire. Bull. Soc. Path. Exot. 17: 547-551. Buschke, A. & F. Harry. 1923. Beitrag zur Frage der Sporulations-fahigkeit parasitischer und pathogener Hefen, Derm.. Woch. 76: 357-360. Chalmers, Albert J. & Alexander Marshall. 1914. Tinea capitis tropicalis in the Anglo- Egyptian Sudan — the systematic position of the genus Trichophyton Malmsten, 1845, Jour. Trop. Med. Byg. 17: 257-265, 289-291, 3 pis. Cornbleet, Theodore. 1930. A reagent for demonstrating fungi in skin scrapings and hair, Jour. Am. Med. Assn. 95: 1743, 1744. Cfurtis, Otis F. & Eeginald H. Colley. 1915. Picro-nigrosin, a combination fixative and stain for algae. Am. Jour. Bot. 2: 89-92. Dalmau, Luz Maria. 1929, 1930. Observations on mycologic technique with particular refer- ence to pathogenic fungi, Porto Rico Jour. Puhlic Health and Trop. Med. 5: 302- 311. Ferrari, Angela. 1930. Un nuovo metodo per la colorazione del micelio, Atti 1st. Bot. B. Univ. Pavia IV, 2: 81-87. Greenwood, A. M. & Ethel M. Eockwood. 1930. The skin in diabetic patients, Arch. Derm. & Syphilol. 21: 96-107, 10 figs. Henrici, A. T. 1930. Molds, yeasts, and Actinomycetes, New York, John Wiley & Son, 296 pp. Hufschmitt, G., A. Sartory, E. Sartory & J. Meyer. 1931. Un cas de blastomycose cutanee a foyers multiples, Ann. Derm. Syphiligr. 7: 850-876. Jeffrey, Edward Charles. 1928. Technical contributions, I-V, Bot. Gaz. 86: 456-467, Figs. 1-3. Kater, J. McA. 1927. Cytology of Saccharomyces cereviciae with especial reference to nuclear division, Biol. Biill. 52: 436-448, £ pis. Kraus, Alfred. 1904. Zur Farbung der Hyphomyceten im Horngewebe, Zentralbl. Bakt., I, 37: 153-156. Lepik, E. 1928. Differential staining of Peronosporuceae, Phytopath. 18: 869-872. Linder, David Hunt. 1929. An ideal mounting medium for mycologists. Science 70: 430. Mallinekrodt-Haupt, Asta v. 1926. Vital farbungen mit Indikatorfarben bei Hyphomyceten, Derm. Zeitschr. 46: 263-305. Mallory, F. B. & J. H. Wright. 1924. Pathological Teclmiquc, Philadelphia and London, 666 pp. 74 MEDICAL MYCOLOGY Maneval, W. E. 1924. A method of securing spores of yeasts, Bot. Gas. 78: 122, 123. Pels, Isaac R. & S. Bayne-Jones. 1923. Elastic tissue simulating mycelial filament in skin scrapings, Arch. Derm. Syphilol. 8; 37-43. Priestley, Henry. 1917. Ringworm and allied parasitic skin diseases in Australia, Med. J. Australia [4] 2; 471-475, IS figs. Sartory, Antoine. 1924. Guide pratique des principales manipulations de mycologie para- sitaire a 1 'usage des pharmaciens, Paris, 341 pp. Smith, J. Lorrain. 1907. On the simultaneous staining of neutral fat and fatty acid by oxazine dyes, J. Path. Bact. 12: 1-4. Unna, Paul, Jr, 1929. tJber Piirbung von Fadenpilzen in der Oberhaut, Derm. Woch. 88: 314-321. Unna, Paul. 1S91. Die Farbung der Mikroorganismen im Horngewebe, Monatsch. Derm. 13: 225-237, 286-311. Weston, William H. 1929. A useful modification of Amann's medium, Science 70: 455. Wetmore, Ralph H. 1932. The use of celloidin in botanical technique, Stain Techn, 7: 37-62, 6 figs. CHAPTER VI BOTANICAL NOMENCLATUEE The problem of selecting a name for an organism is a very ancient one. Early plant names were simple nouns in the language in use by various bota- nists. As likenesses and differences were more clearly realized, adjectives were applied to distinguish between closely related groups. In the course of centuries these adjectives became attached to nouns in a definite and usually stable manner. During the period from the introdviction of printing to the middle of the eighteenth century, the noun gradually took on the generic con- cept, and the group of adjectives, the specific concept. As this usage became more prevalent, a binomial nomenclature was approached, until Linnaeus in his Species Plant arum of 1753 used it almost universally. The last half of the eighteenth century was predominantly one of ex- ploration and description of many new plants and animals. The same or- ganism was often named more than once by workers in ignorance of the pub- lications of others. Then the problem of which name to choose became in- creasingly urgent. In general, the principle of priority developed, by which the oldest binomial name for a group was chosen. During the nineteenth cen- tury, these problems became increasingly (difficult, and authors developed codes of rules for their own use. As a result of this, by the close of the nineteenth century varying practice for handling the same situation had grown up in various countries. In America we had two more or less divergent systems which caused much confusion, as one set of workers used one set of names for their plants while another group used different names for the same plants. An attempt to reach a compromise was made in the last decade of the century, but this was ineffective. At the International Congress at Paris in 1900, a committee was appointed to draft a code and report to the Congress at Vienna in 1905. This code, adopted after much discussion, forms the basis for our present code. It was extensively amended at Brussels in 1910 and at Cambridge (England) in 1930. The official edition of the Code with the 1930 amendments has not yet been published. The most active member of the editorial committee died shortly after the Congress. I have been unable to secure from the surviving member, information as to the probable time of publication. As this book goes to press, A. B. Rendle, Jour. Bot. Brit. For. 72 : Supple- ment 1-29, June, 1934) has published his version of the Code with the approval of the surviving member of the editorial committee, so that it is probable that Rendle 's version will not differ materially from the official version. In gen- eral, Rendle 's version has been reproduced in the following pages, but I have 75 7b MEDICAL MYCOLOGY corrected obvious lapsi calami and added examples of the application of the rules based on names of fungi familiar to medical men which illustrate the point as well as the examples given by Rendle. In this book, I have endeavored to follow the spirit of the International Rules, although it has been impossible to apply the letter of the law in some instances. INTERNATIONAL RULES OF BOTANICAL NOMENCLATURE* Chapter I. — General Considerations and Guiding Principles Art. 1. Botany cannot make satisfactory progress without a precise system of nomenclature, which is used by the great majority of botanists in all countries. Art. 2. The precepts on which this precise system of botanical nomenclature is based are divided into principles, rules, and recom- mendations. The principles (Art. 1-9, 10-14, 15-19 1) form the basis of the rules and recommendations. The object of the rules (Art. 19-74) is to put the nomenclature of the past into order and to pro- vide for that of the future. They are ahvays retroactive : names or forms of nomenclature contrary to a rule (illegitimate names or forms) cannot be maintained. The recommendations deal with sub- sidiary points, their object being to bring about greater uniformity and clearness in future nomenclature : names or forms contrary to a recommendation cannot on that account be rejected, but they are not examples to be followed. Art. 3. The rules of nomenclature should be simple and founded on considerations sufficiently clear and forcible for everj^one to com- prehend and be disposed to accept. Art. 4. The essential points in nomenclature are: (1) to aim at fixity of names; (2) to avoid or to reject the use of forms and names which may cause error or ambiguity or throw science into confusion. Next in importance is the avoidance of all useless creation of names. Other considerations, such as absolute grammatical correctness, regularity or euphony of names, more or less prevailing custom, re- gard for persons, etc., notwithstanding their undeniable importance, are relatively accessory. Art. 5. In the absence of a relevant rule, or where the consequences of rules are doubtful, established custom must be followed. Art. 6. Botanical nomenclature is independent of zoological no- menclature in the sense that the name of a plant is not to be rejected simplj' because it is identical with the name of an animal. If, how- ever, an organism is transferred from the animal to the plant kingdom, its validly published names are to be accepted as botanical nomen- clature in the form prescribed by the rules of botanical nomenclature ; and if an organism is transferred from the plant to the animal king- dom its names retain their status in botanical nomenclature. Art. 7. Scientific names of all groups are usually taken from Latin or Greek. When taken from any language other than Latin, or formed •This entire section set in narrow measure is reprinted from International Rules of Botanical Nomenclature adopted by the Fifth International Botanical Congress, Cambridge, 1930. Supplement to The Journal of Botany, June, 1934. Printed and published by Taylor and Francis. t Art. 19 is both a principle and a rule. BOTANICAL NOMENCLATURE 77 in an arbitrary manner, they are treated as if they were Latin. Latin terminations should be used so far as possible for new names. Art. 8. Nomenclature deals with: (1) the terms which denote the rank of taxonomic groups (Art. 10-14) ; (2) the names which are applied to the individual groups (Art. 15-72). Art. 9. The rules and recommendations of botanical nomenclature apply to all groups of the plant kingdom, recent and fossil, with cer- tain distinctly specified exceptions. Chapter II. — Categories of Taxonomic Groups, and the Terms DENOTING THEM (Art. 10-14, RcC. I, II). Art. 10. Every individual plant, interspecific hybrids and chimseras excepted, belongs to a species (species), every species to a genus (genus), every genus to a family (familia), every family to an order (ordo), every order to a class (classis), every class to a division (divisio). Art. 11. In many species varieties (variefas), forms (forma), and races or biological forms (forma hiologica) are distinguished; in parasitic species special fonns (forma specialis) , and in certain cul- tivated species modifications still more numerous ; in many genera sec- tions (sectio) are distinguished, in many families tribes (tribus). Recommendation I. In parasites, especially parasitic fungi, authors who do not give specific value to forms characterized from a biological standpoint, but scarcely or not at all from a morphological standpoint, should distinguish within the species special forms (forma specialis) characterized by their adaption to different hosts. Art. 12. Finally, if a greater number of intermediate categories are required, the terms for these subdivisions are made by adding the prefix sub (sub) to the terms denoting the categories. Thus subfamily (suh- familia) denotes a category between a family and tribe, subtribe (suh- tribus) a category between a tribe and a genus, etc. The classification of subordinated categories may thus be carried, for wild plants, to twenty-three degrees in the following order: Regnum vegetabile. Divisio. Subdivisio. Classis. Subclassis. Ordo. Subordo. Familia. Subfamilia. Tribus. Subtribus. Genus. Subgenus. Sectio. Sub- sectio. Species. Subspecies. Varietas. Subvarietas. Forma. Forma biologica. Forma specialis. Individuum. If this list of categories is insufficient it can be augmented by the intercalation of supplementary categories, provided that this does not in- troduce confusion or error : e. g., series and subseries are categories which can be intercalated between subsection and species. Recommendation II. The arrangement of species in a genus or in a subdivision of a genus is made by means of typographic signs, letters or numerals. Art. 13. The definition of each of these categories varies, up to a certain point, according to individual opinion and the state of the science; but their relative order, sanctioned by custom, must not be altered. No classification is admissible which contains such alterations : e. g., a form divided into varieties, a species containing genera. Art. 14. The fertilization of one species by another may give rise to a hybrid (hyhrida) ; that of a modification or subdivision of a species by another modification of the same species may give rise to a half- breed (mistiis). 78 MEDICAL MYCOLOGY Chapter III. — Names of Taxonomic Groups (Art. 15-72, Rec. Ill — L). Section 1. — General Principles: priority (Art. 15-17, Rec. III). Art. 15. The purpose of giving a name to a taxonomic group is not to indicate the characters or the history of the group, but to supply a means of referring to it. Art. 16. Each group with a given circumscription, position, and rank can bear only one valid name,* the earliest that is in accordance with the Rules of Nomenclature. Art. 17. No one may change a name (or combination of names) without serious motives, based either on more profound knowledge of facts or on the necessity of giving up a nomenclature that is con- trary to the Rules. Recommendation III. Changes in nomenclature should be made only after ade quate taxonomic study. Section 2.— The Type Method (Art. 18, Rec. IV-VII). Art. 18. The application of names of taxonomic groups is deter- mined by means of nomenciatural types. A nomenclatural type is that constituent element of a group to which the name of the group is permanently attached, whether as an accepted name or as a syno- nym. The name of a group must be changed if the type of that name is excluded (see Art. 66). The type of the name of an order or suborder is a family, that of the name of a family, subfamily, tribe or subtribe is a genus, that of a generic name is a species, that of the name of a species or group of lower rank is usually a specimen or preparation. In some species, however, the type is a description or figure given by a previous author. Where permanent preservation of a specimen or preparation is im- possible, the application of the name of a species or subdivision of a species is determined by means of the original description or figure. Note. — The nomenclatural type is not necessarily the most typical or repre- sentative element of a group; it is merely that element with which the name of the group is permanently associated. Recommendations : IV. When publishing names of new groups authors should indicate carefully the subdivision which is the type of the uew name: the type-genus in a family, the type-species in a genus, the type-variety or specimen in a species. This type deter- mines the application of the' name in the event of the group being subsequently divided. When describing new species, varieties or forms of parasitic plants, especially Fungi, the host plant of the type should be indicated. V. When revising a genus an author should state which species he accepts as the nomenclatural type. VI. In selecting a nomenclatural type for a genus of non-vascular Cryptogams, botanists should, where possible, choose a species that will fix the generic name as it is now commonly applied. *In genera and groups of higher rank the valid name is the earliest name pub- lished with the same rank, provided that this is in conformity with the Eules of Nomenclature and the provisions of Arts. 20 and 21. In subdivisions of genera the valid name is the earliest name published with the same rank, provided that this name and its combination with the generic name are in conformity with the Eules of Nomenclature. In species and groups of lower rank the valid name is the binarj^ or ternary combination containing the earliest epithet published with the same rank, provided that this combination is in conformity with the Eules of Nomenclature. BOTANICAL NOMENCLATURE 79 VII. The utmost importance should be given to the preservation of the orginal ("type") material on which the description of a new group is based. In micro- scopic Cryptogams the preparations and original drawings, in fleshy Fungi water- colour drawings and specimens suitably piepared or dried, should be preserved. The original account should state where this material is to be found. Section 3. — Limitation of the Principle of Priority: publication, start- ingf-points, conservation of names (Art. 19-22). Art. 19. A name of a taxonomic g-ronp has no status under the Rules, and has no claim to recognition by botanists, unless it is validly published (see Section 6, Art. 37). Art. 20. Legitimate botanical nomenclature begins for the differ- ent groups of plants at the following dates: — {a) Phanerogamae and Pteridophyta, 1753 (Linnaeus, Species Plan- tarum, ed. 1). (&) Muscineae, 1801 (Hedwig, Species Mnscorum). (c) Sphagnaceae and Hepatieae, 1753 fLinnasus, Species Pkmtarum, ed. 1). (d) Lichenes, 1753 (Linnasus, Species Plantarum, ed. 1). {e) Fungi: Urediuales, Ustilaginales and Gasteromycetes, 1801 (Per- soon, Synopsis methodica Fungorum) . (/) Fungi cfeteri, 1821-32 (Fries, Systema mycologicum) . (g) Algae, 1753 (Linnasus, Species Plantarum, ed. 1). Exceptions. — Nostocaceae homocysteae, 1892-93 (Gomont, Monographic des Oscillariees in Ann. Sci. Nat. ser. 7, Bot. vi. 91; vii, 263). — Nostocaceae heterocysteae, 1886-88 (Bomet et Flahault, Revision des Nostocacees heterocystees in Ann. Sci. Nat. ser. 7, Bot. iii, 323 ; iv, 344; v, 51 ; vii, 177). — Desmidiaceae, 1858 CRalis, British Desmidiaceae) . — Oedogoniaceae, 1900 (Hirn, Monographic und Iconographie der Oedogoniaceen in Act. Soc. Sci. Fen7i. xxvii, No. 1). {h) Myxomycetes, 1753 (Linnaeus, Species Plantarum, ed. 1). The nomenclature of Fossil Plants of all groups begins with the year 1820. It is agreed to associate generic names which appear in Linnseus's Species Plantarum., ed. 1 (1753) and ed. 2 (1762-63), with the first subsequent descriptions given under those names in Linngeus's Genera Plantarum, ed. 5 (1754) and ed. 6 (1764). Art. 21. However, to avoid disadvantageous changes in the nomen- clature of genera by the strict application of the Rules of Nomen- clature, and especially of the principle of priority in starting from the dates given in Art. 20, the Rules provide a list of names which must be retained as exceptions. These names are by preference those which have come into general use in the fifty years following their publication, or which have been used in monographs and important floristic works up to the year 1890. Note 1. — These lists of conserved names will remain permanently open for addi- tions. Any proposal of an additional name must be accompanied by a detailed statement of the cases for and against its conservation. Such proposals must be submitted to the Executive Committee, who will refer them for examination to the Special Committees for the various taxonomic groups. 80 MEDICAL MYCOLOGY Note 2. — The application of conserved names is determined by nomenclatural types, or by substitute-types where necessary or desirable. Note 3. — A conserved name is conserved against all other names for the group, whether these are cited in the corresponding list of rejected names or not, so long as the group concerned is not united or reunited with another group bearing a legitimate name. In the event of union or reunion with another group, the earlier of the two competing names is adopted in accordance with Art. 56. Note 4. — A conserved name is conserved against all earlier homonyms. Art. 22. "When a name proposed for conservation has been provi- sionally approved by the Executive Committee, botanists are author- ized to retain it pending the decision of the next International Botanical Congress. Section 4. — Nomenclature of the Taxonomic Groups according to their Categories (Art. 23-35, Rec. VIII-XX). § 1. Names of Groups ohove the Bank of Family. Recommendations : VIII. Names of divisions and subdivisions, of classes and subclasses, are taken from their chief characters. They are expressed by words of Greek or Latin origin in the plural number, some similarity of form and termination being given to those which designate groups of the same nature. IX. Orders are designated preferably by the name of one of their principal families, with the ending -ales. Suborders are designated in a similar manner, with the ending -ineae. But other terminations may be used for these names, provided that they do not lead to confusion or error. § 2. Names of Families and Suhfamilies, Tribes, and Subtrihes. Art. 23. Names of families are taken from the name or former name of one of their genera and end in -aceae. Exceptions : (1) The following names, sanctioned by long usage, are treated as exceptions to the rule: Palmae, Gramijieae, Cruciferae, Leguminosae, Guttiferae, Uinbelliferae, Labiatae, Compositae. Botanists are authorized, however, to use as alternatives the appropriate names ending in -aceae. (2) Those who regard the Papilionaceae as constituting an independent family may use that name, although it is not formed in the prescribed manner. Note.— To avoid disadvantageous changes in the nomenclature of families by the strict application of the Rules, and especially of the principle of priority, a list of names which must be retained as exceptions will be provided (Appendix II). Art. 24. Names of subfamilies (subfamiliae) are taken from the name of one of the genera in the group, with the ending -oideae, simi- larly for tribes {tribus), with the ending -eae, and for subtribes {sub- tribus), with the ending -inae. § 3. Names of Genera and Subdivisions of Genera. Art. 25. Names of genera are substantives (or adjectives used as substantives), in the singular number and written with an initial capi- tal, which may be compared with our family names. These names may be taken from any source whatever, and may even be composed in an absolutely arbitrary manner. Recommendation X. Botanists who are forming generic names show judgment and taste by attending to the following recommendations: — (a) Not to make names long or difficult to pronounce. (b) Not to dedicate genera to persons quite unconnected with botany, or at least with, natural science, nor to persons quite unknown. t BOTANICAL NOMENCLATURE 81 (c) Not to take names from barbarous languages, unless those names are frequently cited in books of travel, and have an agreeable form that is readily adaptable to the Latin tongue and to the tongues of civilized countries. (d) To indicate, if possible, by the formation or ending of the name the affinities or analogies of the genus. (e) To avoid adjectives used as nouns. (/) Not to give a genus a name whose form is rather that of a subgenus or section (e. g. Eutorula, a name given to a genus of Saccharomycetaceae Imperfectae. This, however, being legitimate, cannot be altered). (g) Not to make names by combining words from different languages (rioviina hybrida) . Art. 26. Names of subgenera and sections are usually substantives resembling the names of genera : e. g. Fraxinaster, Archieracium. Names of subsections and other lower subdivisions of genera are pref- erably adjectives in the pleural number agreeing in gender with the generic name and written with an initial capital, or their place may be taken by an ordinal number or a letter: e. g. Pleiostylae, Fimhriati, Bihracfeolata. Recommendations : XI. Botanists constructing names for subgenera or sections will do well to attend to the preceding recommendations and also to the following: — (a) To give, where possible, to the principal division of a genus a name which recalls that of the genus with some modification or addition. Thus Eu may be placed at the beginning of the generic name when it is of Greek origin, -astmrn, -ella at the end of the name when Latin, or any other modification consistent with the grammar and usages of the Latin language: e. g. Eucardamine (from Cardamine) , Drabella (from Draha). (b) To avoid giving to a subgenus or a section the name of the genus to which it belongs, with the ending -oides or -opsis: but on the contrary to reserve this ending for a section which resembles another genus and by then adding -oides or -opsis to the name of that other genus, if it is of Greek origin, to form the name of the section. (c) To avoid taking as the name of a subgenus or section a name which is already in use as such in another genus, or which is the name of a genus. (d) To avoid in co-ordinated subdivisions of a genus the use of names in the form of a noun together with those in the form of a plural adjective; the former should he used chiefly for subgenera and sections, the latter for subsections, series and subseries. XII. When it is desired to indicate the name of a subgenus or section (or other subdivision to which a particular species belongs) in connection with the generic name and specific epithet, the name of the subdivision is placed in parentheses be- tween the two (where necessary, the rank of the subdivision is also indicated) : e. g. Achorion (Sect. Lophophyton) muris. §4. Names of Species (hinary names). Art. 27. Names of species are binary combinations consisting of the name of the genus followed by a single specific epithet. If an epithet consists of two or more words, these must either be united into one or joined by a hyphen. Symbols forming part of specific epithets pro- posed by Linnaeus must be transcribed. The specific epithet, when adjectival in form and not used as a sub- stantive, agrees with the generic name. Recommendations : XIII. The specific epithet should, in general, give some indication of the appear- ance, the characters, the origin, the history or the properties of the species. If taken from the name of a person it usually recalls tlie name of the one who dis- covered or described it, or was in some way concerned with it. 82 MEDICAL MYCOLOGY XrV. Names of men and women, and also of countries and localities used as specific epithets, may be substantives in the genitive (Truchisi) or adjectives (Valeriana, lipsiense). It will be well, in the future, to avoid the use of the genitive and the adjectival form of the same epithet to designate two different species of the same genus. XV. In forming specific epithets botanists will do well to have regard also to the following recommendations: — (o) To avoid those which are very long and difficult to pronounce. (b) To avoid those which express a character common to all, or nearly all, the species of a genus. (c) To avoid using the names of little-known or very restricted localities, unless the species is quite local. (d) To avoid, in the same genus, epithets which are very much alike, especially those which differ only in their last letters. (e) Not to adopt unpublished names found in travellers' notes or in herbaria, attributing them to their authors, unless these have approved publica- tion. (/) Not to name a species after a person who has neither discovered, nor de- scribed, nor figured, nor in any way studied it. (g) To avoid epithets which have been used before in any closely allied genus. (h) To avoid specific epithets formed of two or more (hyphened) words. (i) To avoid epithets which have the same meaning as the generic name (pleonasm). § 5. Names of Groups below the rank of Species {ternary names). Art. 28. Epithets of subspecies and varieties are formed like those of species and follow them in order, beginning- with those of the high- est rank. "When adjectival in form and not used as substantives they agree with the generic name. Similarly for subvarieties, forms and slight or transient modifications of wild plants, which receive either epithets, or numbers, or letters to facilitate their arrangement. The use of a binary nomenclature for subdivisions of species is not admis- sible. It is permissible to reduce more complicated names to ternary combinations. Art. 29. The same epithet may be used for subdivisions of different species, and the subdivisions of one species may bear the same epithet as other species: e. g. Rosa JundzilUi var. leioclada and Bosa glutmosa var. leioclada, Viola tricolor var. hirta in spite of the existence already of a different species named Viola hirta. Art. 30. Two subdivisions of the same species, even if they are of different rank, cannot bear the same subdivisional epithet, unless they are based on the same type. If the earlier subdivisional name (ternary combination) was validly published, the later one is illegitimate, and must be rejected. The ternary combinations Biscutella didyma subsp. apula Briq. and Biscutella didyma var. apula Halacsy may both be used because they are based on the same type, and the one includes the other. The following is incorrect: Erysimum hieracifolium subsp. strictum var. longisiliquum and E. hieracifolium subsp. pannonicum var. longisiliquum — a form of nomenclature which allows two varieties bearing the same name in the same species. Recommendations : XVI. Eecommendations made for specific epithets apply equally to epithets of subdivisions of species. XVII. Special forms (forma specialis) are preferably named after the host species ; if desired, double names may be used : e. g. Puccinia HieracU f . sp. villosi, Pucciniastrum Epilohii f. sp. Aiieti-Chamaenerii. XVIII. Botanists should avoid giving a new epithet to any subdivision of a species which includes the type either of the specific name or of a higher sub- BOTANICAL NOMENCLATURE 83 divisional name. They should either repeat that epithet or use one of the customary- epithets, typicus, genuinus, originarvus, etc. E. g. Andropngon caricosus subsp. mollissimus var. mollissim'us Hackol; Arthraxon ciliaris Bcauv. subsp. Langsdorfii var. genuinus Hackel. XIX. Botanists proposing new epithets for subdivisions of species should avoid such as have been used previously in the same genus, whether for species or for subdivisions of other species. § 6. Names of Hybrids and Half-hreeds. Art. 31. Hybrids or putative hybrids between species of the same genus are designated by a formula and, whenever it seems useful or necessary, by a name. (1) Sexual hybrids. — The formula consists of the names or specific epithets of the two parents in alphabetical order and connected by the sign X. When the hybrid is of known experimental origin the fonnula may be made more precise by the addition of the signs 9 $ , the name of the female (seed-bearing) parent being placed first. The name, which is subject to the same rules as names of species, is distinguished from the latter by the sign x before the name : e. g. X Salix capreola {Salix anrita x caprea) . (2) Asexual hybrids (graft hybrids, chinueras, etc.). — The formula consists of the names of the two parents in ali)habetical order connected by the sign +. The name has a "specific" epithet different from that of the corresponding sexual hj'brid (if any), and is preceded by the sign +: e. g. + Solanum tubingense {Solanum nigrum + 8. Lycopersi- cum) . Art. 32. Bigeneric hybrids (hybrids between species of two genera) are also designated by a formula and, whenever it seems useful or necessaiy, by a name. The formula consists of the names of the two parents connected by a sign, as in Art. 31. The name consists of a new "generic" name usually formed by a combination of the names of the parent genera, and a "specific" epithet. All hybrids (whether sexual or asexual) between the same two genera bear the same "generic" name. (1) Sexual hybrids. — In the formula the connecting sign x is used. The name is preceded by the sign x : e. g. x Odoniioda Boltonii (Coch- lioda Noezlianax Odontoglossum Vuylstekeae) . (2) Asexual hybrids. — In the formula the connecting sign + is used. The name is preceded by the sign +. The "specific" epithet is differ- ent from that of the corresponding sexual hybrid (if any) between the same species. E. g. + Laburnocytisus Adami {Laburnum anagyroides +Cytisus purpurens) . Art. 33. Ternar.y hybrids, or those of a higher order, are designated, like ordinary hybrids, by a formula and, whenever it seems useful or necessary, bj^ a name. Such as are trigeneric or polygeneric are given new "generic" names usually formed by a combination of the names of the parent genera. Recommendation XX. Half-breeds, or putative half-breeds, may be designated by a name and a formula. Names of half-breeds are intercalated among the sub- divisions of a species, and are preceded by the sign x. In tlie formula the names of the parents are in alphabetical order. When the half-breed is of known experi- mental origin the formula may be made more precise by the addition of the signs 9 $, the name of the female (seed-bearing) parent being placed first. 84 MEDICAL MYCOLOOY Art. 34. When different hybrid forms of the same parentage (pleomorphic hybrids; combinations between different forms of a col- lective species, etc.) are united in a collective group, the subdivisions are classed under the binary name of the hybrid, like the subdivisions of a species under that of a species. Examples: x Mentha niliaca /3 Lamarckii (=: M. longifolia x rotundifolia) . The preponderance of the characters of one or other parent may be indicated in the formulae in the following manner : Mentha longifolia > x rotundifolia, M. longi- folia X < rotundifolia. The participation of a particular variety may also be indicated : e. g. Salix caprea x daphnoides var. pulchra. § 7. Names of Plants of Horticultural Origin. Art. 35. Forms and half-breeds among cultivated plants receive fancy epithets, preferably in common language, as different as possible from the Latin epithets of species or varieties. When they can be at- tached to a species, a subspecies or a botanical variety this is indi- cated by a succession of names : e. g. Pelargonium zonMe Mrs. Pollock. Section 5. — Conditions of effective Publication. Art. 36. Publication is effected, under these Rules, either by sale or distribution of printed matter or indelible autographs to the gen- eral public, or to specified representative botanical institutions*. No other kind of publication is accepted as effective : communica- tion of new names at a public meeting, or the placing of names in collections or gardens open to the public, does not constitute effective publication. Section 6. — Conditions and Dates of valid Publication of Names (Art. 37-45, Rec. XXI-XXIX). Art. 37. A name of a taxonomic group is not validly published un- less it is both (1) effectively published (see Art. 36), and (2) accom- panied by a description of the group or by a reference to a previously and effectively published description of it. Mention of a name on a ticket issued with a dried plant without a printed or autographed description does not constitute valid publica- tion of that name. Note. — In certain circumstances a plate or figure with analyses is accepted as equivalent to a description (vide Art. 43, 44). Art. 38. From January 1, 1935, names of new groups of recent plants, the Bacteria excepted, are considered as validly published only when they are accompanied by a Latin diagnosis. Art. 39. From January 1, 1912, the name of a new taxonomic group of fossil plants is not considered as validly published unless it is ac- companied by illustrations or figures showing the essential characters, in addition to the description. Art. 40. A name of a taxonomic group is not validly published when it is merely cited as a synonym. Art. 41. A group is not characterized, and the publication of its name is not validated, merely by mention of the subordinate groups included in it : thus the publication of the name of the order is not *The preparation of a list of representative botanical institutions is referied to tlie Executive Committee (see App. VII). BOTANICAL NOMENCLATURE validated by mention of the included families; that of a family is not validated by mention of the included genera ; that of a genus is not validated by mention of the included species. E. g. the generic name Ibidinm Salisbuiy (Trans. Hort. Soc. i, 291: 1812) was published merely with the mention of four included species: as Salisbury sup- plied no generic description, the publication of Ihidium was invalid. Art. 42. A name of a genus is not validly published unless it is ac- companied (1) by a description of the genus, or (2) by the citation of a previously and effectively published description of the genus under another name, or (3) by a reference to a previously and effectively published description of the genus as a subgenus, section or other sub- division of a genus. An exception is made for the generic names published by Linnaeus in Species Plantarum, ed. 1 (1753) and ed. 2 (1762-63), which are treated as having been validly published on those dates (see Art. 20). Note. — In certain circumstances a plate witli analyses is accepted as equivalent to a generic description (see Art. 43). Art. 43. The name of a monotypic new genus based on a new species is validated (1) by the provision of a combined generic and specific de- scription (descriptio generico-specifica) , (2) by the provision of a plate with analj^ses showing essential characters; but this applies only to plates and generic names published before January 1, 1908. Art. 44. The name of a species or of a subdivision of a species is not validly published unless it is accompanied (1) by a description of the group, or (2) by the citation of a previously and effectively published description of the group under another name, or (3) by a plate or figure w^th analyses showing essential characters; but this applies only to plates or figures published before January 1, 1908. Art. 45. The date of a name or of an epithet is that of its valid publication (see Art. 19, 36). For purposes of priority, however, only legitimate names and epithets published in legitimate combinations are taken into consideration* (see Art. 60). In the absence of proof to the contrary, the date given in the work containing the name or epi- thet must be regarded as correct. On and after January 1, 1935, only the date of publication of the Latin diagnosis can be taken into account for recent plants except Bacteria. For fossil plants, on and after January 1, 1912, the date is that of the simultaneous publication of the description and figure (or, if these are published at different dates, the later of the two dates). Botanists will do Avell in publishing to conform to the following recommendations :■ — ■ XXI. Not to publish a new name without clearly indicating whether it is the name of a family or a tribe, a genus or a section, a species or a variety; briefly, without expressing an opinion as to the rank of the group to which the name is given. Not to publish the name of a new group without indicating its type (see Eecom- mendation IV). XXII. To avoid publishing or mentioning in their publications unpublished names which they do not accept, especially if the persons responsible for these names have not formally authorized their publication (see Recommendation XV 85 •A legitimate name or epithet is one that is strictly in accordance with the Rules. 86 MEDICAL MYCOLOGY XXIII. When publishing names of new groups of plants in works written in a modern language (floras, catalogues, etc.), to publish simultaneously the Latin diagnoses of recent plants (Bacteria excepted) and the figures of fossil plants, which will make these names valid according to the Eules. XXIV. In describing new groups of lower Cryptogams, especially among the Fungi, or among microscopic plants, to add to the description a figure or figures of the plants, with details of microscopic structure, as an aid to identification. XXV. The description of parasitic plants should always be followed by the indication of the hosts, especially in the case of parasitic fungi. The hosts should be designated by their Latin scientific names and not by popular names in modern languages, the significance of which is often doubtful. XXVI. To give the etymology of new generic names and also of new epithets when the meaning of these is not obvious. XXVII. To indicate precisely the date of publication of their works and that of the placing on sale or the distribution of named and numbered plants when these are accompanied by printed diagnoses. In the case of a work appearing in parts, the last published sheet of the volume should indicate the precise dates at which the different fascicles or parts of the volumes were published as well as the number of pages in each. XXVIII. When works are published in periodicals, to require the publisher to indicate on the separate copies the date (year and month) of publication and also the title of the periodical from which the work is extracted. XXIX. Separate copies should always bear the pagination of the periodical of which they form a part ; if desired they may also bear a special pagination. Section 7. — Citation of Authors' Names for purposes of precision (Art. 46-49, Ree. XXX-XXXII). Art. 46. For the indication of the name (unitary, binary, or ter- nary) of a group to be accurate and complete, and in order that the date may be readily verified, it is necessary to cite the author who first published the name in question. Art. 47. An alteration of the diagnostic characters or of the cir- cumscription of a group does not warrant the citation of an author other than the one who first published its name. When the changes have been considerable, an indication of their nature and of the author responsible for the change is added, the words mutatis charact., or pro parte, or excl. gen., excl. sp., excl. var., or some other abridged indication being employed. Examples: Phyllanthus L. em. (emendavit) Miill. Arg.; Myosotis L. pro parte, K. Br.; Globularia cordifolia L. excl. var. (em. Lam.). Art. 48. When a name of a taxonomic group has been proposed but not published by one author, and is subsequently validly published and ascribed to him (or her) by another author who supplied the descrip- tion, the name of the latter author must be appended to the citation with the connecting word " ex. " The same holds for names of garden origin cited as "Hort." E. g. Capparis lasiantka R. Br. ex DC; Ges- neria Donklarii Hort. ex Hook. If it is desirable or necessary to abbreviate such a citation, the name of the publishing author, being the more important, must be retained. Where a name and description bj' one author are published by an- other author, the word ai^ud is used to connect the names of the two authors, except where the name of the second author forms part of the title of a book or periodical in which case the connecting word in is used instead. BOTANICAL NOMENCLATURE 87 Art. 49. When a genus or a group of lower rank is altered in rank but retains its name or epithet, the original author must be cited in parentheses, followed by the name of the author who effected the alteration. The same holds when a subdivision of a genus, a species, or a group of lower rank is transferred to another genus or species with or without alteration of rank. Examples: Mcdicago polymorpha L. var. orbicularis L., when raised to the rank of a species, becomes Medicago orhicuJaris (L.) All. Sorbus sect. Ari^i Pers., on transference to Py)'us, is cited as Pijrus sect. Aria (Pers.) DC. Recommendations : XXX. Authors ' names put after names of plants are ablireviated, unless they are very short. For this purpose preliminary particles or letters that, strictly speaking, do not form part of the name are suppressed, and the first letters are given without any omission. If a name of one syllable is long enough to make it worth while to abridge it, the first consonants only are given (Br. for Brown) ; if the name has two or more syllables, the first syllable and the first letter of the following one are taken, or the two first when both are consonants (Juss. for Jussieu, Eich. for Eichard). Wlien it is necessary to give more of a name to avoid confusion between names begiiming with the same syllables, the same system is to be followed. For instance, two syllables are given together with the one or two first consonants of the third; or one of the last characteristic consonants of the name is added (Bertol. for Bertoloni, to distinguish from Bertero ; Michx. for Michaux, to distinguish from Micheli). Christian names or accessory designations, serving to distinguish two botanists of the same name, are abridged in the same way (Adr. Juss. for Adrien de Jussieu, Gaertn. fil. or Gaertn. f. for Gaertner filius). "When it is a well-established custom to abridge a name in another manner it is best to conform to it (L. for Linnseus, DC. for De Candolle, St. Hil. for Saint- Hilaire). In publications destined for the general public and in titles it is preferable not to abridge. XXXI. When citing a name published as a synonym, the words ''as synonym," or pro synon. should be added to the citation. "When an author published as a synonym a manuscript name of another author, the word ex should be used to con- nect the names of the two authors: e. g. Myrtus serratus Koenig ex Steud. Nomencl. 321 (1821), pro synon., a manuscript name of Koenig 's published by Steudel as a synonym of Eugenia laurina Willd. XXXII. The citation of authors earlier than the starting point of the nomen- clature of a group is indicated, when considered useful or desirable, preferably be- tween brackets or by the use of the word ex. This method is especially applicable in mycology when reference is made to authors earlier than Fries or Persoon. Section 8. — Retention of Names or Epithets of Groups which are re- modelled or divided (Art. 50-52). Art. 50. An alteration of the diagnostic characters, or of the cir- cumscription of a group, does not warrant a change in its name, except so far as this may be necessitated (1) by transference of the group (Art. 53-55), or (2) by its union with another group of the same rank (Art. 56-57), or (3) by a change of its rank (Art. 58). Examples: The genus Myosotis as revised by E. Brown differs from the original genus of Liimffius, but the generic name has not been changed, nor is a change allowable. — Various authors have united with Centaurea Jacea L. one or two species which Linnaeus had kept distinct; the group thus constituted must be called Centaurea Jacea L. sensu ampl. or Centaurea Jacea L. em. "Visiani, or cm. Godron, etc. : the creation of a new name such as Centaurea vulgaris Godr. is superfluous. Art. 51. When a genus is divided into two or more genera, the generic name must be retained for one of them, or (if it has not been retained) must be re-established. When a particular species was 88 MEDICAL MYCOLOGY originally designated as the type, the generic name must be retained for the genus including that species. When no type was designated, a type must be chosen according to the regulations which will be given (Appendix I). Art. 52. When a species is divided into two or more species, the specific epithet must be retained for one of them, or (if it has not been retained) must be re-established. When a particular specimen was originally designated as the type, the specific epithet must be retained for the species including that specimen. When no type was designated, a type must be chosen according to the regulations to be given (Appendix I). The same rule applies to subdivisions of species ; for example, to a subspecies divided into two or more subspecies, or to a variety di- vided into two or more varieties. Section 9.^ — Retention of Names or Epithets of Groups below the Rank of Genus on transference to another Genus or Spe- cies (Art. 53-55). Art. 53. When a subdivision of a genus is transferred to another genus (or placed under another generic name for the same genus) without change of rank, its subdivisional name must be retained, or (if it has not been retained) must be re-established unless one of the following obstacles exists: (1) that the resulting association of names has been previously published validly for a different subdivi- sion, or (2) that there is available an earlier validly published sub- divisional name of the same rank. E. g. Saponaria sect. Vaccaria DC, transferred to Gypsophila, becomes Gypsophila sect. Vaccaria (DC.) Gren. & Godr. Art. 54. When a species is transferred to another genus (or placed under another generic name for the same genus), without change of rank, the specific epithet must be retained or (if it has not been re- tained) must be re-established, unless one of the following obstacles exists: (1) that the resulting binary name has been previously and validly published for a different species, (2) that there is available an earlier validly published specific epithet. When the specific epithet, on transference to another genus, has been applied erroneously in its new position to a different plant, it must be retained for the plant on which the group was originally based : e. g. the specific epithet of Pinus Mertensiana Bong, was trans- ferred to Tsuga by Carriere, who, however, erroneously applied the new combination Tsuga Mertensiana to another species of Tsuga, namely, T. heterophylla (Raf.) Sarg., as is evident from his descrip- tion: the epithet Mertensiana (Bong.) must be retained for Pinus Mertensiana Bong, when that species is transferred to Tsuga; the cita- tion in parentheses (under Art. 49) of the name of the original author, Bongard, indicates the type of the epithet, Tsuga Mertensiana (Bong.) Sargent, non Carriere. Art. 55. When a variety or other subdivision of a species is trans- ferred, without change of rank, to another genus or species (or placed under another generic or specific name for the same genus or species), the original subdivisional epithet must be retained or (if it has not been retained) must be re-established, unless one of the fol- lowing obstacles exists: (1) that the resulting ternary combination BOTANICAL NOMENCLATURE 89 has been previously and validly published for a subdivision based on a different type, even if that subdivision is of a different rank; (2) that there is an earlier validly published subdivisional epithet available. When the epithet of a subdivision of a species, on transference to another species, has been applied erroneously in its new position to a different plant, the epithet must be retained for the plant on which the group was originally based. Example: The variety micranthum Oren. & Godr. (Fl. France, i, 171: 1847) of Helianthermi/m italiouTn Pers., when transferred as a variety to H. penicillatum Thib., retains its varietal epithet, becoming H. penicillatum var. micranth^tm (Gren. & Godr.) Grosser (in Engl. Pflanzenreich, Heft 14, 115: 1903). Section 10. — Choice of Names when two Groups of the same Rank are united, or in Fungi with a pleomorphic Life-cycle (Art. 56, 57, Rec. XXXIII-XXXV). Art. 56. When two or more groups of the same rank are united the oldest legitimate name or (in species and their subdivisions) the old- est legitimate epithet is retained. If the names or epithets are of the same date, the author who unites the groups has the right of choos- ing one of them. The author who first adopts one of them, definitely treating another as a synonym or referring it to a subordinate group, must be followed. Recommendations: XXXni. Authors who have to clioose between two generic names should note the following recommendations: — 1. Of two names of tlie same date to prefer the one which was first accom- panied by the description of a species. 2. Of two names of the same date, both accompanied by descriptions of species, to prefer the one which, when the author made his choice, in- cluded the larger number of species. 3. In cases of equality from these various points of view to prefer the more correct and appropriate name. XXXIV. "When several genera are united as subgenera or sections under one generic name, the subdivision including the type of the generic name used may bear that name unaltered (e.g. Anarrhinum sect. Anarrhiniim), or with a prefix (An- thriscus sect. Eu-Anthriscus) , or a suffix (Stachys sect. Stachyotypiis). These pre- fixes or suffixes lapse when the subdivisions are raised to generic rank. XXXV. When several species are united as subspecies or varieties under one specific name, the subdivision whicli includes the type of the specific epithet used may be designated either by the same epithet unaltered (e. g. Stachys recta subsp. recta), or with a prefix (e.g. Alchemilla alpina subsp. eii-alpina), or by one of the customary epithets (typicus, origmarius, genuinvs, verus, veridicus, etc.), in- dicating that it is the type subdivision. Art. 57. Among Fungi with a pleomorphic life-cycle the different successive states of the same species {anamorphoses, status) can bear only one generic and specific name (binary), that is the earliest which has been given, starting from Fries, Sy sterna, or Persoon, Synopsis, to the state containing the form which it has been agreed to call the per- fect form, provided that the name is otherwise in conformity with the Rules. The perfect state is that which ends in the ascus stage in the Ascomycetes, in the basidium in the Basidiomycetes, in the teleutospore or its equivalent in the TJredinales, and in the spore in the TJstilaginales. Generic and specific names given to other states have only a temporary value. They cannot replace a generic name already existing and apply- ing to one or more species, any one of which contains the ''perfect" form. 90 MEDICAL MYCOLOGY The nomenclature of Fnngi wliicli have not a pleomorphic life-cycle follows the ordinary rules. Section 11.^ — Choice of Names when the Rank of a Group is changed. Art. 58. AVhen a tribe becomes a family, when a subg'enus or sec- tion becomes a genus, when a subdivision of a species becomes a species, or when the reverse of these changes takes place, and in gen- eral when a group changes its rank, the earliest legitimate epithet given to the group in its new rank is valid, unless that name or the resulting association or combination is a later homonym (see Art. 60, 61). E. g. the section Campanopsis R. Br. (Prodr. Fl. Nov. Holl. 561 : 1810) of the genus Campanula was first raised to generic rank by Schrader, and as a genus must be called Wahlenbergia Schrad. (Cat. Hort. Goett.: 1814). not Campanopsis (R. Br.) 0. Kuntze {Eev. Gen. ii, 378: 1891). Recommendation XXXVI. 1. When a sub-tribe becomes a tribe, when a tribe becomes a subfamily, when a subfamily becomes a family, etc., or when the inverse changes occur, the root of the name should not be altered but only the termination (-inae, -eae, -oideae, -aceae, -ineae, -ales, etc.), unless the resulting name is re- jected under Section 12 or the new name becomes a source of error or there is some other serious reason against it. 2. When a section or a subgenus becomes a genus, or the inverse changes occur, the original name should be retained unless it is rejected under Section 12. 3. When a subdivision of a species becomes a species, or the inverse change oc- curs, the original epithet should be retained unless the resulting combination is re- jected under Section 12. Section 12.— Rejection of Names (Art. 59-69, Rec. XXXVII). Art. 59. A name or epithet must not be rejected, changed, or modi- fied merely because it is badly chosen, or disagreeable, or because an- other is preferable or better known (see also Art. 69). Art. 60. A name must be rejected if it is illegitimate (see Art. 2). The publication of an epithet in an illegitimate combination must not be taken into consideration for purposes of priority (see Art. 45). A name is illegitimate in the following cases : — (1) If it was superfluous when published, i. e. if there was a valid name (see Art. 16) for the group to which it was applied, with its par- ticular circumscription, position and rank. (2) If it is a binary or ternary name published in, contravention of Art. 16, 50, 52 or 54, i. e. if its author did not adopt the earliest legiti- mate epithet available for the group with its particular circumscrip- tion, position, and rank. (3) If it is a later homonym (see Art. 61) (except as regards Art. 54 and 55). (4) If it is a generic name which must be rejected under Art. 67. (5) If its specific epithet must be rejected under Art. 68. Art. 61. A name of a taxonomic group is illegitimate and must be rejected if it is a later homonym^ that is, if it duplicates a name pre- viously and validly published for a group of the same rank based on a different type. Even if the earlier homonym is illegitimate, or is gen- erally treated as a synonym on taxonomic grounds, the later homonym must be rejected. BOTANICAL NOMENCI.ATimE Examples: The generic name Tapei7ia7ithus Boiss. ex Benth. (1848), given to a genus of Labiatae, is a later homonym of Tapeinanthus Herb. (1837), a name pre- viously and validly published for a genus of Amaryllidaceae ; Tapeinanthus Boiss. ex Benth. must, therefore, be rejected, as was done by Th. Durand (Ind. Gen. Than. 703: 1888) who renamed it TMispeinanta. — The generic name Amblyanthera Miill. Arg. (1860) is a later homonym of the validly published generic name Amblyanthera Blume (1849), and must, therefore, be rejected, although Ambly- anthera Blume is now reduced to OsbecMa L. (1753). — Astragalus rhisanthus Boiss. (Diagn. Fl. Or. ser. 1^ ii, 83: 1843) is a later homonym of the validly published name Astragalus rhisanthus Koyle (Illstr. Bot. Himal. 200: 1835), and it must, therefore, be rejected, as was done by Boissier, who renamed it A. cariensis (Diagn. ser. 1, ix, 57: 1849). Note. — Mere orthographic variants of the same name are treated as homonyms — see Art. 70. Art. 62. A name of a taxonomic group must be rejected if, owing to its use with different meanings, it becomes a permanent source of confusion or error. A list of names to be abandoned for this reason (Nomina amhigua) will form Appendix IV. Examples: The generic name Alsine L., being used by various authors for three genera of Caryophyllaceae (Stellaria L., Spergularia J. & C. Presl, Mirmiartia L.), has been a permanent source of confusion and error (see Sprague in "Kew Bul- letin," 1920, 308).— The name Eosa villosa L., 8p. Fl. ed. 1, 491 (1753), is re- jected, because it has been applied to several different species, and has become a source of confusion. Art. 63. A name of a taxonomic group must be rejected when its application is uncertain (nomen duhium) : e. g. Ervum soloniense L. (Cent. II. PI. 28: 1756) is a name the application of which is uncer- tain; it must, therefore, be rejected (see Schinz and Thell. in Vier- teljahresschr. Nat. Ges. Zurich, Iviii, 71: 1913). Recommendation XXXVII. Wlien the correct application of a nomen dubium has been established by subsequent investigation (of types, etc.), authors adopting it should, for purposes of precision, cite the name of the author who published the additional certifying evidence as well as that of the original author. It is also desirable to add the date of certification. Art. 64. A name of a taxonomic group must be rejected if the charac- ters of that group were derived from two or more entirely discordant elements, especially if those elements were erroneously supposed to form part of the same individual : e. g. the characters of the genus Schrehera L. (Sp. PI. ed. 2, 1662; 1763; Gen. PI. ed. 6, 124: 1764) were derived from the two genera Cuscufa and Myrica (parasite and host), see Retzius (Ohs. vi, 15: 1791). A list of names to be abandoned for this reason (Nomina confum) will form Appendix VI. Art. 65. A name or epithet of a taxonomic group must be rejected when it is based on a monstrosity. Art. 66. The name of an order, suborder, family or subfamily, tribe or subtribe must be changed when it is taken from the name of a genus which is known not to belong to the group in question — e. g. if the genus Portidaca were excluded from the family now known as Portulacaceae, the residual group could no longer bear the name Port- ulacaceae, and would have to be renamed. Art. 67. Names of genera are illegitimate in the following special cases and must be rejected : — (1) When they are merely words not intended as names: e. g. Anonymos Walt. (Fl. Carol, 2, 4, 9, etc.: 1788) must be re- jected as being a word applied to 28 different genera by Walter to indicate that they were without names. 9] 92 MEDICAL MYCOLOGY (2) When they coincide with a technical term currently used in morphology unless they were accompanied, when originally published, by specific names in accordance with the binary method of Linnffius. On and after Jan. 1, 1912, all new generic names coinciding with such technical terms are uncondition- ally rejected. (3) When they are unitary designations of species: e. g. Ehrhart (Phytophylacmm: 1780; and Beitr. iv, 145-150: 1798) pro- posed unitaiy names for various species known at that time under binary names : e. g. Phaeocephalnm for Schoenus fuscus, and Leptostachys for Carex leptostachys. These names, which resemble generic names, should not be confused with them, and must be rejected, unless they have been published as generic names by a subsequent author. (4) When they consist of two words, unless these words were from the first combined into one, or joined by a hyphen. Art. 68. Specific epithets are illegitimate in the following cases and must be rejected : — (1) When they are merely words not intended as names: e. g. Viola "qiialis" Krocker {Fl. Siles. ii, 512 and 517: 1790) ; Atriplex "nova" Winterl (in l7id. Hort. Bot. Univ. Pest. fol. A 8, recto et verso: 1788), the word "nova'' being here used in connection with four different species of Atriplex. (2) When they are merely ordinal adjectives being used for enu- meration. (3) When they exactly repeat the generic name with or without the addition of a transcribed symbol. (4) When they were published in works in which the Linnean system of binary nomenclature for species was not consis- tently employed. Art. 69. It cases foreseen in Art. 60-68 the name or epithet to be rejected is replaced by the oldest legitimate name, or (in a combina- tion) by the oldest legitimate epithet. If none exists, a new name or epithet must be chosen. Where a new epithet is required, an author may, if he wishes, adopt an epithet previously given to the group in an illegitimate combination, if there is no obstacle to its employment in the new position or sense. Section 13.— Orthography of Names (Art. 70, 71, Eec. XXXVIII- XLIV). Art. 70. The original spelling of a name or epithet must be re- tained, except in the case of a typographic error, or of a clearly unintentional orthographic error. When the difference between two generic names lies in the termination, these names must be regarded as distinct, even though differing by one letter only. This does not apply to mere orthographic variants of the same name. Note 1. The words "original spelling" in this Article mean the spelling em- ployed when the name was validly published. 2. The use of a wrong connecting vowel or vowels (or the omission of a connecting vowel in a specific epithet, or in that of a subdivision of a species) is treated as an unintentional orthographic error which may be corrected (see Eec. XLIV). BOTANICAL XOMENCLATURE 93 3. In deciding wlietlicr two or more sliglitly different names should be treated :is distinct or as orthographical variants, the essential con- sideration is wliether they may be confused with one anotlier or not : if there is serious risk of confusion, they sliould be treated as orthographic variants. Doubtful cases should be referred to the Executive Committee. 4. Specific and other epithets of Greek origin differing merely by having Greek and Latin terminations respectively are orthographic variants. Epithets bearing the same meaning and differing only slightly in form are (considered as) orthographic variants. The genetive and ad- jectival forms of a personal name are, however, treated as different epithets (e.g. Lysimachia Eemsleyana and L. Hemsleyi). Recommendations : XXXVIII. Wlien a new name is derived from a Greek word containing the spiritiis asper (rough breathing), this should be transcribed as the letter h. XXXIX. When a new name for a genus, subgenus or section is taken from the name of a person, it should be formed in the following manner: — (a) When the name of the person ends in a vowel the letter a is added (thus Ashbya after Ashby; Blaheslea after Blakeslee), except when the name already ends in a, when ea is added (e. g. Collaea after CoUa). (b) When the name of the person ends in a consonant, the letters ia are added (e. g. Magnusia after Magnus, Guilliera after Guillier), ex^ cept when tlie name ends in er, when a is added, e. g. Kernera after Kerner). (c) The syllables which are not modified by these endings retain their original spelling, even with the consonants fc and w or with groupings of vowels which were not used in classical Latin. Letters foreign to botanical Latin should be transcribed, and diacritic signs suppressed. The Germanic a, o, ii become ae, oe, ue ; the French e, e, e become generally e. In works in which diphthongs are not represented by special type, the diaeresis sign should be used where required, e.g. C'cphaelis, not CepJiaelis. (d) Names may be accompanied by a prefix or a suffix, or modified by ana- gram or abbreviation. In these cases they count as different words from the original name. Examples: Durvillea and Urvillea; Lapeyrousea and Peyrousea; Bouchea and Ubochea; Gerardia and Graderia; Thaxtera, Thax- teriola. XL. When a new specific or other epithet is taken from the name of a man, it should be formed in the following manner: — (a) When the name of the person ends in a vowel, the letter i is added (thus Caoi from Cao), except when the name ends in a, when e is added (thus Faverae from Favera). (&) When the name ends in a consonant, the letters u are added (thus Magrmsii from Magnus, Gttilliermoudii from Guilliermond) , except when the name ends in -er, when i is added (thus Thaxteri from Thaxter). (c) The syllables which are not modified by these endings retain their origi- nal spelling, even when the consonants k or w or with groupings of vowels which were not used in classical Latin. Letters foreign to botanical Latin should be transcribed and diacritic signs suppressed. The Germanic a, o, ii become ae, oe, ue, tlie French e, e, e become generally e. The diaeresis sign should be used where required. (d) When epithets taken from the name of a person have an adjectival form they are formed in a similar way (e. g. Geranium Eohertianum, Verbena Hasslerana). XLI. The same provisions apply to epithets formed from the names of women. When these have a substantival form they are given a feminine termination (e. g. Cypripedium JSookerae, Bosa Beatricis, Scabiosa Olgae, Omphalodes Luciliae.) 94 MEDICAL MYCOLOGY XLII. New specific (or other) epithets should be written in conformity with the original spelling of the words from which they are derived :ind in accordance with the rules of Latin and latinization. Examples: silvestris (not sylvestris) , sinensis (not chinensis). XLTII. Specific (or other) epithets should be written with a small initial letter, except those which are derived from names of persons (substantives or adjectives) or are taken from generic names (substantives or adjectives). XLIV. In the formation of specific (or other) epithets composed of two or several roots taken from Latin or Greek, the vowel placed between the two roots becomes a connecting vowel, in Latin i, in Greek o; thus menthifolia, salvUfolia, not Menthae folia, salviaefolia. When the second root begins with a vowel and euphony requires, the connecting vowel should be eliminated (e. g. lepidantha) . The connecting vowels ae should be retained only where this is required for etymological reasons (e. g. caricaeformis from Carica, in order to avoid confusion with car- iciformis from Carex). In certain compounds of Greek words no connecting vowel is required, e. g. hrachycarpus and glycyphyllus. Art. 71. When the spelling of a generic name differs in Linnffius' Species Plantarum, ed. 1, and Genera Plantanoii, ed. 5, the correct spelling is determined by the following regulations: — (1) If Linnffius subsequently to 1753-54 consistently adopted one of the spellings, that spelling is accepted, e. g. Thuja (not Thmja) . (2) If Linnfeus did not do so, then the spelling which is more cor- rect philologically is accepted, e. g. Agrostemma (not Agro- stema) . (3) If the two spellings are equally correct philologically, and there is a great preponderance of usage in favor of one of them, that one is accepted, e. g. Rhododendron (not Rhododen- drum). (4) If the two spellings are equally correct philologically and there is not a great preponderance of usage in favor of one of them, then the spelling that is in accordance or more nearly in accordance with the Recommendations is accepted, e. g. Ludwigia (not Ludvigia), Ortegia (not Ortega). Section 14. — Gender of Generic Names. Art. 72. The gender of generic names is governed by the following regulations : — (1) A Greek or Latin word adopted as a generic name retains the gender assigned to it by its author: e. g. Orchis (f.), St achy s (f.). (2) Generic names which are modern compounds formed from two or more Greek or Latin words take the gender of the last. If the ending is altered, however, the gender will follow it. Examples of names formed from Greek* words: The generic name Andropogon L. was treated by Linnaeus as neuter, but it, like all other modern compounds in which the Greek masculine word pogon is the final element (e. g. Centropogon, Cymbopogon, Rhisopogon), is now treated as masculine. Similarly all modern com- pounds ending in -codon, -myces, -odon, -panax, -stemon and other masculine words are masculine. The generic name Dendromecon Benth., Eomecon Hance and Ees- peramecmi E. L. Greene are treated as feminine, because they end in the Greek fem- inine word mecon, poppy: the fact that Bentham and E. L. Greene respectively as- ♦Examples of names formed from Latin words are not given, as these offer few difficulties. BOTANICAL NOMENCLATURE 95 cribed the neuter gender to the names Dendromecon and Hesperomecon is imma- terial. Similarly all modern compounds ending in -ackne, -carpha, -cephala, -chlamys, -daphne and other feminine words are treated as feminine. The generic names Aceras 11. Br., Aegiceras Gaertn. and Xanthoceras Bunge are neuter because they end in the Greek neuter word ceras ; the fact that Kobert Brown and Bunge respectively made Aceras and Xanthoceras feminine is immaterial. Sim- ilarly all modem compounds ending in -dendron, -nema, -stigma, -stoma and other neuter words are neuter. Names ending in -anthos (or anthus) and those in -chilos (or -chilus) ought strictly speaking to be neuter, since that is the gender of the Greek words anthos and cheilos. These names, however, have been with very few exceptions treated as masculine, hence it is agreed to assign that gender to them. Similarly those ending in -gaster, which should strictly speaking be feminine, are treated as masculine in accordance with botanical custom. Examples of compound generic names where the termination of the last word is altered: Hymenocarpus, Dipterocarpus and all other modern compounds ending in the Greek masculine carpos (or carpiis) are masculine. Those in -carpa or -carpaea, however, are feminine, e. g. CalKcarpa and Polycarpaea; and those in -carpon, -carpum or -carpium are neuter, e. g. Folycarpon, Ormocarpum and Pisocarpimn. (3) Arbitrarily formed generic names or vernacular names used as generic names take the gender assigned to them by their au- thors. Where the original author has failed to indicate the gender, the next subsequent author has the right of choice. Examples: Taonabo Aubl. (Hist. PI. Guiane, i, 569: 1775) is feminine; Aublet's two species were T. deniata and T. puncta,ia. — Agati Adans. (Fam. ii, 326: 1763) was published without indication of gender: the feminine gender was assigned to it by Desvaux {Jovrn. Bat. i, 120: 1813), who was the first subsequent author to adopt the name, and iiis choice is decisive. Section 15. — Various Recommendations (Rec. XLV-L). XLV. Wlien writing in modern languages botanists should use Latin scientific names or those immediately derived from them, in preference to names of another kind or origin (popular names). They should avoid the use of the latter unless these are very clear and in common use. XLVI. Every friend of science should oppose the introduction into a modern language of names of plants which are not already there, unless they are derived from Latin botanical names by means of some slight alteration. XL VII. Only the metric system should be used in botany for reckoning weights and measures. The foot, inch, line, pound, ounce, etc., should be rigorously ex- cluded from scientific language. Altitude, depth, rapidity, etc., should be measured in metres. Fathoms, knots, miles, etc., are terms which should disappear from scientific language. XL VIII. Very minute dimensions should be reckoned in /i (micromillimetres, microns, or thousandths of a millimetre) and not in fractions of millimetres or of lines, etc. ; fractions encumbered with cipiiers and commas easily give rise to mis- takes. XLIX. Authors should indicate clearly and precisely the scale of the figures which they publish. L. Temperatures should be expressed in degrees of the centigrade thermometer of Celsius. Chapter IV. — Interpretation and Modification op the Rules (Art. 73, 74). Art. 73. A small permanent International Executive Committee is established with functions including the following:— (1) Interpreting the Rules in doubtful cases, and issuing con- sidered "Opinions" on the basis of the evidence submitted. (2) Considering Nomina co7iservanda, Nomina amhigua, Nomina dubia and Nomina confusa, and making recommendations thereon to the next International Botanical Congress. 96 MEDICAL MYCOLOGY (3) Considering all proposals for the modification of the Rules and reporting thereon to the next Congress. (4) Reporting on the effects of modifications of the Rules accepted at the preceding Congress. Art. 74. These Rules can be modified only by competent persons at an International Botanical Congress convened for the express purpose. Modifications accepted at one Congress remain on trial until the next Congress, at which they will receive sanction unless undesirable conse- quences, reported to the Executive Committee, show need for further amendment or rejection. *Appendix I. — Regulations for determining types. *Appendix II. — Nomina conservanda familianim. Appendix III. — Nomina generica conservanda. *Appendix IV. — Nomina ambigua. *Appendix V. — Nomina dubia. *Appendix VI. — Nomina confusa. *Appendix VII. — Representative Botanical Institutions recognized under Art. 34. Appendix VIII. — Nomenclature of Garden Plants. ♦Drafts of these Appendixes will be prepared for submission to the next Inter- national Congress. CHAPTER VII MUCORALES In the Mucorales or Zygomycetes, the thallus is usually coenocytic, al- though in some of the higher forms it is secondarily divided into cells. Under certain conditions the hyphae may fragment into hyphal bodies, etc., which occasionally develop further as sprout mycelia. In an extremely unfavorable environment, small portions of hyphae develop into thick-walled chlamydo- spores. In most forms the reproductive structures are aerial. The haploid my- celium produces sporangia or conidia, the diploid mycelium produces zygo- spores. Sporangia are typical of the Mucoraceae and the Endogonaceae, conidia of the Entomophthoraceae, a group of parasites upon insects. The sporangia form nonmotile, endogenous sporangiospores. In the successively higher genera there is a tendency for reduction in the number of sporangio- spores produced by a sporangium until the sporangium is practically reduced to the level of a conidium which produces mycelium directly without the medium of a spore formation. The sexual act is essentially isogamous in spite of heterothallism. The higher Mucorales tend more and more toward heterogamy. The products of the sexual act, the zygospores, are primarily resting spores and have never been reported in many species. The relationships of the Mucorales are wholly obscure. Vuillemin (1886, 1912) and Lotsy (1907) connect this order through Basidioholus, a genus dividing its life cycle between the digestive tracts of beetles and frogs, to the Conjugales of the green algae and derive the other Entomophthoraceae and Mucoraceae from this genus. Davis (1903) considers in general terms the green algae, especially the isogamous forms, such as Cladophora or the Siphonales. As intermediate forms are wholly lacking and apparently such simple structures as the Mucoraceous sporangium are still unknown in other groups, phylogenetic speculation is unfruitful. The present tendency is to connect the Mucorales with the more primitive Phycomycetes. It is quite po.ssible that the Mucorales include phylogenetically heterogeneous organisms, which are only similar in the copulation of their coenocytic gametangia. In their classification the Mucorales are divided into two groups, one of which forms sporangia, the other conidia. The sporangia! group includes the large family of Mucoraceae and a small family, the Endogonaceae, consisting of small, truffle-like fungi usually growing under leaf mold or in the soil. The conidial group contains the Entomophthoraceae consisting of two tribes, the Basidioboleae found in the intestinal tracts of beetles and amphibia and the Entomophthoreae mostly parasitic on living insects, although capable of 97 98 MEDICAL MYCOLOGY continuing growth and reproduction saprophytically on dead insects. Only the Mucoraceae have been found parasitic on mammals. Basidioholus hominis has recently been reported but has been so poorly described that its relation- ships are still obscure. Mucoraceae. — This family is commonly saprophytic on plant or animal remains, more rarely parasitic on other Mucoraceae, on higher plants, or on animals. They play a large part in the decay of organic substances and a fow species have some economic importance because of fermentations, such as alcoholic fermentation by Mucor javanicns or starch hydrolysis by Rhizopns Oryzae (Wehmer 1907). Except in Haplos%)orangium, where the hyphae are early divided into multinucleate segments by septa, the thallus consists of branched coenocytic hyphae without septa ; in senescence or in the development of reproductive structures, septa are formed irregularly to cut off older vacuolate sections from the younger portions. Furthermore, the hyphae of some genera, in high concentrations of sugars and anaerobic conditions, break up into oidia which may develop further by sprouting ; this sprout mycelium may ferment sugars very much as the true yeasts. (Fig. 2.) In Mortierella and Syncephalis, hyphal branches may fuse where they come in contact with each other, so that the mycelium becomes an anastomosing network. In general, heterothallic species have no definite sexual dimorphism, the strains usually differing slightly in physiologic characters or the positive (female) strain being better developed. Only a few details are known concerning the internal structure of the hyphae. The hyaloplasm of Mortierella reticulata and Rhizopus nigricans (Moreau 1913) contracts into peculiar strands parallel to the hyphal axis. The nuclei are very small throughout (1-3/a in diameter). They divide simul- taneously, both directly and indirectly, in the same hyphal region. The hyphae generally spread out evenly within and upon the substrate. Bhizopus and Absidia have more or less well-differentiated stolons, each con- sisting of a node provided with appressoria or holdfasts, from which radiate new stolons. The appressoria of the forms parasitic on other Mucoraceae are further modified; thus in Mortierella Bainieri they grasp the host hyphae as claws or spirals, and in Piptocephalis Freseniana, they penetrate the interior of the hypha and there branch into a small tuft, as haustoria. Thick-Avalled hypnospores are formed under unfavorable conditions, while in Mucor sphaerosporus the mycelium may form true sclerotia. The hypno- spores usually arise endogenously ; multinucleate protoplasmic portions of varying circumference draw together and, inside the original hyphal mem- brane, surround themselves with a special thick wall (Fig. 3). The stipitate hypnospores (mycelial conidia or stylospores) of Mortierella (Fig. 8, d) and Syncephalis are cut off in scattered or racemose groups on short branches of the mycelium (H. Bachmann, 1900). Under suitable environmental conditions both hypnospores and sclerotia develop to new mycelia. Asexual reproduction takes place through sporangia with sporangiospores. The parts of the mycelium from which the sporangia develop swell consider- MUCORALES 99 ably, and their nuclei divide repeatedly. In forms with stolons, the sporangio- phores branch almost exclusively from the nodes and are then firmly attached to the substrate by the group of rhizoids, but in Ahsidia they may branch directly from the stolons midway between nodes. In the simpler species the sporangiophores are unbranched (Fig. 4, 7) ; in the more highly organized, they are forked, racemose, corymbose, cincinnal, etc. (Fig. 4, 3, 9). The tip of each branch swells up to a sporangium, allowing Fig-. 4. — Sporangia. 1, Ahsidia Truchisi (after Lucet & Costantin) ; 2, Pirella circinans (after Bainier 1883) ; 3, i, Circinella umhellata, showing dehiscence and columella (after Tieghem 1873); 5, Mucor Mucedo (after Brefeld 1872); 6, Rhisopus niger (after Bainier): 7, Rhisoims reflexus (after Bainier 1883) ; S, Rhizopus parasHicus (after Costantin 1900) ; 9, Sporodinia grandis (after Lendner 1908). the protoplasm of the swollen hyphal portion to migrate into it, and is finally abjointed (cut off by a septum). In the tribe, Mortierelleae, the septum is plane or slightly convex, in the Mucoreae it forms a dome projecting far into the sporangium. This dome is usually called a columella. It is generally smooth, cylindric or pyriform and remains attached to the stalk long after the spores are shed. In Piloholus roridus (Tieghem 1875) and Pilaira anomala 100 MEDICAL MYCOLOGY (Brefeld 1881) on dung-, the sporangiopliore swells under the sporangium to a large head. On the absorption of more water, the sporangiopliore bursts at the point of insertion of the columella, shooting off the sporangium and columella often to a height of one meter and with an audible sound. In the Mucoreae the sporangial wall consists fundamentally of cellulose which subsequently is so inerusted with calcium oxalate crystals that it be- comes fragile ; simultaneously the cellulose is hydrolyzed to a more soluble hygroscopic compound, so that it finally dissolves and the crystals scatter. In the majority of genera, only the base of the sporangial wall remains, form- Fig. 5. — Showing the development of the sporangium of Sporodinia grandis. (After Harper 1899.) ing a basal collar about the columella. In the Mortierelleae the wall is equally soluble, but the oxalate crystals are not formed. In Piloholus it is cuticular- ized, except at the base, and permanent. At first, most of the cytologic processes within the sporangium are the same (Fig. 5, i). The content of the young swelling is divided into a central zone, filled mainly by sap and penetrated by a few protoplasmic threads, and a rich peripheral zone, containing most of the nuclei (Fig. 5, 2). The border between the two is differentiated into a foamy protoplasmic layer permeated by narrow, flattened vacuoles. These fuse laterally and form between the vacuolate central portion and the protoplasmic periphery a cleavage cavity; its bordering surfaces are covered with a plasma membrane which is thickened MUCORALES 101 into a wall on the side next the stalk. The central portion remains sterile and becomes the columella (Fig. 5, 4). Its nuclei no longer show any membrane or nucleoli and degenerate, although occasionally fusions or amitotic divisions appear. The peripheral, spherical cap forms the fertile sporogenous layer. There are three types of spore formation. In Piloholus crystallmus (P. microsporus) and P. oediinis (Harper 1899) vacuoles divide the whole sporog- enous protoplasm into uni-, rarely multinucleate, portions, the so-called proto- spores, which round up, swell, and undergo several nuclear divisions before separating into multinucleate portions. These portions again round off, are surrounded by a membrane, and become 2-spored. "We shall find suggestions of this mode of spore formation in the Endomycetales. {Coccidioides and Profomyces, pp. 147, 157.) In Circinnella conica (Moreau 1913) and C. minor (Schwarze 1922) de- velopment is simpler ; here the protospores are surrounded by a membrane with- out further splitting and become spores directly. In most of the other genera, as in Sporodinia (Hai-per 1899), Phycomyces, Ehizopus, Mucor, Ahsidia, and Zygorhynchus (Swingle 1903, Moreau 1913, Green 1927), the protospore stage is omitted. The division of the nuclei in Phycomyces, probably also in the other genera, occurs simultaneously in the whole sporangium. The sporog- enous protoplasm splits directly into multinucleate, rarely uninucleate, por- tions Avhich round off, form a membrane, and develop directly into spores without further nuclear division. The unicellular sporangiospores are gen erally ellipsoid or spherical, hyaline or dully colored; resting free in the sporangium or embedded in a granular gel, probably developed from within themselves, which swells rapidly in water. At germination they swell con- siderably and develop into a mycelium through one or more germ tubes. The original sporangial type discussed above, represented by Mucor, Sporodinia, etc., is modified in higher forms. Either the individualization of protospores is retarded without being suppressed or the sporangia decrease successively in differentiation, size, and spore number until they appear and function as conidia. By retardation of spore formation we have a series from Choanephora to PiptocephaJis. Poorly nourished individuals of Choanephora cucurbitarum exhibit sporangiophores and sporangia like those of Mucor. On the ends of the brown, smooth sporangiospores are 2-3 hyaline processes from each of which as many as 20 hairs may arise. AVhere there is abundant food supply, the spores develop, either on swollen tips of vertical hyphae or on the short secondary branches, exogenously by budding not endogenously by cleavage. Spore formation is ontogenetically retarded and transferred from the interior of the sporangium to its surface. Exogenous spore formation appears more clearly in CunninghameUa in- vestigated by Moreau (1913). In C. echinulata and C. Beriholletiae, the ends of the sporophores swell to sporangia whose content is differentiated into a watery inner, and a rich outer, zone. The peripheral layer pushes out into small spherical sacs on short sterigmata, with 3-8 nuclei each. These sacs are ]02 MEDICAL MYCOLOGY ab jointed and transi'ormed into spores, corresponding in size and form to those of Mncor but borne on the outer, rather than on the inner, surface of sporangia. The series may be continued in Blakeslea trispora (Fig, 6). Under certain environmental conditions, e. g., saturated atmosphere, this species forms typical multispored sporangia with large pyriform columellas (Fig. 6, 1). These sporangia show a marked tendency toward degeneration: with a slight alteration of cultural conditions, they decrease in size and spore number, and Fig. 6. — Blakeslea trispora. Modifications of sporangia: 1, original form; Z, reduced form without columella ; 3-5, formation of exogenous sporangioles ; 6, sporangiospore from sporan- giole. {1-5 X260; 6 X720.) (After Thaxter 1914.) the columella shrinks or disappears. The resulting forms only distantly re- semble the original sporangium (Fig. 6, 2). Where conditions of growth are normal, the spore protoplasm migrates into protrusions, borne on spherical sterigmata (Fig. 6, 3). Meridianal fission divides each of these into 3 spores adorned with little apical tufts of hair (Fig. 6, 5) . The mature protuberance separates from its sterigma or with its sterigma from the sporangium and is disseminated (Fig. 6, 4). Here spore formation is further retarded; between MUCORALES 103 the differentiation of sporangial content into a sterile and a fertile zone and the individualization of single spores, the sporangium also develops into numerous "partial sporangia," each of which forms a .small number of sporangiospores. In Syncephalastrum the ability to form sporangia of the Mucor type has entirely disappeared, and the extramatrical partial sporangia have reached a higher stage of development (Fig. 7). Several palmately joined sterigmata develop into long cylindric tubes which receive as many as 20 nuclei each. 9 8)0 0® ^J'm SS ©% V Fig-. 7. — Syncephalastrum cinereum. Development of extramatrical partial sporangia. (X950.) (After Moreau 1914.) When these have reached their full length, their content splits into uni- or multinucleate portions which round off and are surrounded by walls. The spores are finally liberated by the dissolution of the sporangial wall (Thaxter 1897, Moreau 1913). In Syncephalis, after the destruction of the partial sporangium, the spores remain connected with the adjacent cufflike part of the sporangial wall; the spore wall itself remains thin and insignificant while the sporangial wall is thick and occasionally sculptured. In S. aurantiaca, the partial sporangia divide by septa into as many locules as there are spores. When these septa 104 MEDICAL MYCOLOGY split, the partial sporangia divide into oidial members, each of which contains a spore ; thus the spores are completely surrounded by a sporangial wall, in- separable from their own wall. The functionless sporangium remains in open connection with the sporifer- ous hypha ; it remains capitate and does not collapse until after the maturing of the partial sporangia. It may be regarded as a degenerate form only in so far as the sterile inner part is no longer separated from the peripheral spore protoplasm by the columellar wall. In Piptoceplialus, the sporangium loses its capitate form, and shrinks to a verrucose basal cell, with an apical partial sporangium which the degeneration of the basal cell frees from the sporiferous hypha (Brefeld 1872, Tieghem 1875). The partial sporangia break up into monosporous members whose sporangial wall is fused with the spore membrane. The sporangiophores, therefore, have become conidiophores, rec- ognizable as sporangiophores only through their phylogeny. In the Thamnidium-Chaetocladium series, the sporangiospores are nu- merically much reduced, their functions being assumed by sporangia which successively degenerate to conidia. In Tliamnidium elegans, the main axis possesses an apical multispored sporangium which has a columella. Under certain conditions, dichotomous branches terminating in sporangia are formed from the main axis. These sporangia, however, are smaller than the terminal ones, have no columella, become loosened as a whole from the sporangiophores, and contain few, generally 4, spores. Spores are liberated not by deliquescense but by disintegration of the sporangial wall. kSuch reduced sporangia are called sporangioles. The spores in both types of sporangia behave similarly as regards germination and further development. "When well nourished, the sporangioles persist through several generations and finally become as large and multispored as the sporangia. Conversely, with poor food supply a termi- nal sporangium may turn into a sporangiole, often containing but one spore. This line of development is continued through Chaetostylum Fresenii (Thamnidiuni chaetocladioides) . Here true sporangia are borne only with adequate nourishment, while Avhere the food supply is limited, the terminal sporangia abort. The terminal sporangia which have declined in these two species, disap- pear in Chaetocladium, where the sporangioles also degenerate. They become monosporous, the spore membranes fusing with sporangial walls. In Chaeto- cladium Jonesii this double nature of the spore wall is evident, for on germina- tion the sporangial wall separates from the spore. In C. Brefeldii, however, this differentiation disappears, and the germ tubes protrude directly from the wall. Again the sporangium has been transformed to a conidium. In another series, Mortierella and Haplosporangium show a similar degen- eration. The sporangium of Mortierella is separated from the sporangiophore by a septum (Fig. 8). Since its contents are not differentiated into fertile and sterile zones, the spores arise directly by cleavage of the whole proto- plasm. Their number is notably reduced, in some species to 2 or 4. Haplo- MUCORALES 105 sporangium bisporale (Thaxter 1914) exhibits this condition as a general rule. The sporangia remain very small and retain only 1 or 2 spores. The spore wall is delicate, the sporangial wall thick and sculptured. The same process of reduction as in the unrelated Thamnidium and Chaetocladium has occurred here and has led to the formation of 1- or 2-spored sporangioles, biologically functioning as conidia. Both sexual and asexual reproduction are known in most Mucoraceae. The type of reproduction in the homothallic forms depends mainly on con- ditions of nutrition ; the heterothallic forms require the presence of both sexes. Mycelia of one sex may be cultivated alone indefinitely without the appear- ance of normal sexual reproduction, which appears promptly whenever the opposite sex is brought into the vicinity. Fig. 8. — Mortierella niveovelutina. a, g, hyphal anastomosis ; h, spoiangiospore ; c, sporangia ; d, stylospores or aerial conidia ; e, chlamydospores ; /, sporangia attached to sporan- giophores. When the environmental conditions are favorable, the mutual approach of two sexually mature (in heterothallic forms also dynamically opposite) hyphae results in the formation of outgrowths toward each other. Each out- growth is cut off from the hypha close behind the tip by a septum laid down from the wall inwards. The tip cell is the gametangium, the hypha is the suspensor. As the homothallic forms are bisexual, apparently there occurs in their hyphae at sexual reproduction, a spatial sepai-ation of + and - energids. In some species, the copulating branches arise from ordinary hyphae, in others they are developed on special branches, the zygophores (Fig. 9). The two separating walls between the gametangia are gradually dis- solved from the middle toward the edge, and the zygote becomes a hypnospore 106 MEDICAL MYCOLOGY by the formation of a many-layered wall. This spore is called a zygospore. In the homothallic species when the copulating branch finds no mate, the gametangium surrounds itself with a thick many-layered wall and is called an azyg-ospore or, less properly, a chlamydospore. The same thing happens Fig. 9. — Zygospores. 1, Ahsidia glauca (after Lendner) ; 2, Mucor hiemalis (after Lendner 1908) ; 3, Parasitella simplex (after Burgeff 1924) ; -i, Zygorhynchus heterogamus (after Blakeslee 1913) ; 5, Rhizopus nigricans (after Bary) ; 6, Sporodinia ginndis (after Bainier 1882) ; 7, Phycomyces nitens (after Van Tieghem & Lemonier) ; 8, Spinellus fusiger (after Bainier 1882) ; 9, Mucor racemosus (after Bainier 1883) ; 10, Circinella spinosa (after Bainier 1882). if the cultures are placed in an unfavorable environment, such as high tem- peratures. In the heterothallic forms, similar phenomena may occur if the MUCORALES 107 copulating branches belong to two different species; in this case, they cease growing and transform the; ganietniigia (in case they liave ali'(!ady been cut off as such) into azygospores. This inconii)lete hybridization, however, does not seem to occur between all species, for, while it occurs between Phycoriiycea nitens + and Mucor Mucedo - and conversely, it does not between Phycomyces nitens and Rhizopus nigricans (Blakeslee 1904-1927), While both gametangia are usually of the same size and thus externally suggest isogamy, in individual species their size relationships show a notable tendency to heterogamy. Thus, in the homothallic ZygorJiyncJins Moelleri, the copulating branches are unequally developed. In the heterothallic Absidia Orchidis the gametangia are of unequal diameter, the resulting zygospores being conic. In Piptocephalis, the zygospore grows upward from the point of fusion so that it is borne upon the top of the copulation branch. In Syncephalis Fig. 10. — Development of Uie zygospore of Sporodinia yrandis. (After Keene 1914.) nodosa, one copulation branch coils around the other in a helix (Thaxter 1897) ; the zygospore does not arise at the point of fusion but comparatively distant, on the outer portion of the helix near the septum separating the gametangium from the suspensor. Cytologically all the; above-mentioned processes behave similarly. Sporo- dinia grandis has been more complete studied. Its young gametangia con- tain more than a thousand nuclei each (Fig. 10). While the separating walls between the gametangia are dissolved, the nuclei undergo almost simultaneous division ; their cytoplasm intermingles and their nuclei subsequently pair and fuse. Those without mates, especially those near the periphery, degenerate and disappear. Meanwhile at the surface a wall of several layers has been formed, the suspensors collapse, and the zygospores presently lie free upon their substrate. It is characteristic of this process that no dynamic differentiation occurs oetween the + and - energids in spite of their spatial separation. Thus, both 108 MEDICAL MYCOLOGY gametangia are cytologically equivalent, and fertilization is isogamous with reference to the nuclei. In Sporodinia, since there is no individualization of gametes, two coenocytic gametangia copulate and accomplish multiple fertiliza- tions. In ZygorrhyncJius Dangcardi, all but 4 gamete nuclei degenerate in the young zygote. The surviving four fuse in pairs very late after the endo- spore has been formed. A similar retardation of caryogamy has been observed in Phy corny ces nit ens (Burgeff 1915) in which the nuclei in zygospores 5 months old and ready to germinate, still lie in pairs. The wall of the zygospores in the more carefully studied species of Mucor, Sporodinia, and ZygorrhyncJius consists of 5 layers (Vuillemin 1904). The in- nermost layer is thin and granular ; it forms the transition from the protoplasm and to a certain extent is the mother layer. The next is thickest and is called the cartilaginous layer on account of its elasticity. This is covered by a thin sheath, the middle cuticular layer. The fourth or carbonaceous layer is fragile and brown or black ; the outermost cuticular layer is either pale and elastic, or dark and fragile, and often interrupted or fractured. The greatest modifi- cations in the various genera are shown by the surface of the carbonaceous layer which is verrucose or reticulate. The two outer layers are grouped as the exospore, the three inner as the endospore. In Absidia and Phycomyces the zygospores are loosely surrounded by branches from the suspensors (Fig. 9, 1, 7). In Mortierella these branches intertwine with the neighboring hyphae into a solid felt whose outer surface is cuticularized and brown. AVithin this tissue lies the zygospore. The zygospores germinate only after a long resting period. The exospore is ruptured, the endospore puts forth a germ tube which develops to a mycelium or, with insufficient nourishment, directly to a sporangium or a conidiophore. During germination, meiosis of the diploid nuclei occurs. Where the germ tube becomes the fundament of a sporangium (e. g., Phycomyces nitens, Burgeff 1915) meiosis occurs only in the latter which is called a "germ sporangium," and as we shall see later is the precursor of the ascus. The sexual relationships existing at meiosis have been more closely studied for three types. {Sporodinia, Mucor Muceclo, and Phycomyces nitens, Blakeslee 1904, 1906.) In Sporodinia the sporangiospores are liomothallic and the separation of the + and - energids occurs only in the formation of the copulation branches. In the heterothallic Mucor Mucedo the separation of the + and - energids occurs probably in the formation of sporangia ; i.e., the spores are all of one sex in one sporangium, either all + or all -. In the equally heterothallic Phycomyces nitens, the separation of sexes occurs only in the formation of spores. Even so, it is incomplete ; besides the + and - spores there are also unstable, neutral, bisexual spores in whose sporangia the separation into + and - spores is continued (Burgeff 1912). MUCORALES 109 Although some of the following characters are often unstable, they should be noted in the study of the Mucorales. The presence or absence of branching is often difficult to determine. Sometimes typical branching may be found in the small sporangiophores next the substrate, when it is not observed in the larger sporangiophores. If stolons arc present, note their arrangement and the disposition of the sporangiophores upon them ; also note the presence of holdfasts, etc. The nature of the medium influences the height of the spo- rangiophore, which should be determined only in cultures where optimum con- ditions of growth prevail. Malt gelatin (10%) or 10% gelatin to which has been added the residue of white wine from which the alcohol has been distilled, are suitable. The latter is known as Lendner's medium. Report the height of the spo- rangiophore from a colony cultivated at room temperature for at least 8 days, its diameter, the diameter of the sporangium (one of the larger ones), the height and diameter of the columella, the mean diameter of the spores (or their mean dimensions), and the diameter of zygospores and chlamydospores. The spo- rangial membrane may be diffluent, in which case, younger sporangia should be measured, or the sporangia mounted in a mixture of glycerol and water. If the membrane easily becomes fragmented, this should be noted. Note the presence or absence of a collar about the columella and the surface of the latter. Spore shape varies in the same sporangium. When a species is re- ported as having spherical spores, the majority are spherical, although oval or irregular spores may be present. Disregard variations in size unless they are extreme. To find hypnospores use cultures 2 weeks old or more on solid media or on liquid media with much sugar for sprouting cells. Note fermenta- tions in case the sprout cells are abundant. Classification. — There is still considerable disagreement among mycologists as to the subdivisions of this order. It is clearly divisible into three groups which the older mycologists considered families (Mucoraceae, Endogonaeeae, and Entomophthoraeeae). Some of the younger generation would elevate the old families to suborders, the latter two suborders containing a single family each, while the old tribes of the Mucoraceae are elevated to family rank (Fritzpatrick 1930). In any case only members of the tribe IMucoreae and Mortierelleae have so far been reported pathogenic and need be considered here. Since many of the saprophytic genera are difficult to define, and there are strong differences of opinion on synonymy, only the pathogenic genera which have been reported pathogenic to mammals are included in the fol- lowing kej^s. MUCORACEAE Mycelium coenoeytic, forming loose felted colonies ; sporangiophores erect, often variously branched ; sporangia usually with a columella (absent in Moriierella) ; sporangiophores abundant in the Mucoreae, in other tribes often reduced to a few spores in small sporangioles ; zygospores resulting from the copulation of gametangia. 110 MEDICAL MYCOLOGY Key to Pathogenic Genera Sporangium containing a columella; zygospore not surrounded by a layer of interwoven hyphae; sporangioles not formed, sporangial wall thin, not cutinized. Miicoreae. Sporangiophores arising directly from the mycelium, suspensors lacking outgrowtlis ; gametangia essentially alike. Mucor. Sporangiophores arising from aerial arching stolons which develop rhizoids at points of contact with the substratum. Sporangiophores borne on the arching internodes of the stolons between the nodes; sporangia pyriform ; zygospores, when present, vpith prominent circinate out- growths. Absidia. Sporangiophores arising in a fascicle from the node of the stolon; sporangia spherical. Bhisopus. Sporangium lacking a columella; zygospore where known enveloped by a thick layer of in- terwoven hyphae; sporangioles and conidia formed in some cases, when present isolated, not covering an enlargement on the sporangiophore or conidiophore ; sporangiophore erect, tapering upward, usually not branched. Mortierella. MUCOR Mucor Micheli, NovaPlantarum Genera 215. 1729; Linne, Species Plan- tarum 1185, 1753; Gray, Natural Arrangement of British Plants 1: 560, 1821; Fries, Systema Myeologicum 3: 320, 1829. Type species : Mucor Mucedo L. Mycelium abundant both in and on the substratum, lacking stolons and rhizoids ; sporangiophores occurring singly, erect, simple or occasionally branched, each branch terminated by a sporangium which is large, spherical, many-spored with an evanescent sporangial wall neither cutinized nor in- crusted ; columella always present, variable in shape ; sporangiospores spheri- cal to ellipsoid, with a thin, smooth wall ; zj^gospores borne on the mycelium, suspensors lacking outgrowths ; chlamydospores present in some species termi- nal or intercalary, smooth, hyaline ; oidia accompanied by fermentation found in the submersed mycelium. At present there is little conclusive evidence that this typically saprophytic genus is pathogenic for man. Most of the cases originally attributed to this genus were based on misidentification of the organism and belong elsewhere. For descriptions of species of this genus see the systematic accounts of A. Fischer (1892), Lendner (1908), and Povah (1917). Mucor Mucedo L., Species Plantarum 1185, 1753. The case of Fiirbringer (1876) should probably be referred to Absidia corymhifera. Mucor racemosus Fresenius, Beitr. z. Mycol. 12, 1850. Pleurocystis Fresenii Bonorden, Handb. Allgem. Mykol. 124, 1851. f Mucor scarlatinosus Hallier, Zeitschr. f. Parasitenk. 1: 117-184, 290-352, Pis. 3, 4, 1869. Chlamydomucor racemosus Brefeld, Unters. Gesammtegebiet der Mykol. 8: 223, 1890. MUCORALES 111 M^icor scarlatinosus Hallier, a very poorly described organism was sup- posed to have been isolated from a case of scarlatina. It was quite probably a contamination and has been refen-ed here as a possible synonym by A. Fischer, 1892. M. racemosus was reported by Bollinger (1880) from the respiratory tract of birds but was not pathogenic for laboratory animals, by Zurn (1876) in the nasal cavity of a sheep, and by Frank (1890)' in a tumor in a horse, but both determinations doubtful. Savoure (1906) reports that it was not pathogenic for rabbits. Mucor pusillus Lindt, Arch. f. exp. Path, u Pharm. 21: 272. PL 2, Figs. 1-6, 1886. [Saprophyte.] M. ramosus Jakowski, Gazetta Lekarska No. 34: 1888. [Centralbl. Bakt. II, 5: 388, 1889] not Lindt, I.e., p. 275. The fungus reported by Jakowski from the outer ear has been referred here by Vuillemin (1904), while the original author and Barthelat (1903) refer it to Absidia ramosa (Lindt) Lendner. Since there is so little conclusive evidence of pathogenicity, the reader is referred to the systematic accounts of A. Fischer 1892, Lendner 1908, and Povah 1917 for aid in determining cultures. ABSIDIA Absidia Tieghem, Ann. Sci. Nat. Bot. VI, 4: 350, 1876. Lichtheitnia Vuillemin, Bull. Soc. Myc. France 19: 119-127, 1903. Type: Absidia capillata Tieghem. Vuillemin (1903) divided this genus into six genera of which Lichtheimia contained the parasitic fungi so far described. The characters on which the separation was based seem comparatively trivial, and this segregation has not been followed by systematists of the group, although recognized by some medical men. Mycelium forming stolons, often branched, more or less curved producing rhizoids more or less branched at the surface of the substratum; sporangio- phores erect, usually in groups of 2-5 arising from the curved part of the stolon, not from the place of origin of the rhizoids ; sporangia pyriform, erect, with an infundibulifonn apophysis, membrane neither cutinized nor incrusted, diffluent, leaving a small collar at the base ; columella hemispheric, conic, or terminated by a single projection, continuous with the apophysis which is cutinized and of deeper color than the sporangiophore ; spores small, oval usually smooth, rarely echinulate, hyaline ; zygospores formed on the stolons surrounded by circinate filaments, cutinized, growing from one or both of the suspensors. This genus differs from Rhizopus by the development of the sporangiophores from the internodes, by the pyriform sporangia, by the columella continuous with the apophysis and by the suspensors provided with circinate filaments. 112 MEDICAL MYCOLOGY Key to Pathog-enic Species Growth good at ordinary temperature Spores mostly spherical, SA/x. in diameter; columella usually somewhat spinescent A. corymbifera Spores elongate or oval, 4-5x2-3/^; columella smooth A. ramosa Growth poor at ordinary temperature, optimum about 37° C. Sporangia 36-70^, columella 60^, spores ovoid 4x2-3ja; some growth at 51° C. A. Truchisi Sporangia 30-38ai, columella 26/x, spores 3.2x3.75;U,; no growth at 51° C. A. Begnieri Absidia corymbifera (Cohn) Saccardo & Trotter in Saccardo, Sylloge Fungorum 23: 825, 1912. Mucor corymbifer Cohn in Lichtheim, Zeitschr. Klin. Med. 7: 147, Pis. 6-8, 1884; Barthelat, Ann. Parasitol. 7: 25-30, 1904. Lichtheimia corymbifera Vuillemin, C. R., Acad. Sei. Paris 136: 516, 1903. Mucor corymbifera var. typica LicJitheimi Lucet & Costantin, Arch, de Parasitol. 4: 380, 1901. Absidia Lichtheimi Lendner, Mat. Fl. Cryptog. Suisse 3: 143, 144, 1908. Many cases in the literature dealing with bronchomycosis (Paltanf 1885, etc.). Lang & Grabauer (1923) discuss the clinical and pathologic aspects fully and summarize earlier cases. This fungus has also been reported from the ear by Huckel 1884, Siebenmann 1889, and Graham 1890. Mycelium white, then clear gray, completely covering the substrate; hyphae often up to 15/x in diameter, branched, hyaline under microscope. Sporangiophores resupinate, branching as a corymb, terminated by sporangia, occasionally a few small sporangia on short pedicels. Sporangia hyaline, pyriform up to lO/x in diameter with mean 45-60/x, small sporangia 10-20'/^, wall hyaline, smooth, diffluent, often with a basal collar; columella hemispheric, 10-20/x, smooth or sometimes papillate, smoke-gray or brownish continuous with the infundibuliform apophysis, spores nearly spherical 2-4/t, smooth, hyaline, occasionally up to 6ju,. Growth has been reported good on moist bread, potato, carrot, sugar media with slightly acid reaction, and Sabouraud agar; growth poor in liquid media; unfavorable conditions of humidity or lack of oxygen cause abundant pro- duction of gemmae; growth possible at 12-15°, optimum 36°, killed at 55° C. Absidia italiana (Costantin & Perin) Dodge, comb. nov. Lichtheimia italiana Costantin & Perin. Bull. Soc. Med. Chir. di Pavia 35: 1922. Lichtheimia italica Pollacci & Nannizzi I miceti patogeni dell'uomo e degli animali 3: No. 26, 1924; Perin, Arch, di Clin, e Patol. Med. 2: 5, Oct., 1923; Trattato Micopatol. Umana 1: 55-73, 1925. I have been unable to locate any of the original descriptions of this organ- ism. Some have reduced it to synonymy with A. corymbifera. The following notes are based on Perin (1925). MUCORALES 113 Isolated from sputum and from tissue fragments from the lungs. Patho- genic for laboratory animals. Sporangiophores branched, lacking septa, primary axes resupinate, sec- ondary fertile axes erect with only two or three branches ; rhizoids variable. Sporangia up to 66 x 58/*, columella 45/a broad, varying from hemispheric to conic, pedicel 20/x in diameter. Spores slightly ovoid, 3.2-4.2 x 2-2. 8/a. Colony on agar, white, cottony, gradually becoming grayish and finally brownish. Gelatin slowly liquefied. On liquid media, small flocci in the depths, a cottony pellicle above. Milk coagulated and acidified. Absidia ramosa (Lindt) Lendner, Mat. Fl. Ciyptog. Suisse 3: 144-146, 1908. Mucor ramosus Lindt, Archiv. f. exp. Path. & Pharmakol. 21: 269, 1886. Lichtheimia ramosa Vuillemin, Arch, de Parasitol. 8: 562-572, Figs. 1-4, 1904. Not Mucor ramosus Bulliard which is a synonym of M. aspergilJus Scopoli transferred by Link 1824 to Sporodinia. Perhaps not the organism of Jakow- ski (1888) which may be M. pusillus Lindt fide Vuillemin. Reported by Jakowski (1888) in human ear. Originally isolated on damp bread, found pathogenic to laboratory animals, and reported by Vuillemin from lesions and mucus of the nose in horses, also adenitis of lower jaw. Found in generalized infection in swine by M. Christiansen (1922) and in cases of abortion in cows by Bendixen & Plnm (1929). Sporangiophores branched as in A. Corymhifera, usually lacking cross- walls. Primary axes resupinate like stolons, but not recurved. Fertile axes little branched, with fewer umbels, especially compound umbels, than in A. corymtifera. Primary axes bear rhizoids frequently instead of terminal spo- rangia. Rhizoids variable. Sporangia much as in A. corymhifera, the diffluent membrane covered with fine granulations. Spores elongate, oval, or sub- cylindric, 4.8 x 2.8ju, (4.6 x 2.6, 5.2 x 3/x) brownish yellow. Columella very rarely conic, mostly ovoid, not spinescent, 57.5 x 4/x where it separates from the apophysis, 35/* in maximum diameter; blue, darkening with age. Distinguished from A. corymhifera by nonspinescent columella and sub- cylindric spores. Near A. diibia, intermediate between the subgenera Lich- iheimia and Tieghemella. Var. Rasti Lendner, Mat. Fl. Cryptog. Suisse 3: 146, 1908. Rising 1 cm. at the most above the substrate, mycelium constantly bluish gray, sporangia more abundant, present everywhere, larger, spores slightly more elliptic, often abnormal in shape. Var. Zurcheri Lendner, Mat. Fl. Cryptog. Suisse 3: 146, 1908. Rising to 4 cm. above the substrate, mycelium pure white, culture less vigorous, sporangia formed only at the surface, the deeper filaments remain- ing pure white, spores not abnormal in shape. Both varieties grow well on potato at 45° C. Absidia Regnieri (Lucet & Costantin) Lendner, Mat. Fl. Cryptog. Suisse 3: 146, 147, 1908. 114 MEDICAL MYCOLOGY Mucor Reg7neri Lucet & Costantin, Arch, de Parasitol. 4: 362-384, 1901; Barthelat, Ann. Parasitol. 7: 34, 35, 1904. Lichtheimia Regnieri Vuillemin, C. R. Acad. Sci. (Paris) 136: 516, 1903. Isolated from another stable with an environment similar to that of A. Truckisi, pathogenic for rabbit. Mycelium lax, weak vegetative growth, of a uniform color of gray slightly tinged with blue. Sporangiophores in corymb or umbel, the outer rays longer, unequal, swollen below the sporangium so that the collar seems to divide the columella into two parts, 3-8/x, in diameter, sometimes up to 12.5 or even 19/*. Sporangial membrane smooth and transparent leaves little trace of a collar. Columella ovoid, pyriform, with a clear, brown color which ex- tends a certain distance down the sporangiophore, frequently small W.lfi, sometimes 23/* and rarely up to 35/*. Spores usually round, mean 3.2 to 3.75, some 2.5/*. Besides the typical spores some are avoid (3.8 x 5, 3.2 x 2.9/*) some are irregular to almost polyhedral. Zygospores unknown. The above characters were obtained at 25° C. on solid media. Below 20° growth normal, sporangia abundant, becoming 30-38/* and pedicels 3.8 to 6.5/*. At 51-52°, slight or no growth ; pedicels simple, sporangia 19/*, spores few. Killed at 55-56°. Absidia Tnichisi (Lucet & Costantin) Lendner, Mat. Fl. Cryptog. Suisse 3: 146, 1908. Mucor Truchisi, Lucet & Costantin, Arch, de Parasitol. 4: 362-384, 1901. Barthelat, lUd. 7: 31-34, 1904. Mucor corymhifer Cohn, var. Truchisi Sartory, Champ. Paras. Homme Anim. 91, 1921. Isolated from a hoi-se infected with Trichophyton minimum. Pathogenic for rabbits. Mycelium lax and in general vigorous, whitish or very light gray, becom- ing darker in old cultures on solid media. Sporangiophores in corymbs or in an umbel with branches of unequal length, the outer usually longer, swollen below the sporangium so that the collar seems to divide the columella into two parts, 2-7/* thick, secondary axes 55-195/*; sporangia spherical with trans- lucent membrane, smooth, about 35/* in diameter, spores regularly ovoid to slightly elongate, mean 4 x 2.5/*, smaller 3.75 x 2.5/*, and larger 4.5 x 3/*. Columella pyriform, brown at the base, becoming lighter toward the apex; mean breadth 20-26/*, smaller 4-15, larger 30/*. Zygospores unknown. At low temperature, 10-18°, mycelium little developed, fine and delicate ; fructifica- tion little developed, pedicels always simple, sporangia up to 70/t, and spo- rangiophores 14/* in diameter. At 51-52° C. for 5 days, growth rich, fruiting abundant. Sporangia 26/*, columella 17-19/*, pedicels 5-6/*, spores 5 x 2.5- 3/*. At 53° for 17 days growth still noticeable; killed at 55-56°. Growth very good on raw potato. Absidia cornealis (V. Cavara & Saccardo) Dodge, n. comb. Mucor cornealis V. Cavara & Saccardo in V. Cavara, Ann, di OttalmoL 42: 650-674, 1 pi, 1913, MUCORALES 115 Producing lesions in the cornea, Italy. Mycelium loosely interwoven, white, cinereous-plumbeous on milk, bread, and potato; growth abundant at 37° C, very slow at 15° C, and at 51° C. ; sterile hypliae large, branched, 14-15/* in diameter, hyaline, corymbose or racemose branched at the tips ; branches of sporangiophores either alternate or opposite, simple or dichotomous, 80-300 x 7-8/i., leaving the main axis at about an angle of 45° slightly enlarged at the tips; sporangia spheric or sub- spheric with a thin membrane 40-44/t in diameter (rarely up to 50-55/a or as small as 15-22/a), columella obovate, pyriform, more or less light fuscous, 22-24/x broad ; spores thin-walled, hyaline becoming light yellowish, spherical, 4-4. 5/a, rarely ovoid, zygospores unknown. RHIZOPUS Rhizopus Bhrenberg, Nova Acta Acad. Leopold. 10: 198, 1820. Bhizomucor Lucet & Costantin, Rev. Gen. Bot. 12 : 81, 1900. The type species is Rhizopus nigricans Ehrenberg. The type of Rhizomucor is R. parasiticus Lucet & Costantin. Aerial mycelium of creeping stolons, with holdfasts at the nodes which at- tach the hyphae to the substrate. Sporangiophores arising in groups at the nodes, sometimes solitary, enlarged above into a columella, as in Absidia. Spo- rangia white at first, becoming black ; spherical or nearly so with base slightly flattened ; membrane not cuticular, uniformly incrusted and entirely diffluent without leaving a basal collar. Columella hemispheric, often flattening after dehiscence, suggesting the pileus of a mushroom. Spores spherical or ovoid, even angular, hyaline or brownish, cuticular walls, smooth or striate, rarely spinulose. Zygospores without covering from outgrowth of suspensors, form- ing in the substrate and on the stolons. Suspensors straight, swollen, without appendages. Key to Pathog-enic Species Spores irregular, angular, subspheric, oval. B. parasiticus. Spores spherical, smooth or echinulate, but not angular. Columella conici or subcylindric (black tongue). E. mger. Columella ovoid or pyriform, pathogenic for rabbit. Clilamydospores not produced. E. rhizopodiformis. Chlamydospores present. E. equinus. Rhizopus equinus Costantin & Lucet, Bull. Soc. Myc. France 19: 200, 1903. Isolated from a horse, pathogenic for rabbit. Found in generalized in- fection in swine by M. Christiansen (1922) and in bovine fetal membranes and fetus by Theobald Smith (1920) who referred his species to R. rhizo- podiformis. Mycelium at first white, then gray after the formation of sporangia. Spo- rangiophores at first isolated and without rhizoids, straight or curved, later in bouquets, frequently provided with rhizoids, cutinized, pale ochraceous ; 50-220/i, sometimes up to 600/x long, 3-12.3/a in diameter. Sporangia 30-115/x in di- 116 MEDICAL MYCOLOGY ameter. Columella 45-51 x 31-41ju. Spores round, sometimes slightly an^lar, smooth, 4/A. Chlamydospores eitriform, 30 x 25 or 40 x 26/a, or spherical, 20^16 in diameter, forming ordinarily on the mycelium. Zygospores unknown. Var. annamensis, P. N. Bernard, Bull. Soc. Myc. France 30: 230-232, PI. 14, 1914; Bull. Soc. Path. Exot. 7: 430, 1914. Isolated from sputum of an Annamite, aged thirty-two, 32 in 1911; sputum blackish, as if mixed with carbon grains. Cough with a little dyspnea. Whole left lung infected. No previous history except bronchitis, with com- plete recovery. Hospitalized for cough in fall, 1910, worse July, 1911, but alive in December, disease seemingly arrested and localized. Intravenous inoculations in pigeons, no effect ; nor intraperitoneal, in guinea pig ; rabbit succumbed by both methods, but not by subcutaneous inoculation. Organ- ism recovered. Isolated sporangiophores without rhizoids 72-450'/a x 8-12/^ usually 150 x 12/x. Sporangiophores in pairs near each other, pedicels short 78-210 x 8 - 12/A, 30-45/x between pedicels, without rhizoids. Sporangiophores with rhi- zoids singly or in pairs, the latter 138-144 x 8-9/a, occasionally pedicels up to 420-780/i long. Sporangia oblate spheroid, 48-84/a. Columella 18-48|U high X 24-52/i broad. Cutinization from pedicels to rhizoids and stolons, less ac- centuated on the columella. No collar after sporangial dehiscence. Inter- calary chlamydospores 36 x 24/* eitriform ; or spherical 30-42/* in diameter ; ovoid 60 x 48-42 x 30/t, numerous even in aerial portions. Spores round, smooth, 4/t, not cutinized. Growth good on Sabouraud agar for several months, then suddenly stopped fruiting on carrot and potato glucose; optimum 37-39° C, tube filled with mycelium in 3-4 days; sporangia in 5 days; no growth at 5° C. ; killed at 100° moist heat in 15-20 minutes. Rhizopus niger (Ciaglinski & Hewelke) Barthelat, Mucorinees patho- genes et les Mucormycoses 55, 56, 1903; Arch, de Parasitol. 7: 46, 47, 1903. Miicor niger Ciaglinski & Hewelke, Zeitschr. Klin. Med. 22: 626, 1893. Isolated in cases of black tongue ; not pathogenic for guinea pigs or rab- bits. Also found later by Sendziak (1894). Stolons provided with numerous rhizoids, forming a snow-white layer. Sporangiophores erect, straight, fasciculate, terminated by spherical sporangia, which become black at maturity. Columella at first cylindric 2-3 times as long as wide, later enlarging and becoming hemispheric ; after the dehiscence of the sporangium assuming the appearance of an open umbrella. Spores oval, smooth, gray, black in mass. Growth good on potato and in bread gelatin, optimum 25-27° C, growth ceases at 37° C. Rhizopus parasiticus (Lucet & Costantin) Lendner, Mat. Fl. Cryptog. Suisse 3: 115, 1908. Bhizomucor parasiticus Lucet & Costantin, Rev. Gen. Bot. 12: 81, PI. 3, 1900; Arch, de Parasitol. 4: 384-408, 1901. [See p. 394.] MUCORALES 117 Mycosis of lung of coiintiy woman, final recovery after several months of treatment with arsenic. See Arch, de Parasitol. 4: 386-389, 1901, for case history and results of inoculations. Nonpathogenic in subcutaneous inocu- lations. Mycelium gray (lead color) to mouse gray then grayish brown to yellow. Stolons and rhizoids irregular. Sporangiophores branched in simple clusters, or in corymbs 12-14/x in diameter, 1-2 cm. long ; sporangia 35-80/x,, with mem- brane covered with fine crystalline needles; columella ovoid, pyriform, cutin- ized slightly brownish, 30-70//, long by 24-56/a in diameter; lateral sporangia similar but much smaller ; pedicels rarely ramifying a second time ; spores irregular or reniform, smooth, 4 x 2.5/x. Zygospores unknown. Growth on most media very good, but less on peptone broth and on very acid or alkaline media, poor on coagulated sera, amniotic liquid, white of egg, cider, apples, or pears, the latter, however, is good if glycerin or glucose is added. Growth more rapid on solid than on liquid media. [Very full description given in Arch, de Parasitol. 4: 384-408, 1901.] Optimum about 37° C. ; growth starts at 22°, at 51-52° very abnormal vegetative growth but no spores. It needs much oxygen. Rhizopus rhizopodiformis (Cohn) Zopf, Die Pilze 317, 1890. Mucor rhizopodiformis Cohn in Lichtheim, Zeitschr. Klin. Med. 7: 148, 1884. Rhizopus Cohnii Berlese & de Toni in Saccardo, Syll. Fung. 7: 213, 1888. Pathogenic for the rabbit Avhen injected into peritoneum and veins. Ziegenhom (1886) was unable to modify pathogenicity of spores. T. Smith (1920) reports this organism on membranes and in lungs and digestive tracts of fetus but probably it was R. equinns. Mycelium white, then mouse-gray, rising as a spider web above the sub- strate. Stolons forming rhizoids at point of contact with substrate, in brown- ish bouquets. Sporangiophores isolated or grouped, erect or incurved, short, 120-125/i, not branched, with brownish membrane enlarging in an apophysis. Sporangia spherical, 60-110/t usually about 66/a, blackish at maturity, smooth with incrusted membranes. Columella forming with the apophysis an ovoid or pyriform organ, 50-75/i broad, membrane smooth and brownish. Spores usually spherical, small, 5-6/x, without angles, smooth, hyaline. Zygospores and chlamydospores unknown. Cultural characters are similar to those of Absidia corynibifera (p. 112). Optimum temperature 37-38° C, sporangia after 48 hours, mycelium changing from white to gray. At 12-15° spores germinate on third day, sporangia on fourth or fifth day. At 45° the mycelium is arrested and spores are killed at 68°. Doubtful Position Rhizomucor septatus (Bezold in Siebenmann) Lucet & Costantin, Arch, de Para.sitol. 4: 362, 1901; Barthelat, Mucorinees pathogenes et les mucormy- coses 52, 1903. Mucor septatus Bezold in Siebenmann, Die Schimmelmycosen 97. Wies- baden, 1889. 118 MEDICAL MYCOLOGY Mycelium colorless, sporangiophores brown, in branching cluster, some- times terminating in an umbel, with small rhizoids at the base with a mecii diameter of lO/i,; secondary pedicels, 3-4 in number, are short with crosswalls at point of branching; sporangia pale grayish brown, spherical, transparent membrane, smooth or slightly papillate, 32/x, in diameter ; columella also brown, spherical or slightly ovoid, mean diameter of 27/*; spores spherical or ovoid, smooth, clear yellowish or brownish, 2.5 - 4/a. Referred to Mucor racemosus by A. Fischer, and Lendner. It also resembles M. hifidus Fresenius. It differs from R. parasiticus in smooth sporangial wall devoid of crystals, sporangia smaller, and sporangiophores always septate. MORTIERELLA Mortierella Coemans, Bull. Acad. Sei. Belgique II, 15: 536, 1863. The type species is Mortierella polycephala Coemans. Mycelium within the substrate or forming a closely appressed weft over its surface, not typically aerial; sporangiophores erect, simple or branched, usually tapering to a delicate tip just below the sporangium, often swollen below; sporangia spherical without columella, wall soon disappearing; stylo- spores unicellular, spherical, echinulate, suggesting sporangia with a single spore; zygospores enveloped by a thick layer of densely woven hyphae which arise just below the gametangia and tend to obscure the details of conjugation. Mortierella niveovelutina Ciferri & Ashford, Porto Rico Jour. Publ. Health & Trop. Med. 5: 134-143, 1 pi, 1929. Isolated from a Porto Rican with a patch of inflamed papules covering the antero-extemal aspect of the right thigh and extending around behind, ending below at the insertion of the adductor muscles. At first glance the appearance reminded one of psoriasis, but when fading, the eruption became discrete and parts once thickly studded with red nodules disappeared without leaving a trace of the former thickly infiltrated red nodular area. Intense pruritus present. Healed by the application of salicylic acid, ichthyol, and sulphur ointment. Colony white, velvety ; mycelium of highly branched hyaline hyphae with apical branches normally bifurcate, occasionally Avith three forks, continuous or later scantily septate, with frequent and complex anastomoses, without rhizoids, 2-3/i. in diameter; hypnospores in chains in liquid media, less abun- dant in solid media, from spherical to elongate with a smooth membrane ; stylospores or aerial hypnospores normally very abundant, singly or in chains of up to 12 cells, generally 2-3/* in diameter with a smooth epispore ; sporangio- phores 30-80/1 long, straight erect, of uniform diameter, never branched, con- taining an apical septum immediately beneath the sporangium ; a single spo- rangium for each sporangiophore, approximately spherical, 30-90/1 in diameter normally about 60/i, irregularly dehiscent with smooth, diffluent membrane, in part more or less firmly fixed in the sporangiophore ; numerous spores in each sporangiophore (15-20 or more) elliptico-apiculate, with the extremities more MUCORALES 119 or less pointed, 3.0-3.5 x 4-4.5/a, producing one or two germ tubes, simple then repeatedly dichotomous ; zygospores not seen. Ferments glucose feebly ; as- similates well the monohexoses, peptone and ammonium sulphate. Growth good on bread and liquid media, such as Raulin's fluid, Difco malt extract, and Lendner's dealcoholized white wine agar. Growth slow on many other media reported. Mortierella sp. Costantin, Bull. Soc. Myc. France 8: 57-59, 1892. Isolated from a cat. Too poorly described for identification. BIBLIOGRAPHY Barthelat, G. J. 1903. Les mucorinees pathogenes et les mucormycoses, These de Paris. 127 pp., 13 figs.; Arch, de Parasitol. 7: 1-116, 13 figs. Bernard, P. Noel. 1914. Sur un Ehizopus pathogene de I'homme: RMzopus equinus Lucet & Costantin var. annamensis. Bull. Soc. Path. Exot. 7: 430-437. — . 1914. Sur un Ehizopus pathogene de I'homme, Bull. Soc. Myc. France 30: 230-232, PI. 14. Bodin, Eugene & Pierre Savoure. 1904. Eecherches experimentales sur les mycoses internes. Arch, de Parasitol. 8: 110-136. Casagrandi, Carmelita. 1931. Sulla presenza di Basidioboli nell'uomo, Biv. Biol. 13: 1-8, 1 fig. Also Boll. Soc. Internas. Microbiol. Ses. Ital. 9: 63, 64. Cavara, Vittoriano. 1913. Una forma nuova di cheratomicosi (cheratomicosi mucorina), Ann. Ottalmol. 42: 650-674, 1 pi. — . 1913. Sull'importanza patogena per I'occhio di alcune specie di Mucor, Ann. Ottalmol. 42: 729-771. Christiansen, M. 1922, Mycoses generalisees chez le pore, determinees par des mucorinees, C. B. Soc. Biol. 86: 461-463. — . 1922. Generel mucormykose hos svin, K. Veterinaer og landtoh^jsTcole. Aarsshrift. 1922: 132-190, 2 pis., 11 figs. Ciaglinski, A. & O. Hewelke. 1893. tJber die sogenannte Schwarze-Zunge, Zeitschr. Klin. Med. 22: 626-632, 6 figs. Ciferri, Eaffaelle & Bailey K, Ashford. 1929. A new species of Mortierella isolated from the human skin, Porto Bico Jour. Puilic Health. & Trop. Med. 5: 134-143, 1 pi. Cohnheim. 1865. Zwei Falle von Mycosis der Lungen, Arch. Path. Anat. Physiol. Klin. Med. 33: 157. Costantin, Julien. 1892. Note sur un cas de pneumomycose observe sur un chat par M. Neumann, Bull. Soc. Myc. France 8: 57-59. Costantin, Julien & Adrien Lucet. 1903. Sur un Ehizopus pathogene. Bull. Soc. Myc. France 19: 200-216, Pis. 9, 10. Dessy, G. 1933. La chemiotherapie des mycoses. III. Mucoromycose. IL Experiences in vivo. Boll. Ses. Ital. Soc. Internas. Microhiol. 9: 201-206. Ernst, H. C. 1918. A case of Mucor infection. Jour. Med. Bes. 34: 143-146, PI. 3. Frank. 1890. [Aspergillus fumigatus, Mucor racemosus], Deutsche Zeitschr. Thiermed. Vergl. Path. 16: 296, 297. Gasperini, Gustavo. 1927. II dinamismo citoplasmatico nelle Mucorinee sottoposte a varie azioni e singolaramente a quelle degli elettroliti, Ann. Ig. 37: 193-231, 3 pis. Gortner, E. A. & A. F. Blakeslee. 1914. Observations on the toxin of Ehizopus nigricans. Am. Jour. Physiol. 35: 353-367. Graham, Harry. 1890. Mucor corymbifer in the external auditory meatus. Lancet 2: 1379. Herla, Victor. 1895. Note sur un cas de pneumomycose chez I'homme, Bidl. Acad. Boy. Med. Belgique. IV, 9: 1021-1031, 1 pi. ]20 MEDICAL MYCOLOGY Hiickel, Armand. 1884. Zur Keniitnis der Biologie des Mucor corymbifer, Beitr. Path. Anat. Allg. Path. 1: 115-131, PI. S. Korte, W. E. de. 1923. A paramycetoma (?) of the forearm, Jour. Path. Pact. 26: 189-192. Lang, F. J. «& F. Grubauer. 1923. tJber Mucor- und Aspergillusmykose der Lunge, Arch. Path. Anat. Phys. [Virchow] 245: 480-512, 17 figs. Leinati, Fausto. 1928. Micosi rare in animali. Osservazioni cliniche sperimentali, Atti R. Accad. Fisiocrit. Siena X, 3: 83-92, 3 pis. — . 1929. Ulcere micoticlie sperimentali dello stomaco, Atti R. Accad. Fisiocrit. Siena X, 4: 147-223, ^3 figs. Lichtheim, L. 1884. tJber pathogene Mucorineen und die durch sie erzeugten Mykose des Kaninchens, Zeitschr. Klin. Med. 7: 140-177. Lindt, Wilhelm. 1886. Mittheilungen iiber einige neue pathogene Schimmelpilze, Arch. Exp. Path. Pharm. 21: 269-298. Lueet, Adrien & Julien Costantin. 1899. Sur une nouvelle Mucorinee pathogene, C. B. Acad. Sci. 129: 1031-1034. — . 1900. Ehizomucor j)arasiticus, espece pathogene de I'homme, Rev. Gen. Bot. 12: 81- 98, PL 3. — . 1901. Contribution a 1 'etude des mucorinees pathogenes. Arch, de Parasitol. 4: 362- 408. Macfarlan, Douglas. 1924. Fungus of the tongue. Laryngoscope 34: 720-722. Martins, Cesar. 1928. Mycose pulmonaire a Ehizomucor parasiticus, C. R. Soc. Biol. 99: 957, 958. Motta, Eoberto. 1926. Contributo alio studio delle micosi del condotte uditivo esterno (Studio clinico e micologico), Atti R. Accad. Fisiocrit. Siena. IX, 17: 603-631, 9 figs. Paltauf, Arnold. 1885. Mycosis mucorina. Ein Beitrag zur Kenntnis der menschlichen Fadenpilzerkrankungen, Arch. Path. Anat. Physiol. [Virchow] 102: 543-565, PI. 6. Parisot, Jacques & Pierre Simonin. 1922. Mycose pulmonaire associee; reactions biologiques et recherches exp^rimentales, Rev. Med. de l-Est. 50: 8-11. Pena Chavarria, Antonio & Janet H. Clark. 1924. The reaction of pathogenic fungi to ultraviolet light and the role played by pigment in this reaction, Am. Jour. Hyg. 4: 639-649, 6 figs. Podack, Max. 1899. Znr Kenntnis des sogenannten Endothelkrebs der Pleura und der Mucormykosen, Deutsch. Arch. Klin. Med. 63: 1-78. Eedaelli, Piero. 1924. Contributo sperimentale alio studio della Moniliasi e della Lichtheimiasi, Bif. Med. 40: 665, 666. Savour^, Pierre. 1905. Eecherches experimentales sur les mycoses internes et leurs parasites. Arch, de Parasitol. 10: 5-70. Sendziak, Johann. 1894. Beitrag zur Aetiologie der sogenannten schwarzcn Zunge, Monatschr. Ohrenheillv. KehlJcopf-, Nasen- u. RachenkranTch. 28: 112-119, 228. Smith, Theobald. 1920. Mycosis of the bovine fetal mem.branes due to a mould of the genus Mucor, Jour. Exp. Med. 31: 115-122, 1 fig. Spillmann, L. So Ph. Lasseur. 1922. Un cas d 'epidermomycose, Congr. Derm. Syphiligr. Langue Frang. 1: 42, 43. *Stange, G. 1892. Experimenteller Beitrag zur Pathogenitat der Mucorineen, Inaug. Diss. Dorpat. Vuillemin, Paul. 1903. La serie des absidiees, C. R. Acad. Sci. 136: 514-516. — . 1904. La Lichtheimia ramosa (Mucor ramosa Lindt) champignon pathogene distinct du L. cormybifera, Arch, de Parasitol. 8: 562-672, Figs. 1-4. Ziegenhorn, Otto. 1886. Versuche iiber Abschwachung pathogenen Schimmelpilze, Arch, Exp. Path. Pharm. 21: 249-268. CHAPTEE VIII ASCOMYCETES The Ascomycetes are those fungi in which meiosis occurs in characteristic sporangia with endogenous spore formation. These sporangia are called asci and their spores, ascospores. Their thallus is generally well developed; its hyphae (in contrast to those of the Phycomycetes) are regularly divided by septa into uni-, bi- or, rarely, multinucleate cells. Under certain environ- mental conditions, they may continue growth by sprouting ; in the yeasts and a few other forms, only the sprout mycelium is known. The imperfect forms reach the culmination of development in this group, especially among the pathogens of the higher plants. Besides oidia, hypno- spores, etc., the most varied types of conidia are found, often produced in highly specialized organs which at times approach the perfect (sexual) forms in complexity. In certain families, several imperfect forms may be produced successively or even simultaneously in the same species, a condition usually referred to as polymorphism. In case the perfect stage is unknown, these im- perfect stages are given a name and classified among the Fungi Imperfecti. The sexual organs of the primitive groups with which we are concerned more or less resemble those of the Phycomycetes, especially those of the Muco- rales. In the most primitive family we have gametes differentiated and set free to copulate in pairs. These produce a diploid ascogenous hypha. These conditions approximate those in the Oomycetes, although the ascogenous hypha and ascus seem to be a new development. Also in the Ascoideaceae we have a proliferation of the gametangium or ascus which is suggestive of the Oomycetes. Aside from these very primitive forms there are simple isogamous or heterogamous copulation branches very much as we found in the Mucorales. In the higher groups, there is an extensive functional and morphologic differentiation, the male being differentiated as an antheridium and the female as an ascogonimn. A unicellular antheridium approaches a unicellular ascogo- nium and is surrounded by the filamentous end of the ascogonium, known as the trichogyne. In the Plectascales, the only group of interest to medical men, there is not a great differentiation of trichogyne from the ascogonium. In most groups plasmogamy has lost its obligatory character and becomes facultative. Morphologically this functional disturbance first affects only the antheridia ; these disappear and amphimictic fertilization is replaced by many deuterogamous processes. (For details, see Gaumann & Dodge 1928.) Grad- ually this functional degeneration extends to the female organs which also disappear in many groups. Eventually no sexual organ is formed and plas- mogamy becomes pseudogamous. In conjunction with this degeneration, there is a shifting in the signifi- cance of the sexual organs for the formation of fructifications. In the lower 121 122 MEDICAL MYCOLOGY groups the fructification is initiated by the formation of the sexual organs. In many of the higher forms, external stimuli cause the fructifications to de- velop independent of sexual organs which are later formed within the fruc- tification. Plasmogamy is not followed directly by caryogamy but one or several dicaryons are formed. In the lower forms, the dicaryon migrates directly into an ascus which is the product of the plasmogamy. In the higher forms, plasmogamy is increasingly retarded and the fertilized gametangium develops into one or more hyphae. These take up the dicaryon and by conjugate divi- Fig-. 11. — Pyronema confluens. 1, Two antheridia arising from a dichotomous hypha, a trichogyne is in contact with each. 2, An ascog-onium showing fusion in pairs of the sexual nuclei. S, An older stage, showing the beginning of ascogenous hyphae. i, Young ascogenous hypha. 5, An older hypha in which wall formation is in progress. 6, Older hvphae in which the binucleate cells are building out to form croziers. (i and 3 X660, 2 Xl.060, Jf-S Xl,230.) (After Gwynne Vaughan & Williamson 1931.) sion, branch and form asci. Such dicaryotic hyphae are therefore called ascogenous hyphae; biologically they offer the advantage that one gametan- gium can create a number of asci. In most of the higher Ascomycetes, the asci develop from croziers at the end of the ascogenous hyphae. Each of the ascogenous hyphae arising from the ascogonium contains a number of dicaryons and develops by repeated forking, more or less vertically toward the top of the future fructification ASCOMYCETES 123 (Pig. 11, n) . Subsequently it divides by septa so that in the vicinity of the ascogonium the cells contain 2-8 dicaryons and farther away only one (Fig. 11, 4). A cell with only one dicaryon puts forth a lateral process whereby the nuclei are rather far separated (Fig. 11, 6) ; shortly the process bends around into a crozier, and the nuclei begin to divide conjugately (Fig. 12, 1). The spindles lie approximately parallel to each other. After the division, the crozier is ab jointed from both the tip and stipe cells which contain one nucleus each (Fig. 12, 2). In the simplest case the nuclei of the crozier fuse to a diploid nucleus, the primary ascus nucleus, and the crozier develops an ascus (Fig. 12. 3). In another type, the crozier develops a new crozier which in turn may develop still another. In any case it is only the terminal crozier that develops mi m wmm Fig. 12. — Pyronema confluens. 1, Older ascogenous hypha with a new ascogenous hypha budding out to form croziers, the tip cell uninucleate. 3, A crozier, showing fusion in the ascus cell. S, Prophase in the two nuclei of a crozier, each showing twelve chromosomes. 4, First mitotic telophase in the ascus with twelve whole chromosomes going to each pole. 5, Metaphase of the second division in the ascus, showing six chromosomes. 6, Third division in the ascus ; the lower nuclei are in the late metaphase, the next shows the anaphase, and that nearest the apex an ea.rly telophase in which six chromosomes can be counted at the pole. 7, Mycospliaerelln Fragariae, a tvpical peritheciuni. 8, Ascospores. (/ and 2 Xl,230 : S-6 XI, 760; 7 X360 ; S X800.) (After Gwynne Vaughan & Williamson, 1931 and Klebahn 1918.) an ascus. In a third type, the dicaryon of the crozier divides without form- ing any new crozier. The original crozier develops a branch which later may form a new crozier whereon an scus may arise directly; or caryogamy may again be retarded with the result that a tuft of croziers is formed. Occa- sionally the stipe and tip of the crozier fuse, the stipe nucleus generally migrating into the tip cell. This proceeds to develop a binucleate branch which gradually forms a crozier that may develop an ascus by fusion of its nuclei or repeat crozier formation. 124 MEDICAL MYCOLOGY In the higher Ascomycetes, there are, in addition to the crozier type, a whole series of other developmental forms of ascogenous hyphae which need not concern us at present. As far as is known, the further development of the asci is the same in all Ascomycetes. The primary ascus nucleus, which has arisen from the fusion of the dicaryon (Fig. 12, 3), undergoes three divisions, at least one^ of which is meiosis ; the eight daughter cells cut out eight ascospores from the cytoplasm of the ascus by free cell formation. The cytoplasm not included in the spores is called epiplasm, which, besides nourishing the ascospores, pro- vides substances for the sculpturing of the spore walls. In certain forms, the number of nuclear divisions may be limited to two or may increase to sixteen, in the former case producing but 4 ascospores and in the latter 64,936. Where the ascospores are thick-walled, they usually possess a typical germ pore or a meridional fissure. In the latter case, the halves of the ascospore wall sepa- rate in germination like the two valves of a mussel. According to the Anglo-Saxon school, represented by Harper, B. 0. Dodge and Gwynne-Vaughan (nee Fraser) the nuclear fusion in the young ascus is not the first and only fusion ; but is preceded by another fusion in the ascogonium directly after plasmogamy. The ascogenous hyphae, accord- ing to this conception, do not contain haploid dicaryons but undivided diploid nuclei which only after the formation of the croziers come together as di- caryons. Because of this double fertilization, the primary ascus nucleus is tetraploid and contains 2x double chromosomes. At the first ascus division (Fig. 12, 4) meiosis occurs with each daughter nucleus containing 2x simple chromosomes. The second step is homeotypic (Fig. 12, 5), the 2x simple chromosomes are halved so that each daughter nucleus still contains 2x simple chromosomes. In the third step (brachymeiosis) (Fig. 12, 6) one-half of the undivided chromosomes migrates to each pole, so that each daughter nucleus of the third division contains x simple chromosomes. Although the cytologic reports are somewhat contradictory and in part may be interpreted by either hypothesis, the students of Continental Europe and some in America prefer the interpretation of Dangeard and Claussen. First imperfect forms, then sexual organs arise on the haplont. Between these sexual organs plasmogamy occurs, while the male and female nuclei pair as a dicaryon. These dicaryons migrate into the ascogenous hyphae and divide conjugately. The ascogenous hypha thus represents a special diploid phase, the dicaryophase, which ends with caryogamy (fusion) in the young asci. Caryogamy is followed directly by meiosis, usually producing 8 haploid ascospores. In the higher Ascomycetes this scheme of development is further complicated, since the haploid thallus proceeds to form fructifications on or in which the ascogenous hyphae complete their development. As in most red algae and in the sporophyte of the mosses, the dicaryophase is to a certain extent parasitic on the haplont and nourished by it. In the simplest case, these fructifications form an undifferentiated mass of tissue, a stroma on or in which the asci are formed. A fructification of this ASCOMYCETES 125 type is called an ascostroma. In the liigher forms the hyphal tissue of the stroma undergoes differentiation both in form and histologic structure, and develops the fructifications which furnish important characters for classifica- tion. Only one of these structures in its simplest form need concern us here. For a further consideration of the higher Ascomycetes see Gaumann & Dodge (1928) and the recent fundamental work of Nannfeldt (1932). The perithecium consists of a solid, often pseudoparenchymatous wall and a cavity in which the asci are borne (Fig. 12, 7). The more primitive types are usually spherical ; the asci lie irregularly in the interior and are only liberated by the decay of the perithecial wall. In the higher types, there are more elaborate mechanisms for spore dispersal. BIBLIOGRAPHY Gaumann, Ernst A. & Carroll William Dodge. 1928. Comparative morphology of fungi. New York, McGraw Hill Book Company, 701 pp., 398 jigs. Nannfeldt, J. A. 1932. Studien iiber die Morphologic und Systematik der nicht-lichenisierten inoperculaten Discomyceten, Nova Acta E. Soc. Sci. Upsaliensis IV, 8: 2:1-368, Pis. 1-20. CHAPTER IX ENDOMYCETALES The Endomycetales include those forms in which an ascus arises directly as the product of a sexual act, wherever this occurs. They comprise eleven families distributed among four diverging lines of degeneration. In the most primitive family, the Spermophthoraceae, the mycelium is nonseptate and coenocytic, the gametes are differentiated and freed from the gametangium, and a septate uninucleate secondary mycelium results. From this primitive family we have four diverging lines of degeneration, each line having a char- acteristic spore shape. In the Ashbyaceae, the mycelium, when present, may be septate, but the cells are usually coenocytic and have the elongate fusiform ascospore of the Spermophthoraceae (Fig. 13, 1-3). In the Ascoideaceae — Endomycetaceae line, the mycelium becomes uninucleate, the ascospores, which in the early stages of its development are fusiform, become cucullate (Fig. 13, 10), or the rim assumes an equatorial position, producing a saturnine spore (Fig. 13, 6). The position of the Pichiaceae is not clear. Here the ascospores are hemispheric or slightly angular (Fig. 13, 7, 8). They may pos- sibly be derived from GuiUermondeUa (Fig. 13, 5), a member of the Ashbyaceae, or more probably Hanseniospora, one of the Endomycetaceae with rough cucul- late spores, by the loss of the rim (Fig. 13). In the two remaining lines, the spores are ellipsoid or spherical and have probably diverged from the Spermophthoraceae through Dipodascus. One line retained strong evidence of sexuality, gradually losing it with extreme degenera- tion, producing its spores saprophj'tically, and early reducing the ascospore number to 8, 4, or fewer. This line is represented by the Eremascaceae and Saccharomycetaceae. The other line promptly discarded traces of sexuality, produced its spores in the host tissue, very rarely under saprophytic condi- tions, and retained the large number of ascospores of the Dipodascaceae. This latter line is represented by three strictly parasitic families, one comprising predominantly mammalian parasites, the other two plant pathogens. There is a small residue of species usually placed in the Saccharomycetaceae, in which the ascospores copulate in pairs. Their systematic jDosition is not clear. Guil- liermond has recently (1931) suggested that this group of species has been derived from the Taphrinaceae, the end member of the fourth line mentioned above. Key to Families Gametes fusiform, set free from the gametangium, copulating in pairs, producing an ascogenous hypha; ascospores fusiform. Spermophthoraceae. Gametes not set free, gametangial copulation the usual type, or the ascospores develop parthenogenetically. Ascospores fusiform to acicular. Ashbyaceae. 126 ENDOMYCETALES 127 Ascospores, cucullate, saturnine. Mycelium multinucleate, conidia produced, ascus many-spored, proliferating. Ascoideaceae. Mycelium uninucleate, degenerating to sprout mycelium; conidia not differentiated; ascospore number usually 4 or fewer; not proliferating. Endo'mycetaceae. Ascospores hemispheric or angular; sprout mycelium uninucleate; ascospores 4 or fewer. Pichiaceae. Ascospores ellipsoid or spherical. Mycelium multinucleate, ascus resulting from copulation of two hyphal tips. Ascus many-spored. Dipodascaceae. Ascus 8- or 4-spored. Eremascaceae. Mycelium uninucleate, usually sprout mycelium, asci resulting from copulation of two cells or from parthenogenesis or apogamy. Saccharomycetaceae. No trace of copulation; asci many-spored, rarely reduced to 8; mycelium often scanty in the tissue but then developing readily in culture; asci usually thick-walled, often differentiated as a resting spore, usually abundant in host tissue, rare in culture. Ascospores developing directly and filling the ascus. Coccidioideaceae. Ascospores developing in tetrads about the wall of the ascus. Protomycetaceae. Ascospores developing directly, but reduced in numbei', not filling the ascus, mycelium developing in host tissue. Taphrinaceae. Fig. 13. — Spore shapes in the Endomycetales. J, Nematospora; S, Coccidiascus; S, Mono- sporellaj ■'/, Schwanniomyces ; 5, Nadsonia; 6, Williopsis ; 7, S, Pichia; 9, Guilliermondella ; 10, Endomyces ; 11, Saccharomyces. (After Guilliermond 1928.) Spermophthoraceae. — In 8'permophthora Gossypii on cotton, the mycelium is nonseptate and coenocytic. It produces gametangia with numerous fusiform gametes (Fig*. 14, 1). After the dehiscence of the g-ametangium, the gametes fuse in pairs, and the resulting zygote germinates immediately by a septate, uninucleate, diploid mycelium (Fig. 14, 2-7). The asci are borne directly on this mycelium without crozier formation (Fig. 14, 8). The ascospores are fusiform, but smaller and shorter than the gametes. The differentiation of gametes is suggestive of remote derivation from tlie Oomycetes rather than from the Zygomycetes or Mucorales, as suggested by Gaumann (1926). In neither group is there any structure in any way comparable to the diploid ascogenous mycelium of this family. 128 MEDICAL MYCOLOGY Ashbyaceae. — This family has been little studied cytologically. In Pied- raia Hortai, forming hard nodules on human hair, the mycelium is thick-walled, septate, and more or less agglutinated into a solid mass surrounding the repro- ductive structures (Fig. 15, 1, 2, 12). Young cells contain up to 8 nuclei, but mature cells are mostly uninucleate, although one cell figured by Horta (1911) suggests a binucleate condition. The functions of gametangium and ascus, performed by distinct structures in the Spermophthoraceae, are performed by a single stinicture, usually called the ascus, which arises as the terminal cell of a hypha. The gametes are differentiated within this structure but unite in pairs without being set free (Fig. 15, 3-11). The resulting zygotes then Fig. 14. — Spermophthora Gossypii. 1, gametangium showing immature gametes within ; Z-%, stages of copulation of gametes, ascogenous filament with young ascus ; 8, asci showing as- cospores ; 9, germinating ascospores. (After Guilliermond 1928.) elongate to produce the 8 uninucleate, fusiform, or crescent-shaped ascospores with 2 (rarely 3) filiform appendages (Fig. 15, 13-16). The details of the cytology have not yet been reported. The ascospores germinate directly to mycelium which penetrates beneath the cuticle of the hair, forming a pseudo- parenchymatous palisade which eventually ruptures the cuticle and expands to produce the typical nodule. In Pieclraia veneznelensis, little is known of its life history, but the ascospore number is reduced to four and the filiform ap- pendages practically disappear. Langeron (1929) and Brumpt & Langeron (1934) suggest that Piedraia is related to the sooty molds which it resembles slightly in general appearance, but in the curious development of gametes and ENDOMYCETALES 129 Fig. 15. — Fiedraia Hortai. 1, S, sections of masses on hair, showing developing asci ; S-11, development of ascospores in ascus ; 12, mycelial mass on hair ; IS, 15, 16, ascospores ; 1^, ascospores emerging from ascus. 130 MEDICAL MYCOLOGY in the lack of ascogenous hyphae, it has nothing in common with that group, and is intermediate between the Spermophthoraceae and the Ashbyaceae. In Eremofheciuni (Fig. 16), the mycelium is multinucleate and rarely sep- tate. The genus has not been carefully investigated cytologically, but the gametangium (or ascus?) resembles that of Spermophthora in shape. The spore number is somewhat less. The spores are fusiform, rounded at one end, taper- ing to a long filiform appendage at the other. They are arranged in the ascus with the rounded ends in contact and the filiform appendages gathered in a fasicle at the poles, giving the whole spore mass the appearance of a huge nuclear spindle. This grouping suggests that figured by Horta (1911) for Piedraia. The protoplasm of the spore is much denser in the end opposite the appendage, and germination takes place only in the end of the spore with the dense protoplasm. No septum has been detected separating the spore into two cells. Yig 16. — Ashbya Gossypii. 1, mature spore; Z, S, germinating spores; i, mycelium; 5, 6, de- velopment of gametangium; 7, mature ascus with ascospores. (After Guilliermond 1928.) In Ashhya Gossypii, a parasite on cotton, the mycelium is septate but the cells are multinucleate. Sexuality has been lost, the asci developing partheno- genetically. The nuclei divide twice, forming the tetrads which precede spore formation. This is reminiscent of sporangiospore formation in Piloholus and will be encountered several times in other lines of this group. The number, both of nuclei and of nuclear divisions, is reduced and stabilized so that ordinarily either 8 or 16 spores are produced. The spores are usually rounded at one end and taper at the other into a long slender projection, sug- gestive of a flagellum, but without motility. In Nematospora, which is also parasitic on plants, degeneration has pro- ceeded further until sprout cells as well as mycelium are produced ; the spores are long fusifonn to acicular and reduced to 8 per ascus (very rarely 16, or ENDOMYCETALES 131 further reduced to 4 or 2 in N. Nmjpuyi) . The same fiagellar appendage is also present but usually shorter (Fig-. 17). Two other little known genera may either beh)ng here or in the next family. In (Joccidiascus mycelium is absent, the asci develop following isogamous copu- lation, and the spore number is fixed at 8 (Fig. 13, 2). This species has been found in the digestive tract of Brosophila but has not been cultivated. In Mono- sporella both mycelium and copulation are absent. The ascus produces a single acicular spore parthenogenetically (Fig. 13, 3). The members of this genus have also been found in the digestive tract of invertebrates and not cultivated. Fig. IT. — Nematospora Coryli. 1, mature ascospore ; 2, 3, geiminating: ascospores ; i, 5, sprout celLs ; b", developing ascus; 7, mature ascus. (After Guilliermond 1928.) PIEDRAIA Piedraia Fonseca & Area Leao, Inst. Oswaldo Cruz, Suppl. das Mem. 4: 124-127, 2 pis., 1928. Trichosporon Behrend, Berliner Klin. AVoch. 27: 464-467, 1890. TriclwHporum Vuillemin, C. R. Acad. Sci. 132: 1369-1371, 1901. Arch, de Parasitol. 5: 38-66, 12 figs., 1902, non Fries 1825, 1849. Fries, Syst. Orb. Veg. 306, 1825, published a new genus Trichosporum which is spelled Tn'chosponum in the index. No species was attributed to this genus at the time. I can find no mention of the genus in his Sy.stema mycologicum 132 MEDICAL MYCOLOGY 1832. Its first place of effective publication may be considered to be his Summa Veg. Scand. 2 : 492, 1849, where he treats twelve species, many of them f oi-ming the first section of liis Sporotrichum in the Systema. For spelling we are equally puzzled, as it is Trichosporum in the text and Trichosporium in the index. Saccardo and later authors have used the latter spelling. In any case Fries' use of the former precludes the possibility of using Trichosporum for another genus, and TricJiosporum of Vuillemin, Schachter, and later medical men, must be renamed. One might consider the retention of Trichosporon Behrend, since it differs by the last two letters, but the frequent interchange of spellings of genera, such as Microsporum and Microsporon, as well as the fact that the spelling Trichosporum has had the wider usage in the present century, makes it a permanent source of error and confusion, so that I am in favor of abandoning it altogether. Ponseca and Area Leao (1928) have reported asci and ascospores for Trichosporum Hortai and have transferred this species to Piedraia. No one has suggested asci in the European species while practically all investigators have noted these structures in the South American species whether they have called them asci or not. Also the lesions in the hair in the case of the European species are much more serious, causing irregular splitting of the hair, suggesting that the European species may belong in some other group of fungi, perhaps remotely related to the Gymnoascaceae or the Eremascaceae. Until more is learned about these imperfectly described species, we prefer to leave them as an appendix of doubtful species of Piedraia rather than to transfer them elsewhere. Piedraia is very imperfectly characterized. Mycelium thick-walled, septate, agglutinating into solid masses on the hair; asci 8-spored; ascospores large, fusiform with acute ends prolonged into filiform appendages. Piedraia Hortai (Brumpt) Fonseca & Area Leao, Inst. Oswaldo Cruz. Suppl. das Mem. 4: 124-127, 2 pis., 1928. Tnchosporum sp. Horta, Mem. Inst. Oswaldo Cniz. 3: 87-107, Pis. 5, 6, 1911. Trichosporum Hortai Brumpt. Precis Parasitol. 1913. Forming characteristic hard, black, adherent, small, spherical or long conic nodules on hair, Brazil. Hyphae septate, 8-12^ in diameter slightly brownish, thick-walled; asci not clearly seen in culture; ascospores fusiform (Fig. 22), curved, greenish yellow, each end acute and prolonged into an appendage about 30;u, long, the body of the spore being about 30 x 10/a. On Sabouraud agar, colonies small, dark brown, very adherent to the medium, velvety, margin somewhat lighter, finally becoming folded. After 10 weeks the whole colony is black. Growth much better on carrots where the lighter colored margin is lacking. Piedraia Sarmentoi Pereira f., Kev. Med. Cimrg. Brasil 38: 49-52, 6 pis., 1929; C. R. Soc. Biol. 104: 680, 1930. ENDOMYCETALES 133 Abundant on the hair of young people in the state of Rio Grande do Sul, Brazil. General microscopic appearance on hair similar to that of P. Hortai. Hyphae up to Tfi in diameter. Asci more spherical, up to 30/x long, 8-spored, ascospores 35-40 x 7-8fi, filiform appendages 7-8 rarely lO/x long. In cultures only terminal and intercalary chlamydospores seen. Growth rapid on Sabouraud agar, colonies white, low margins dentate, creamy then entirely black, fuliginous, easily detachable although penetrating the substrate deeply. Growth slow on potato, and pigment formed very late. On carrot, colony creamy white, pigment first appearing in spots, becoming diffused, fuliginous, cerebriform. Piedraia surinamensis Dodge, n. sp. TricJiosporum sp. Aars, Arch. Derm. Syphilol. 22: 401-409, 1930. Nodosities on hair commonly up to 500/*, rarely 1 mm. long, mostly about 100/A in diameter exclusive of the diameter of the hair, composed of thick- walled cells 4:-6fx in diameter. Asci occurring singly, 32-44 x 20/a; ascospores fusiform, 42 x 6^, with two, seldom three, filiform appendages at the ends. On maltose agar and honey agar, dark brown to black, hard, slightly velvety colonies. Piedraia colombiana Dodge, n. nom. Dematium sp. Juhel-Renoy & Lion, Ann. Derm. Syphiligr. Ill, 1 : 765-772, 2 pis., 1890. TricJiosporum giganteum Vuillemin, Arch, de Parasitol. 5: 38-66, 12 figs., 1902 non Unna 1895. Producing nodules on the hair in Colombia. Cells 4-5 X 5-6/x sometimes 8-12/xi long, yeastlike and proliferating when young, later forming a mycelium, in old age again becoming more or less yeastlike. Coils were observed in cultures, but it is not certain whether they were functional antheridia and ascogonia. [Malcolm Morris (1879) had pre- viously noted asci with spores, but he does not describe them in sufficient detail. Peiia Chavarria & Rotter (1983) describe the ascocarps, asci, and spores in some detail but fail to give measurements of the ascospores.] Brumpt & Langeron (1934) studying a case from Medellin, Colombia, state that the asci were 50 x 30/a, containing 8 ascospores ; spores thick-walled, 40 x 6-9/* with very short filiform appendages, 4-5/t long. On Sabouraud agar, colonies white at first, becoming yellowish, cerebri- form, not penetrating the substrate and easily separable. In old cultures the cerebriform appearance is lost. On maltose agar, colonies darker and more adherent. On sugar beets and carrots, growth good. Gelatin liquefaction begins within 8 days. Piedraia venezuelensis Brumpt & Langeron, Ann. Parasitol. Hum. Comp. 12: 155-158, Figs. 27-32, 1934. Infected hairs sent by Machado of Caracas from a case of 2 years' dura- tion, apparently the first case reported from this region. 134 MEDICAL MYCOLOGY Nodules on the hair, large ovoid or fusiform ; asci 35-40/* long, only 4- spored; ascospores 25-30 x 10-14/* very thick-walled, lacking filiform append- ages but ending in long slender points up to 10-12/* long, often somewhat curved. Not cultivated. Species of Doubtful Position The following species are rather imperfectly described and may belong elsewhere, although they were placed here by the original authors. In some cases I have been unable to locate the original description and know the name only by a brief mention in the literature. Trichospomm Beig-elii (Rabenhorst) Vuillemin, Arch, de Parasitol. 5: 38-66, 12 figs., 1902. Pleurococcus Beigeli Klichenmeister in Rabenhorst, Hedwigia 6: 49, 1867. Sclerotium Beigelianum Hallier, Paras. Unters. 75, PI. 2, Figs. 24, 25, 1868. Zoogloea Beigeliana Eberth, Centralbl. Med. Wiss. 11: 307, 1873. Hyalococcus Beigeli Schroeter, Kryptog. Fl. Schlesien 3: 152, 1886. Chlamydotomus Beigeli Trevisan in Saccardo, Syll. Fung. 8: 1042, 1889. Micrococcus Beigeli Migula, Syst. Bakterien, 1: 193, 1900. Attacking the hair in wigs and switches, Germany. Vuillemin (1902) isolated what he called this fungus from the hairs of the moustache. The report of this fungus from pubic hair in Nigeria by Manson-Bahr (1932) probably should be referred to Favotrichophyton. Cells 3-5/t in diameter, spherical or angular by mutual pressure, surround- ing the hair in a spherical mass of gel ; sporangia mostly 1/* thick, containing 12-20 spores. Vuillemin describes the fungus isolated by him as follows: Cells 2.5-4/*, mostly 3-4/*, wall thick, held together by gelification of the outer layer of the wall. Larger chlamydospores in the interior of the stroma, 6/* in diameter. On maltose agar, gelatin, carrot, beet, and potato, a yellowish gray, humid colony, becoming convoluted, surface drying chalky white. Cells cylindric, 4-5/* in diameter, forming a more or less dichotomous mycelium about 2/i in diameter, producing hypnospores filled with reserves up to 12/* in diameter. Gelatin and serum not liquefied. Producing a pellicle on the surface of liquid media, growth suggesting- that of Geotrichum lactis. Trichospomm ovale Paoli, Giorn. Ital. Mai. Ven. Pelle 54: 566-572, 1913, non Unna ap. Vviillemin 1902. Isolated from infected hairs of the moustache with the usual trichosporic nodules, hair not breaking readily. Spores in nodules 3-4/i, hyphae present. In cultures, mycelium of branched, septate hyphae bearing terminal chains of spores often suggesting clostero- spores. Not pathogenic for guinea pig or rabbit. On Sabouraud agar, colony whitish, moist, radially furrowed, slightly velvety, becoming yellowish. On gelatin, colony similar but growth much ENDOMYCETALES 135 slower, gelatin liquefied in 13-15 days. On broth, grayish white pellicle settling after 6-7 days and replaced by another pellicle. From the brief description, the systematic position of this organism is uncertain. Many characters suggest much closer relation to the Favotricho- phyfon ochraceuni group rather than to the yeasts (near Mycotornla) . Clini- cally it seems very close to the European T. Beigeli, if it be not the same organism. Trichosporum ovcides Behrend, Berliner Klin. Woch. 27: 464-467, ld90. Trichosporum ovale Unna ap. Vuillemin, Arch, de Parasitol. 5: 38-66, 12 figs., 1902. ITrichosporum giganteum. Unna, Deutsch. Med. Zeitschr. 1895: 255-256, 1895, non Vuillemin 1902. Isolated from sycosis in Germany, forming brownish yellow nodes on the hair, leaving the hair shaft nearly intact. Spores ovoid, 2-4.5 x 1-4/x, sometimes in-egular. Colonies cerebriform, blackish brown on several media. Trachsler (1896) tried to separate T. ovoides and T. giganteum, but the differences between the strains are so slight that they are probably not sig- nificant. Trichosporum glycophile DuBois, Ann. Derm. Syphiligr. V, 1: 447-456, 1910. Causing irritation in female genitalia. The hairs which have been moistened with urine (patient was a mild diabetic) have nodosities and terminate in brushlike tips, often breaking a few millimeters from the skin. The mycelium proliferates between the fibrillae of the hair and terminates at the surface under a layer of sporiform elements. Not pathogenic for the guinea pig, rabbit, or rat. The following description of morphology based on cultures from 5% maltose agar, 5% peptone agar, or 2.5% maltose + 2.5% peptone agar. The yeast cells 5-6/x in diameter. The hyphae slender, septate, at long intervals, uniform at least in the terminal portion. As the hyphae become older, they become thicker, more tortuous, and end in a terminal swelling. Spores verticillate, regularly spaced. Chlamydospores large. On solid sugar media, abundant mycelium below the surface while on the surface a small gray colony scarcely covers the hair. On solid protein media, colonies yellowish, smooth, humid, proliferating on surface with cen- tral elevation and sometimes Avith little knobs, margin with a circular elevation connected with the center by furrows. Cross inoculations showed these two types of colonies to be the same organism. On Sabouraud agar, colonies smooth gray, or yellowish, growth rapid. On broth agar and peptone agar, yeast form prevails on the surface, growth slow, colony gray, smooth with a slight central knob, colony bordered by fine hyphae at the end of 3 weeks. On liquid media, grayish white pellicle, dry but not poAvdery, pellicle settling in 10-12 days and a new one forms. On potato, colonies gray yellowish, moist, rugose. 136 MEDICAL MYCOLOGY Gelatin not liquefied, but fine hyphae perpendicular to the line of the stab seen and a tuft of mycelium at the bottom. The systematic position of this species is not at all clear. Its behavior on the hair suggests relationships with the European species previously placed in Trichosporuni. On the other hand, this organism caused considerable irrita- tion of the genitalia while the so-called Trichosporuni of Vuillemin and others caused no irritation. Its morphology in culture would relate it to Mycotorula as has already been pointed out by Langeron & Talice (1932), but none of the other species of the group have been reported able to penetrate the hair. Trichosporuni equinum Fambach teste Fonseca, Rev. Med. Cirurg. Brasil 38: 256, 1930. Producing white nodules on hair of horse, becoming yellowish or darker. Trichosporum Foxi Castellani, 1908. Pichiaceae. — The systematic position of this family is not clear. It may have arisen from Nematospora or Ashhya or, more probably, as another line of degeneration from the Ascoideaceae. The most primitive member is Guillier- mondella which produces considerable mycelium with conidia. Asci 4-spored, either lateral or intercalary, follow isogamous copulation or may develop par- thenogenetically. Spores sickle-shaped [Dekker (1931) figures them as kidney- shaped very close to those of Pichia]. A somewhat more degenerate species, GuilUermondella Vuillemini (Endomyces albicans Vuill. non Johan-Olsen, En- domycopsis albicans Dekker) from a case of thrush, still retains both mycelium and sprout mycelium. The spore number is 4, or fewer. In Zygopichia and Pichia there are cylindric cells and some mycelium for- mation, forming a thick, dry pellicle on the surface of liquid media. In Zygopichia heterogamous copulation precedes ascospore formation, in Pichia the degeneration is complete, and the ascospores develop parthenogenetically. Both genera have kidney-shaped, hemispheric, or angular ascospores. Guilliermondella Vuillemini (Lindau?) Dodge, n. comb. Endomyces albicans Vuillemin, C. R. Acad. Sci. 127: 630-633, 1898; Rev. Myc. 21: 43-45, Pis. 189-190, 1899 not Johan-Olsen 1897 (excl. all syn. based on Oidium albicans Robin). Endomycopsis albicans Dekker (excl. syn.), Verhandel. K. Akad. Wetensch. Amsterdam, Afd. Natuurk. 28: 231, 1931. Endomyces Vuillemini Lindau?, Mikroscopische Pilze 1912 [cited as Landrieu by Castellani & Chalmers, Man. Trop. Med. ed. 2, 1913, but I am unable to verify this citation]. Isolated from a case of thrush. Yeast cells abundant at first, giving rise to mycelium with chlamydospores and finally producing asci, spherical or ellipsoid, 4-5/*, 4-spored, rarely 2-3- spored, axes of ascospores 3.8-3.5 x 1.75-2 x 1.2-lA/x, not staining with nuclear stains (i.e., the deeply staining bodies called by Vuillemin the ''globules internes" are probably nuclei, fide unpublished work of Morris Moore). The ascospores cling together in groups for some time after the asci have dis- appeared. ENDOMYCETALES 137 Ascoideaceae. — In Ascoidea rubescens in the slime flux of trees, the myce- lium is coenocytic, septate, and branched. The mycelium produces conidia either singly or in tufts. The gametangium is a large multinucleate cell. Varitchak has reported the degeneration of all but one pair of nuclei, which proceed to fuse. Walker (1935) was unable to confirm this statement and suggests that the "privileged sexual nuclei" of Ascoidea are degenerating nuclei which she has also seen in other parts of the fungus. By two or three successive divisions the spore initials are cut out of the mass as lenticular, uninucleate portions of the protoplasm. Finally, these spore initials contract and form the typical cucullate spores of this series. Not all of the protoplasm is used up in the process, some remaining behind as the epiplasm, as in other groups of Ascomycetes. The ascospores are extruded from the mouth of the ascus by the proliferation of the cell next below it. This proliferating cell proceeds to form another ascus at the same site. Plasmogamy occurs shortly after spore germination, but the occurrence of caryogamy is still uncertain. The spore sac appears similar to the gametangium of Spermophthora, but gametes are not differentiated and whether there is nuclear fusion in this organ is still questioned. In its later stages the organ behaves as a multi- spored ascus, although ontogenetically it bears little relation to such a structure. The proliferation of the basal cell into the mature organ sug- gests sporangial proliferation in the Saprolegniaceae, but the manner of spore formation is entirely different. It would seem likely that we are dealing with a stage of degeneration from an ancestral form like Spermoph- thora in which the gametangium has ceased to function as such and func- tions as an ascus with a shortening of the stages between gametangium and ascus and with a partial or complete elimination of the sexual act. The fact that the spore initials appear similar to the ascospores of Spermophthora and finally become cucullate as in the Endomycetaceae, suggests that it may be an intermediate stage in the phylogeny of the latter. Endomycetaceae.- — In this family the mycelium is usually uninucleate, soon degenerating to sprout mycelium, conidia are no longer differentiated, the ascospore has become reduced in number per ascus and has assumed a cucul- late or saturnine shape. So far as is known this family is saprophytic, although Hansenula has occa- sionally been isolated from sputum, and Pijper (1928) reports Hanseniospora from a case of onychomycosis. The strains of fungi from these isolations have not been reported pathogenic for experimental animals. It is possible that they may cause irritation in the mucous membranes or aggravate a condition primarily due to some other organism, but reports of the cases should be scrutinized very carefully before admitting them as pathogens. On the other hand, there is a large group of parasites previously referred to Endomyces, having spherical to ellipsoid, smooth spores, which is here treated in the Eremascaceae. 188 MEDICAL MYCOLOGY This family is characterized by the rim around tlie spore, producing a cucullate spore if the rim is on one side, or a saturnine spore if the ring is equatorial. Within the family there is a large isogamous series and a small heterogamous one. In the heterogamous series, which seems to be the more primitive, we have Endoniyces Magnusii {Magnusiomyces) from the slime flux of trees (Fig. 18). The hyphae are generally multinucleate (2-8). In growing hyplial tips this number may mount to 50 (Pig. 18, 1), in weak hyphae it may be as low as one. The hyphae divide easily into oidia ; Avliich are generally multinucleate, rarely uninucleate. In successive divisions the tendency is toward the uninu- cleate condition. Often their wall thickens and the oidia become hypnospores Fig. 18. — Endomyces Magnusii. 1, young- multinucleate hypha ; 2, older hypha ; 3-9, II, development of asci ; 10, hvpnospores. {1, 2, 10 xl.500; S-9, 11 X500.) (After Guillier- mond 1909.) (Fig. 18, 10) ; with the consequence that, under certain environmental con- ditions, a culture may disintegrate into hypnospores after a couple of weeks. With favorable conditions, oidia may develop to sprout mycelia, not by independent development of small outgrowths of the mother cell to sprout cells but by fission of the mother cell. Both daughter cells round oft' and develop to the size of the mother cell (ordinary cell division in contrast to sprouting). When the mycelium is ready to form asci, it divides into numerous short, slender branches with short cells containing few nuclei, often not more than one (Fig. 18, 3, and 4). A branch ends either in a very large cell full of reserves, the ascogouium, or in a narrow hyaline cell, often much twisted, the antheridium. The upper third of the ascogonium swells considerably and ENDOMYCETALES 139 collects the cytoplasm with 2 or 3 nuclei. At the beginning' of copulation, it bends over to meet the antheridium. In this stage, the swollen part contains only one nucleus, the others having^ migrated (Fig-. 18, 5). The narrow an- theridium contains, when young, 1-3 nuclei of which only one remains at the tip. In about three-fourths of the cases, copulation occurs. The antheridium approaches the ascogonium, swells slightly, and ab joints the apical uninucleate gametangium from the stipe cell. Meanwhile the uninucleate tip of the ascogonium is abjointed from its basal cell. Hereupon the walls separating the gametangia dissolve, with the development of tlie zygote to a 4-spored ascus (Fig. 18, 6-9). There often occur numerous variants of this usual course of development. Thus, the antlieridium may approach the ascogonium at the side instead of the tip; or both may be uninucleate without the abjnnction of basal cells; or the ascogonium may develop parthenogenetically. Fig. 19. — Endomyces decipiens. 1-3, stages of copulation; .'(-12, development of asci. (X 1,200.) (After Juel 1921.) The end member of the heterogamous series is Endomyces decipiens, found on fructifications of the mushroom, Armillario. mellea, producing its perfect stage on the lamellae. Sexual organs are almost entirely absent, copulation being heterogamous when present or very rarely isogamous (Fig. 19, 1-3). The asci are lateral on the hyphae. Although sometimes three nuclear divi- sions occur in the ascus, only 4 ascospores are formed (Fig*. 19, 4-12). In cultures the hyphae easily break apart into uninucleate oidia. Sometimes the tips of branches form thick-walled yellowish hypnospores instead of asci. The isogamous series begins with the genus Endomycopsis. In this genus the abundant sprout mycelium is apparently connected with the adaptation of these species to media containing starch and sugars. Wherever the sprout cells arise on aerial mycelium, their diameter is smaller and the Avail some- what thicker than in tlie submersed sprout cells. They are then ver}^ resistant and survive a long period in temperatures up to 55° C. Biolog-ically their significance seems to be that of hypnospores, and because of their exogenous formation they are usually called conidia. 140 MEDICAL MYCOLOGY In Endomycopsis fihuliger, although any two cells may form copulation branches which approach each other, even with the dissolution of the wall the 2 nuclei rarely fuse {Fig. 20, 1). Generally the copulation branches de- velop parthenogenetically, though the separating walls may be temporarily dissolved. In exceptional cases there may occur a pseudogamous anastomosis of two sprout cells, one changing to an ascus. (Fig. 20^ 2-4). In a large number of cases no copulation branches are formed, but the asci, like the sprout cells, arise as lateral outgrowths of the hyphal cells (Fig. 20, 6). These asci are three or four times larger than the ordinary sprout cells. Occasionally, they arise from ordinary swollen hyphal cells, or from swollen sprout cells. When they begin to appear, the formation of sprout cells slows up, but does not cease, with the result that hyphae may form sprout cells and asci simultaneously. One finds even young asci which continue to cut off sprout cells until the beginning of spore formation. Periods Fig-. 20. — Endomycopsis fihuliger. Development of asci. (XoOO.) (After Guilliermond 1909.) of vegetative growth and fructification are thus not sharply differentiated one from the other. Each ascus contains four cucullate spores. At germina- tion, these throw off the exospore and germinate either with a germ tube or with a sprout mycelium. Here sexuality is so completely weakened that only vestiges of the sexual organs remain. In a large number of cases where no copulation branches are formed, the asci arise directly from vegetative hyphae or sprout cells. In the remaining forms of the isogamous series, copulation branches de- velop less frequently and the asci arise parthenogenetically without fusion. Growth of mycelium by sprouting increases proportionally. In two Chinese species, E. Lindneri and E. Hordei, the copulation branch no longer changes directly to an ascus but develops to a short, occasionally branched mycelium where asci arise by swelling of the hyphal cells (Fig. 21, 1, 2) . In most cases the asci are formed directly from the sprout cells without this detour. ENDOMYCETALES 141 In other species which terminate the isogamous series, E. javanensis and E. capsularis, the copulation branches have entirely disappeared (Fig. 21, 6-8). According to the conditions of environment, either the hyphal or the sprouting condition may prevail. By swelling of the terminal cells of the hyphae or by lateral sprouting (sometimes also intercalary), there arise 4- spored asci. Each of these spores is divided into two unequal parts by annular thickenings (Guilliermond 1909). Neither species is possessed of much fer- mentative ability. From Endomycopsis two lines diverge. In Hansemda the ascospore is cucullate, the mycelium has disappeared, although the cells are quite elongate and sometimes in chains. Two of the sprout cells form copulation tubes to- Fig. 21. — Endomycopsis Lindneri. 1-5, E. capsularis ; 6-8, development of asci. (1, 2, 6-8 X470 ; 3-5 X500.) (After Mang-enot 1922 and Guilliermond 1909.) ward each other, the nuclei migrate into the bridge and fuse, the diploid nucleus divides, both daughter nuclei migrate back into the cells, there divide a second time, and develop two ascospores in each fusion cell. In Hansenio- spora the cells of the sprouts mycelium are mostly citriform, sprouting only from the poles. Copulation has not been observed and apparently the asco- spores develop parthenogenetically. Pijper (1928) reports H. Guilliermondii from the nails in a case of onychomycosis. In the other line diverging from Endomycopsis, the ring about the ascospore occupies an equatorial position. In Williopsis saturnus we have vegetative conditions very much as in Hansenula, where this species was formerly placed. M2 MEDICAL MYCOLOGY Copulation is reported to occur between ascospores before germination. The end member of this series is Schwarmiomyces, where the spore is rough as well as saturnine, the spore number is usually 1 per ascus, very rarely 2. Projec- tions simulating copulatory canals are produced, but the asci develop par- thenogenetically. Key to Genera IVEycelium present and well developed, no assimilation of nitrate. Copulation heterogamous ; no sprout mycelium, although the cells of the mycelium break apart into arthrospores; a thin pellicle on malt, also on ethyl alcohol. Endomyces. Copulation isogamous, sprout mycelium also present, pellicle thick, often gelatinous, on malt, none on ethyl alcohol. Endomycopsis. Mycelium absent although a pseudomycelium of sprout cells may be formed. Ascospores cucuUate. Yeast cells elongate, or ovoid ; copulation present ; nitrates assimilated ; thick, wrinkled pellicle on malt, often dry and powdery, fermentation of sugars positive, pellicle on ethyl alcohol. Hansenula. Yeast cells citriform, sprouting only from the poles, no copulation, nitrates not assimilated, no pellicle on malt, only weak fermentation of glucose, no growth on ethyl alcohol. Hanseniospora. Ascospores saturnine, yeast cells ovoid or spherical, copulation of ascospores before germination reported, nitrates assimilated; thick, wrinkled pellicle on malt, fermentation of sugars positive, pellicle on ethyl alcohol. Williopsis. Ascospores saturnine and rough, yeast cells ovoid, copulatory canals formed but not functional, nitrates not assimilated, usually only a thin ring on malt, occasionally a slimy pellicle; fermentation of sugars positive, very poor growth on ethyl alcohol. Schwanniomyces. HANSENULA Hansenula Sydow, Ann. Myc. 17: 44, 1919. Willia Hansen, Centralbl. Bakt. II, 12: 529, 1904 not Willia Miill. The type species is Hansenula anomala (Hansen) Sydow. Yeast cells ovoid to elongate, multiplication by sprouting, the sprout cells often clinging together in chains. On liquid media containing sugar a thick pellicle formed, dry and dull from the included air; fermentation of sugars positive, aesculin hydrolyzed, nitrate assimilated, good growth with pellicle formation on ethyl alcohol; ascospores eucullate, 1-4 per ascus. Hansenula anomala (Hansen) Sydow, Ann. Myc. 17: 44, 1919. Saccharomyces anomalus Hansen, C. R. Trav. Lab. Carlsberg 3: 44, 1891. Ann. Micrographie 3: 467-474, 1891. Willia anomala Hansen, Centralbl. Bakt. II, 12: 529, 1904. Willia anomala var. Beauverie and Lesienr, Jour. Pliysiol. Path. Gen. 14: 991, 992, 1912. Saccharomyces Beauveriei Froilano de Mello, Arq. Ilig. Pat. Exot. 6: 246, 1918 [based on case of Beauverie & Lesieur 1912]. ENDOMYCETALES 143 Reported occasionally from sputum but in none of the cases has patho- genicity been clearly shown. Beauverie & Lesieur (1912) reported a case of phthisis with the organism in the mucopurulent sputum. Grigorakis and Peju (1922) also report a case. Shrewsbury (1930), in a monograph on the genus, reports his strain 209 isolated from sputum from a case of chronic bronchitis along with Monilia, and an unidentified yeast. Shrewsbury was unable to find any pathogenicity for his strain on experimental animals. Pseudomycelium formed on many media, especially in pellicles on liquid media. On carrot at 26° C. in 6 days, cells spherical or ovoid, 3-8 x 2.5-6/*. Sporulation on carrot juice in the pellicle after 55 hours, cells 3-8/a in diameter. Asci spherical, ascospores typically cucullate. Colonies cream white, smooth, and shining. Usually developing a strong odor of fruit ethers. Hansenula bispora (Mattlet) Nannizzi, Tratt. Micopatol. Umana [Pol- lacci] 4: 134, 1934. Saccharomyces (Willia) Mspora Mattlet, Ann. Soc. Beige Med. Trop. 6: 32, 33, 1926. Isolated from stools of patients with various degrees of dysentery. Patho- genicity not proved; animal experiments not yet reported in detail. In potato decoction at 37° C. after 3 days, spherical or ovoid cells con- taining a vacuole and granulations, 2-10/*, budding, larger cells 6-7/* or rarely pyriform 11 x 7/*, thick-walled granular with little or no vacuole, other cells elongate, swollen at one end, the smaller tip of one against the swollen tip of the other and showing isogamous copulation. After about 10 days, oil droplets abundant in most of the cells or occasionally droplets unite into one large drop, copulation forms rarer. Little change in appearance after 30 days. On Gorodkova agar asci appear after 5 days, thick-walled, containing 2 cucullate ascospores 6 x 3/*. On Sabouraud agar colonies white dull, circular with even margin, in age the center becomes yellow and a few radiating folds develop. On gelatin stab, slight liquefaction with some gas formation. In potato decoction, abundant deposit of yellowish white clots which dissolve in the liquid on shaking. Optimum temperature 37° C. Litmus milk slightly acid. Acid and gas on glucose, fructose, maltose, galactose and sucrose; very slight acid on dextrin, no action on lactose and mannite. HANSENIOSPORA Hanseniospora Zikes, Centvalbl. II, 30: 145, 1911. Yeast cells citriform or elongate ovoid, vegetative multiplication by bipo- lar sprouting; no pellicle on malt, spores spherical at first, then cucullate (ability to form spores easily lost on cultivation), only glucose slightly fer- mented, nitrate not assimilated, practically no growth on ethyl alcohol. Hanseniospora Guilliermondii Pijper, Proc. Sect. Sei. K. Akad. Wetensch. Amsterdam 31: 989-992, 2 figs., 1928. Isolated from onychomycosis of European woman in Pretoria; patho- genicity quite probable but not proved. 144 MEDICAL MYCOLOGY Growth good at both room temperature and at 37° C. On fluid media cells citriform or ellipsoid, 5.2 x 2.4/*. In old cultures a slight tendency to- ward "mycelium" formation. Vegetative development by sprouting. No trace of sexual process, spores normally 4 per ascus, cucuUate, becoming spherical on germination. A tubelike protuberance is put out which leaves the spore and takes on the characters of the ordinary* vegetative cell. While sexuality has disappeared there is evidently some physiologic differentiation of the spores. After mordanting with chromic acid, staining with carbol- Fig. 22. — Dipodascus albidus. 1, 2, young copulation branches not yet abjointed ; S, h< diploid nucleus in female copulation ; 5, first step in division of diploid nucleus ; 6, later stage ; 7, 8, later stages of young ascus ; 9, upper portion of nearly mature ascus, the dark points in- dicating degenerate nuclei. (i, 2, 5-9 X900 ; 3 X800 ; 4 X600.) (After Dangeard 1907, Juel 1902.) fuchsin, decolorizing with sulphuric acid, and counterstaining with methylene blue, the equatorial pair of spores were acid-fast while the polar pair were blue. Giant colonies on malt agar, grayish brown, edge lobulated, surface smooth showing very delicate concentric rings corresponding to the days of growth, no radial lines, a central knob present from the beginning and later secondary knobs appear at various places. ENDOMYCETALES 145 Acid formed in raffinose, sorbite, and dextrin and a small amount of gas in glucose and fructose. No action on any of the other common sugars and glucosides. Malt gelatin liquefied slowly. On all liquid media, growth took place at the bottom only, no pellicle formed, even after many months. Dipodascaceae.- — In Dipodascus, the mycelium is septate, but the cells are coenocytic. The copulation of two hyphal cells produces gametangia (Fig. 22, 1-2). Fusion of the gametangial nuclei occurs without differentiation of the gametes. The formation of diploid mycelium has disappeared and the ascospores are formed in the gametangium without differentiation of an ascus (Fig. 22, 3-7). From this stage we have two main lines of divergence, one through Eremascus to the true yeasts, and one through Pericystis to the Coc- cidioideaceae. A third possible line has ended blindly in Actonia, a genus whose life cycle is not well known. In Actonia tropicalis, the mycelium is septate and may multiply by sprout- ing. A round gametangium (sporangium) forms at the end of a filament and develops small motile gametes (zoospores), which are apparently forced out of the gametangium by the invagination of the basal wall. "Whether this phenomenon is comparable to the proliferation of Ascoidea or is related to columellar formation in the Mucorales is uncertain. The further fate of the gametes ("zygospores") was not described. Whorls of sprout cells are some- times produced on the hyphae. After some weeks on Kaulin's medium, asco- spores (?) are formed. "At the end of a filament a round body appears with a well-marked external capsule and a small green body in its center. This body becomes flat and disklike, and from it four ascospores bud off. The four ascospores are surrounded by a limiting membrane which is extremely diffi- cult to see because of its transparency. The greatest care has to be taken not to rupture the membrane. "When rupture does occur, the membrane is coiled up under the parent cell and appears to be double" (Acton 1919). This, if correctly reported by its author, is quite similar to the peculiar spore for- mation we find in Paracoccidioides and reminiscent of partial sporangial forma- tion in the higher Mucoraceae. "When the morphology and cytology of this organism is better known, perhaps it will be found related to Paracoccidioides and Histoplasma rather than to the Dipodascaceae. In Pericystis alvei (Betts 1912, Claussen 1921) the mycelium is differen- tiated sexually, and copulation is heterogamous. Its cytology is unknown, but the ascus is spherical and contains many spores, suggestive of conditions found in the Coccidioideaceae-Taphrinaceae line. ACTONIA Actonia Dodge, n. gen. Mycelium cellulis curtis, crassis gemmiparis; zoosporangia terminalia, zoosporis motilibus, sphericis, e zoosporangiis ab basis invaginatione ejectis;, conidia verticillata, ovoidea gemmipara ; hypnosporae endogenae in hyphis veteribus; asci terminales, spherici, tetraspori. 146 MEDICAL MYCOLOGY Mycelium of short, stout cells, often multiplying by sprouting; zoospo- rangia terminal, producing motile, spherical zoospores which are ejected from the sporangium by the invagination of the basal wall ; conidia in whorls, ovoid, germinating by sprouting; endogenous hypnospores formed on old mycelium. Asci terminal, spherical, 4-spored. Type species is Enclomyces tropicalis Acton non Castellani. If the observations of the author are correct, this is a very curious fungus with a life cycle quite different from any other genus of fungi known, and the only place where motile gametes or zoospores have persisted in the Ascomycetes. It is possible, however, that flagelliform appendages of gametes similar to those in Spermophtliora were observed, or else that a contamination from some member of the Phycomycetes has been confused with some asco- genous fungus. Actonia tropicalis (Acton) Dodge, n. comb. Endomyces iropicalis Acton, Indian Journ. Med. Kes. 6: 591-600, 1919; not Endomyces tropicalis Castellani, Centralbl. Bakt. I, 58: 236-238, 1911. Monilia Actoni Vuillemin, Champ. Paras. Homme Anim. 84, 1931 nom. nud. Producing small creamy patches on the tonsils and uvula in throats of soldiers in Mesopotamia. The patches are difficult to remove but leave no raw bleeding surfaces. There is diffuse inflammation of the uvula, pillars of the fauces, and the posterior pharyngeal wall. In debilitated persons it may extend to the bronchi and bronchioles, causing fatal bronchopneumonia. Both sprout cells and mycelium present ; gametes produced in spherical sporangia. Their development is not altogether clear from Acton's descrip- tion, some of the phenomena suggesting proliferating sporangia of Ascoidea or of the Saprolegniaceae. Chlamydospores present. Ascospores budded off the ascus somewhat as in Paracoccidioides. On 1% sucrose agar, colony ivory white with raised crenate edges; be- coming creamy yellow and sticky in a few daj's, mycelium also penetrating the agar. On litmus milk, groAvth scanty, acid on the third day without coagu- lation. On Kaulin's solution, slight cream colored growth at the bottom of the tube. Ascospores in 2-6 weeks. On carrot, growth luxuriant, sticky, brown. From the stage attained by the Ascoideaceae two other lines of develop- ment diverge. One line has retained the large number of spores in the ascus, developed the ascus as a thick-walled resting spore, with a tendency to delay spore formation until the protoplasm has slipped out of the ascus. The nuclear history of most members of this line is so little known that there is doubt -j.- to whether sex has been retained, although the large nucleus (in some species) in the very young ascus suggests that a fertilization has taken place and the occurrence of spores in tetrads during one stage of development, in several genera, suggests a reduction division. Finally spore number is reduced in some species of the Taphrinaceae to 4 or 8, but so many intermediate forms exist and the number is so inconstant that it has been abandoned as a generic character. Along with this goes the elimination of the thick-walled resting ENDOMYCETALES 147 stag-e of the young- ascus. The collection of asci into more or less definite fructifications or ascocarps within the host tissue has been attempted in the end member (Taphrinaceae) where, in some species the clavate asci form a palisade layer under the epidermis of the host. The other line has early reduced the number of ascospores to a small, definite number, 8, 4, 2, or 1 and, perhaps owing to their habitats, have gradually diminished the amount of mycelium until in the Saccharomycetaceae, true mycelium has disappeared. For further discussion of this line, especially the steps in the gradual disappearance of sexuality, see Chapters XI and XII. Coccidioideaceae.- — This family has four genera causing more or less similar lesions in man and in experimental animals, with no saprophytic species so far recorded. Three of the genera are monotypic, from widely separated and rather restricted localities. Coccidioides is mostly restricted to California, Uruguay, and Argentina. Paracoccidioides to Brazil, while Rhino- sporidium has been reported from Argentina, India, and the Mississippi Valley. Histoplasma is known in the ^Mississippi Valley and in Panama. The large indefinite number of ascospores (Fig. 23, 19) suggests condi- tions found in the Ascoideaceae, but no conidial stage is knoAvn. Along with increasing specialization for strict parasitism in this family, sexuality has ap- parently disappeared without a trace.* The mycelium is septate and multinu- cleate as in the preceding family. In the host, however, it tends to less and less development until occasionally^ in Coccidioianisni as iMarcone and Tokishi<>'e. In 3 906, 8an Felice succeeded in g-rowing mierocolonies, but was unable to keep them alive very long, although he was able to reproduce the disease. In spite of these seem- ing successes, the next few years saw several hypotheses that the organism was a protozoon, the authors explaining the presence of hyphae in the cultures of earlier workers as contaminations in spite of the observations of San Felice who controlled his studies by microscopic examination. In 1916, Lindner and Knutli renamed the organism Monilia co.psidata. During- the decade of the World War, Boquet and Negre and their coworkers made a very thorough study of the problem and developed methods for its cultivation. The last decade has seen their work confirmed by several German workers. In the tissues, cells are spherical or ovoid, or acuminate at the two poles, sprouting, 3-5 x 2.5-3.5(U in diameter, membrane of variable thickness, granular. Occasionally elongate cells seen or larger cells, 5-7/t in diameter. In cultures, hyphae 2/i, in diameter, septa about 10-20/u apart, finely granu- lar, no oil globules, branched. Yeast cells pyriform, thin-walled, at first, becoming thick-walled after the cell separates from the parent cell, containing oil globules which may be large as the cell increases in size up to 8-12^ or even 15-16//,. Thick-walled hyphae formed from these yeast cells, 3-4// in diameter, with septa 10-18// apart and oil globules present. Chlamj^dospores from thick-walled mycelium, usually terminal, slightly polyhedral, 10-18//, protoplasm granular without oil globules. In old cultures the mycelium breaks up into somewhat irregular, thick-walled arthrospores. Asci 4-spored. [The ascospores figured by Tokishige were undoubtedly drops of oil.] Everbeck (1926) reports ascospores, thick-walled, ovoid, 3 x 2//, regularly produced after 14 weeks. In tissues from animal inoculations, yeast cells 3-4//, or, when actively sprouting, 5-6//, in groups of 10-15, and some thick-walled hyiDhae. The anti- bodies appear about the twentieth day of the disease and remain a long time after cure, so that it is very difficult to inoculate a horse which has had the disease. For isolation and early subcultures Boquet & Negre report best results with the following medium : Macerate 400 gm. horse dung in 2,000 c.c. of water for 24 hours in a cool place ; strain through cheesecloth, squeeze, filter, and add 10 gm. peptone and 18 gm. agar for each 1,000 c.c. of filtrate. Sterilize for 30 minutes at 120° C., add 40 gm. glucose per 1,000 c.c, tube and sterilize for 20 minutes at 115° C. For isolation add 20 drops of the following solution to the tubes after inocu- lation : chop 100 gm. lymphatic ganglions of the horse, macerate for 24 hours in 500 c.c. water, strain through cheesecloth, squeeze, filter, add 20 gm. glucose, tube and sterilize for 30 minutes at 115° C. Moisten cultures with this liquid as they begin to dry out. Incubate at 25-30° C. On dung agar, after 4-6 weeks, colonies appear on surface, as small, round, elevated, grayish white, slightly velvety, the size of a pinhead. The colonies 172 MEDICAL MYCOLOGY grow in height and at the periphery and turn brown, becoming contorted and scattered with small white, velvety points, with a white velvety area at the periphery which is festooned. Colonies hard, compact, adherent to the agar. On Sabouraud agar moistened with maceration of ganglia, the young colonies appear as on dung agar, the older colonies more folded and yellowish white, sandy, somewhat darker in age. Time of incubation shorter on suc- cessive subcultures (first in 1 month, fifth in 3 weeks, tenth in 10 days). Be- tween the fifth and tenth subcultures, organism is inoculable into ordinary agar, glucose agar, malt agar, with same aspect as on Sabouraud agar, but less abundant and less velvety growth. On carrot and potato, colony elevated, slightly folded, brownish gray, darkening in age, moist, smooth. On g-elatin, white velvety colonies, medium rapidly liquefied. On milk growth very slow, milk coagulated. Mycelium produced in the water of condensation of horse or sheep serum with 6% glycerol, horse serum agar, ordinary peptone agar, or glucose bean (2% agar with 20 drops of bean decoction added). The mycelium is oidiform, irregular with chlamydospores. While liquid media are not suitable for isolation or early subcultures, growth is possible after a time on peptone (1%) and glucose (5%), if the inoculum is floated on the surface and the culture incubated at 35° C. There is formed a thick whitish pellicle composed of yeast cells, spherical or ovoid, overlying the limpid liquid. Finally mycelium appears in the pellicle. If 0.5% agar is added to increase the viscosity, the pellicle is thick, folded, snow white, composed of yeast cells and, especially, thick-walled hyphae similar to those on Sabouraud agar. The optimum temperature is 37° C, but the medium dries out too rapidly; growth at this temperature is about twice as fast as at 25-30° C. At the higher temperature, the colonies are softer, more spongy, with more velvet, and not so adherent to the substrate. The maximum temperature is 38-40° and the minimum 15-18° C. At lower temperatures growth is very slow in the early subcultures, becoming more rapid after the organism has become adjusted to artificial media. At room temperatures, the colonies are elevated, folded, white, powdery or velvety, composed of very thin-walled hyphae and some sprout cells. After some weeks, short thick-walled hyphae are seen and the septa are more evident. In liquid media development is very slow at 20-25° C. If a culture is removed from 35-36° to 20-25°, it produces a grayish folded pellicle, the yeast cells cease budding and produce slender thin-walled hyphae as on glucose. If the culture is removed from 20-25° to 30-36°, the number of yeast cells increases. In low oxygen tension (under a layer of oil) growth is very slow, large irregular cells 12-15/a in diameter, thick-walled, granular, often in chains of 5-6 cells with large oil globules. Citric acid up to 1 :2,000 favors growth, producing a grayish folded pellicle or small round white colonies floating on or in the medium. Mycelial forms at low tempera- tures similar to those seen in pus. Bierbaum (1919) found slow growth on slightly alkaline horse meat agar with 2% glucose, 2.5% glycerol, and 3-4 c.c. EREMASCACEAE 173 sterile horse serum. Lange (1921) used egg medium with 2% glucose and 1% glycerol, reporting colonies small, brownish yellow, dry, center elevated, margin flattened, edge like a rampart, confluent near water of condensation. No fermentation of sugars, only glucose and sucrose utilized. Ammonia produced from peptone. Milk coagulated, gelatin rapidly liquefied. Zymonema Molardi (Salvat & Fontoynont) Dodge, n. comb. Endomyces Molardi Salvat & Fontoynont, Bull. Soc. Path. Exot. 15: 311- 320, 1922. Endomyces Molardi Fontoynont (nomen nudum) Bull. Mem. Soc. Chirurg. Paris 48: 439-442, 1922. Isolated from lesion on leg of man inoculated by scratching, (?) at first a fistula, then an ulcer, with depigmentation. Lesion finally cured by ex- ternal applications of methylene blue for about one month. Inoculation on thigh of a guinea pig caused abscess which healed spontaneously. Intraperi- toneal injection caused a nephritis which was fatal in 34-44 days. Organism recovered from the blood. Hyphae occasionally 2-3 fields long, 2.5-3.5/i. in diameter, irregularly septate or fragmented, appearing- nearly empty, tinted slightly pale rose by Gram's method. Yeast forms visible. Hyphae 2-6fx in diameter, contracted as septa, thick-walled, very rich in glycogen. Filament may terminate in a simple cell, a group of short, thick-walled, nearly round cells, one to two times as thick as the penultimate cell; or it may end by a spherical cell, 8-22/t in diameter, thick-walled, staining with aqueous eosin (chlamydospore). These chlamydo- spores appear occasionally in groups or short chains. Ascospores appear on glycerol agar or in liquid of glycerol-carrot. Asci spherical or ellipsoid, 4- spored, rarely 8-spored. On glycerol agar, growth rapid, colony thick, creamy, ivory white, ele- vated with little deeper yellow craters in the middle and never reaching the side of the tube. On Sabouraud maltose, growth as on glycerol but colony dirty, yellowish white. On Sabouraud glucose, growth less abundant, with slightly elevated polycyclic plateau, center creamy, white, glistening, then opaque. On fifteenth day fine radiations from central cone. On carrot with glycerol, from streak, growth appears like white porcelain, is thick, glistening, creamy, composed of a mass of granular colonies. Hyphae on walls of tube in fifth month, the whole surface is invaded and becomes crateriform, moist below, dryer above. Growth on potato and turnip with glycerol same as for carrot. Growth from gelatin stab appears like inverted fir tree. On surface plateau 8 mm. in diameter, three zones: (1) outer, 1 mm. broad, fine radiat- ing lines, (2) higher, 3 mm. wide, abrupt drop to outer zone, (3) uppermost, 3 mm. broad, granular. On horse serum, growth is meager, old ivory in color. On solid media, only yeastlike cells appear. In liquids a slight pellicle and very fragile ring appears, sediment is flocculent and abundant, liquid remains clear. On lactose bouillon, growth is good ; even after 5 months lactose is not exhausted. Medium gradually becomes acid. Growth in glycerol bouillon 174 MEDICAL MYCOLOGY poor. In Raulin's liquid, growth is fair, white sediment, liquid clear. In Gedo- elst liquid, growth very good. Coagulated serum not liquefied. Gelatin not liquefied in 30 days. Zymonema crateriforme (Iludelo, Sartory & Montlaur) Dodge, n. comb. Endomyces crateriforme Hudelo, Sartory & Montlaur, C. R. Acad. Sci. 170: 1086-1088, 1920. Saccharomyces, sp. Hudelo, Sartory & Montlaur, Bull. Sci. Pharmacol. 25: 352-357, 1918. Isolated from a lesion in the armpit of a young woman. Lesion oval, 3 cm. in diameter, erythematous, scaly, reminding one of dry seborrheic eczema. Nonpathogenic to rabbits and guinea pigs, by either subcutaneous, intraperi- toneal, or intravenous injections. On scarification an atypical, evanescent lesion results. Submerg'ed mycelium shows yeastlike budding cells, 9-11/a, or oidial cells, 15-16/A long. Conidia sprouting from aerial mycelial branches on some sort of sterigmata are variable but usually ellipsoid or ovoid, 5-7 x 6-9//.. Asci appear on solid media and old cultures after 80 hours at 22-23° C. These are usually terminal, rarely intercalary, 4-5/x in diameter, containing 4 spores each. Spores 3-3.5 x 1.75-2/a, with single smooth membrane. On Sabouraud agar, colony is small and white, becoming- creamy white, center yellow and irregularly crateriform, finally crackled, appearing- like small sponge. On carrot or potato, colony broader, thicker with several small craters, very much folded, cream white, covered with conidia. On broth gelatin, colony not fully developed before liquefaction on seventh day. After 18 days on malt extract, colony appears as thick, dry, folded mat covered with conidia, sediment of yeast cells, feeble fermentation with slightly aromatic odor. Growth similar on beef broth, peptone + glycerol + glucose broth, normal Raulin's solution with g-lucose, lactose, galactose or maltose, or on prune decoction. Sucrose, glucose, fructose fermented, not rafSnose, lactose, galactose, or maltose. Starch paste not liquefied. Milk not coagulated in 34 days. Gelatin liquefied. Zymonema albicans (Okabe) Dodge, n. comb. Endomyces albicans Okabe, Centralbl. Bakt. I, 111: 181-187, 1 pi., 1929, non aliorum. Isolated from 49 cases of thrush, in Japan, and pathogenicity of this strain proved. ["Monilia Candida," a filamentous species with strong- fermentation of sucrose and a strain wdiich completely failed to ferment, also isolated from some of the above cases, but proved to be nonpathogenic] Yeast cells spherical or ovoid, 4-6/a in diameter, with giant cells in old cultures reaching 15-20/x, after a time elongating and forming mycelium. Mycelium 3-4/i, in diameter, cells 20-30/x long, rarely up to 80/x. Chlamydo- spores pyriform, spherical or ovoid, 7-12/i, in diameter. Asci in 45 strains 1-spored, rarely 2-4-spored, 5-6/a in diameter; ascospores spherical to ellipsoid, 3.5-4 X 2.5-3/i., thick-walled, asci found only in the yeast stage, not on the mycelium, produced only on koji extract agar after 4-6 months. EREMASCACEAE 175 On koji extract agar, colony thin, milky white, smooth, surface moist with sharp borders, becoming thicker, duller, yellowish white ; on drying', becoming brownish yellow witli an ashy gray powder, mycelium very scarce. On 10% sucrose peptone agar, mycelium produced, colony verinicose and folded. No pel- licle on liquid media, powdery sediment. On sucrose peptone solution, mycelium floating as woolly masses hanging from the ring. By Lendner method, fermen- tation with glucose, levulose, mannose, maltose shows strong acid and gas ; dextrin and galactose show slight acid and gas, soon disappearing; no action on other sugars tried. Growth better at 37° C, than at 25° C. ; growth pos- sible in ice box or at 40°. Zymonema bonaerense (Greco) Dodge, n. comb. Endoniyces bonaerensis Greco, Argentina medica 1908 ; Origine des Tumeurs. 123-156, 412-421, Figs. 54-68, 225-231, 1916. Producing small miliary abscesses and plaques near eye. Pathogenic for rabbit, forming subcutaneous abscesses. Yeast cells 4-6 x 2-4/^, ovoid, hyphae mostly 2-4^ in diameter, occasionally much thicker and shorter. Asci 4-12/x spherical, spores 1-2/a. Colonies milk-white, shining-, discrete, or becoming confluent. On solid media after about a month, hyphae beg-in to show in the media. No gas but an ethereal odor in some cultures. On Sabouraud agar, growth at 20° and 37° white, creamy, a little thicker in the center, with filaments at the periphery. On beef agar and glycerol agar, growth less, colony flatter, more opaque, less shining white. On potato, colony creamy, very moist, white, later more opaque and grayish. On drying, small secondary yeast colonies form on surface of mother colony. Potato glycerol shows better growth, dryer, mammillate or slightly crateriform, slightly yellowish and cheesy in appearance, medium darkened. On carrot, colony creamy, spreading, white margin toothed. On liquid media no discoloration of media, no pellicle, but sediment produced. Sugars not fermented. Milk coagulated very slowly. Colony on gelatin stab resembles a test tube brush, without radial filaments, no liquefaction. The figures and description are not altogether convincing as to presence of ascospores, and I have hesitated to transfer it to this genus, but it is quite evidently not Endoniyces. Zymonema album, Dodge, n. sp. Monilia sp. Bianchi & Niilo, Bol. Inst. Ch'n. Quirurg. Univ. Buenos Aires 4: 531-538, 6 figs., 1927. Producing bronchomj^cosis. Pathogenic for guinea pig. Spherical yeast cells about Sfi in diameter; ovoid cells 3-5 x 2/*. Cells 10-12;u, in diameter, thick-walled with four deeply staining bodies which may be ascospores. In old cultures on glycerol, carrot, and potato, there are slender mycelium and large chlamydospores. This mycelial form continues on sub- culture. Colonies hemispheric, creamy white, moist, finally becoming confluent with a dryer, whiter margin. Optimum temperature 37°, but fair growth at 20° C. 176 MEDICAL MYCOLOGY On potato glycerol, medium, darkened; serum not liquefied, milk coagulated in 24 hours. In Raulin's solution, sediment, but not turbidity. Drigalski- Conradi slightly acid in 4 days. On neutral red agar, gas in 24 hours, color change in 4 days. No indol. No fermentation of sugars. Acid produced from glucose, fructose, galactose, lactose, and arabinose, no action on mannite. sucrose, raffinose, and inulin. Zymonema bucalis (Niiio & Puglisi) Dodge, n. comb. Monilia hucalis Niiio & Puglisi, Semana Med. 34: 222-229, 1927. The lesions began as perleche, extending gradually over the buccal mucosa which became covered by white plaques, slightly adherent, not continuous, suggesting clots of curdled milk, extending into the tonsillar crypts, produc- ing a burning sensation. Yeast cells and hyphae 2.5-3/x, little branched. In cultures yeast cells 3-7 X 2-5/x; asci spherical, 10/a in diameter, with 2-4 ascospores. Hyphae de- veloping from thick-walled cells, flexuous, little branched, 1.5-3/a in diameter; blastospores lateral; large terminal ehlamydospores present. Colonies at 37° C. on Sabouraud agar, potato-8% glycerol, carrot, and potato agar, circular, confluent, moist, white, little elevated. On liquid media (broth, potato decoction, 2% peptone solution and maltose-peptone solution) turbidity, with the formation of white clots, which settle to the bottom. Drigalski-Conradi medium becomes red, then decolorized; litmus sugar agars become red, then decolorized, and finally become blue. Acid on glucose, sucrose, and lactose, not on mannite. Sucrose not inverted, starch digested. Sugars not fermented. Milk coagulated on the third day, no liquefaction of gelatin nor of coagulated serum. Zymonema Cruzi (Froilano de Mello & Paes) Dodge, n. comb. Endomyces Cruzi Froilano de Mello & Paes, Arq. Hig. Pat. Exot. 6 : 51-60, 1918. Isolated from sputum of patient suffering from bronchitis and asthma. In sputum cells 4-8 x 2-4/^, rarely spherical, with refringent granules. No hyphae or spores present. On potato, cells ovoid or spherical, pseudomycelia 50-80/A long', simple or branched. Spherical or reniform asci present, 2-4- spored. On simple agar, spherical cells in chains of 8 to 16, no asci. On 1% agar + 6 drops of 10% NaOH, mycelial filaments definitely septate, 100/A long with terminal ehlamydospores or lateral conidia at the septa, and arthrospore formation. On 10 gm. agar + 5 drops 10% NaOH, some yeast cells, mycelium, terminal ehlamydospores. On 10 gm. agar + 4 drops 10% NaOH, feeble development, yeast cells, no true mycelium, asci abundant. On simple agar, feeble development, small milk-white colonies. On mal- tose agar (Sabouraud), colony white, then yellowish, verrucose. Colonies on potato, circular, elevated, wax color, opaque. Broth becomes cloudy after 72 hours with flocci in suspension, forming a filiform sediment. On Raulin liquid, thin whitish pellicle becoming thicker and chalky spotted, with small white EREMASCACEAE 177 points above and below with streamers 1-2 cm. long protruding into the liquid, finally breaking off and forming filiform sediment at bottom of the tube. Glucose, maltose, and sucrose fermented. Zymonema Alvarezsotoi (Mazza & Niiio) Dodge, n. comb. Monilia Alvarezsotoi Mazza & Niiio, apud ]\Iazza, Niiio, Quintana & Bem- aseoni, Bol. Inst. Clin. Quiriirg. Univ. Buenos Aires 6: 180-214, 38 figs., 1931. Generalized mycosis of native of Argentina. Fungus isolated from caseous lumps in feces, from lumps in the sputum and from the urine. Patient finally succumbed. Pathogenic to white rat, guinea pig, in intraperitoneal injection, to rabbit intravenously or applied to the mucous membranes. In one rabbit subcutaneous injection resulted in subcutaneous nodules which were surgi- cally removed. The animal recovered completely. On Sabouraud solid media, yeast cells mostly round, variable in size, some sprouting. No hyphae or asci observed. On coagulated human serum, after one month, many yeast cells, mostly round, 1-10/a in diameter. Rare ovoid forms. Few hyphae, composed of chains of fusiform, granular cells of vari- able size. Lateral branches at septa. Occasionally two large cells, approxi- mately of equal size, and with thick membrane, are joined by a narrow neck [isogamous copulation?]. Morphology in Gougerot gelatin slightly different. Hyphae predominate. These are variable in size, flexuous, composed of chains with lateral ramifications. Some spherical or ovoid cells are united directly to the hyphae or by little pedicels [immature asci?]. There are intercalary or terminal swellings resembling chlamydospores. Some cells are cylindric with rounded ends, 10 x 2/i,, vacuolate. In some filaments, internal spores like oidia. Optimum temperature for growth 30-37° C, no growth at 50° C. Gram-positive or negative. With Leishman stain, protoplasm sky blue, chromatin granules garnet. On Sabouraud glucose agar, colony rugose yellowish. Same on Sabouraud maltose. Good growth on agar and carrot. On potato, growth also good with darkening of medium. No growth on Drigalski medium. On potato, with 8% glycerol, good growth in 24 hours with darkening of medium. Colony white, humid, covering the whole surface. Pellicle on top of liquid and deposit at bottom of tube. On carrot, with 8% glycerol, abundant growth in 24 hours, colonies white, humid, covering whole surface of medium. Some days later on upper part of edge some white, agglutinated hyphae appear. Pellicle on surface of liquid ascending walls of tube. On Gougerot gelatin stab, rugose, caramel-colored pellicle on surface of medium, presently darkening on top. On plain gelatin stab at 15° C, dendroid growth along whole length of stab. On gelatin streak, good growth, best at the bottom where some lateral arbor- escences appear. Along coagulated human serum stab, growth slow and dend- roid. In plain broth and Sabouraud broth, turbidity and sediment in clots. In acid Raulin's solution, sediment in clots without turbidity. Starch paste unaltered. No fermentation of sucrose, rafiinose, lactose, maltose, mannite, sorbite, dextrin, or inulin. Slight acidity but no fermentation with glucose, 178 MEDICAL MYCOLOGY fructose, and galactose. No indol formation. Milk not coagulated. Coagu- lated human serum slowly liquefied. No effect on gelatin in 30 days. While no asci or ascospores have been reported in the following species, its pathogenicity, its cultural characters, and morphology all place it in Zymonema rather than Mycoderma, very close to Z. dermatitidis. Zymonema Harteri (Verdun) Dodge, n. comb. Cryptococcus Harteri Verdun, Precis ParasitoL, 1912. Atelosaccharomyces Harteri Beurmann & Gougerot, 1913. Parasaccharomyces Harteri Froilano de Mello, Paes & Sousa, Arq. Hig. Pat. Exot. 6: 33, 1918. Mycelohlastanon Harteri Ota, Jap. Jour. Derm. Urol. 28: [4] 1928. Fig-. 35. — Zymonema Harteri. (After Pollacci & Nannizzi 1926.) Monilia Harteri Vuillemin, Champ. Paras. Homme. Anim. 85, 1931. Mycotorula albicans Langeron & Talice, Ann. ParasitoL Hum. Comp. 10: 47, 1932, jyro parte. Torulopsis Harteri Almeida, Annaes Fac. Med. Sao Paulo 9: 10, 1933. Isolated from a case of generalized blastomycosis presumably contracted in Cochin China. The organism has evidently invaded the intestinal tract, liver, lungs and finally produced subcutaneous nodules. [Case of Harter, C. K. Soc. Biol. 64: 241-242, 1908; De la blastomycose humaine. These Fac. Med. Univ. Nancy 20: 1-222, 1909.] Pathogenic for rabbits and mice. Medi- cation with KI ineft'ective. Sprout cells ovoid or ellipsoid, 4-6 x 3-5/x, somewhat larger in liquid media, with some spherical cells in old cultures. On solid media elongate cells EREMASCACEAE 179 8-15 X 5-6/A appear. Hyphae about 2/x in diameter with occasional swellings up to 3/* seen in Raulin's liquid. Chlamydospores 5-8;u, with very thick walls seen on carrot. Asci not observed (Fig. 35). Growth over a Avide temperature range between 10 and 55° C, with good growth at 37° C, and at room temperature. On glycerol agar, growth abun- dant, white, smooth, velvety in the depths ; cells ovoid, with very large vacuoles in old cultures. On sucrose or glucose agar, growth creamy, abun- dant, shining with only ovoid cells. On ordinary agar, growth poor, granular with little penetration of the substrate. On carrot and turnip, growth creamy, white, smooth or granular, becoming mamillate, or even crateriform in old cultures, developing hyphae ; cells at first ovoid, soon becoming elongate and mycelial. On potato, growth slow, grayish, soon dry. On gelatin, colonies white, granular, with dendroid growth in the medium ; cells elongate or ellipsoid in the depths but no hyphae, ovoid at the surface. On blood serum, colony grayish white. In liquid media, floccose growth which slowly settles without producing ring or pellicle. Milk not coagulated, gelatin not liquefied after 8 months. No fermentation, sucrose only slightly inverted. I have been unable to locate the original descriptions of the following species and know them only from secondarj^ sources. Endomyces pulmonalis Senez, Boletin del Lab. de Bact. Tucuman (Argen- tina) 1: 58-60, 1918. [Reviewed in Bull. Inst. Pasteur 17: 636, 1919; Perin, Micosi polmonari 94-95, 1925; Pollacci & Nannizzi, I, Miceti Patogeni 4: No. 34, 1925.] Isolated from sputum of patient suspected to have tuberculosis. Creamy white colonies, asci ovoid, 4-spored, 10/a in diameter. Zymonema histosporocellularis Haberfeld, Tesis, 108 pp., 21 figs., Sao Paulo, 1919. Mycoderma histosporocellularis Neveu-Lemaire, Precis Parasitol. Hum. 70, 1921. This species is said to be a synonym of Paracoccidioides hrasiliensis (Almeida 1933). OLEINA Oleina Tieghem, Jour, de Bot. [Morot] 1: 289-292, 1887. Raquet mycelium Avell developed, intercalary chlamydospores present ; asci spherical, either intercalary or lateral, no trace of sexuality observed ; ascospores, 8 per ascus, varying from ellipsoid to spherical. No type species was designated. 0. nodosa, (Fig. 35, 3) was found grow- ing on fresh cartilage which was floating in olive oil during some studies of saprophytic fungi growing in oil. In this species the asci are intercalary, the spores ellipsoid, 4 x 6/*. 0. lateralis (Fig. 35, i), in which the asci are lateral and the ascospores spherical, 5/i. in diameter, was found on a bit of water- soaked cotton floated on the olive oil in similar experiments. It was cultivated 180 MEDICAL MYCOLOGY also on cartilage and maintained its distinctive characters. Yeast cell stage unknown, probably not normally present, as the author states that the chlamydospores germinate directly by germ tubes. The normal habitat of this genus is rather problematical. The cultures were made before the technic of pure culture had been highly developed, so that probably the cartilage had not been sterilized and the organism may have been present in it, or it may have been present in the surrounding oil. It is to be hoped that these organisms may be again encountered and studied. There is no trace of sexuality observed, so that this may represent an end member of a series. The relationship of Octomyces Froilano de Mello & Gon- zaga Fernandes is also problematical. OCTOMYCES Octomyces Froilano de Mello & Gonzaga Fernandes, Arq. Hig. Pat. Exot. 6: 237-242, 1918. Mycelium septate, sprout mycelium also present ; asci 1-, 2-, 4-, and 8- spored ; chlamydospores terminal. Type species Octomyces Bettencourti Froilano de Mello & Gonzaga Fern- andes. The type was originally isolated from a contamination at Nova Goa, so that nothing is known of its normal habitat. This genus may be considered as a synonym of Oleina, but it differs in the absence of raquet mycelium, the presence of sprout mycelium, and in the terminal rather than intercalary chlamydospores, which seem to be produced only in liquid media. Both genera have been rather poorly described and need much further study before their systematic position will be known. Neither seems to have a well-developed sexuality, such as found in Eremascus and Zymonema. Octomyces Bettencourti Froilano de Mello & Gonzaga Fernandes, Arq. Hig. Pat. Exot. 6: 237-242, 1918. Isolated from contamination in a Petri dish, at Nova Goa. Mycelium septate, yeast cells spherical with brown granules, asci ellip- soid, spherical, and lanceolate, 1-, 2-, 4-, and 8-spored. Chlamydospores terminal, yeast cells not sprouting. On glucose and maltose agar, colony moist, dirty white streak, same morphology as on potato. On potato, colony dry, veiy wavy, margins indented. On carrot, culture dry, very wavy, dirty white, granular in appearance. In broth, easily dissociable pellicle and sediment, no turbidity, with no fermentation, acid with dextrin, fructose, maltose, no action with lactose, mannite, or glucose. Octomyces Etiennei (Potron) Dodge, n. comb. Saccharomyces Etiennei Potron, Rev. Med. de I'Est. 45: 814-826, 841-855, 3 figs., 1913. EREMASCACEAE 181 Isolated from severe pleiiropiilmonary infection and bronchopneumonia with abundant j^easts appearing in the sputum. Producing local pyogenesis in rab- bit and guinea pig. Cells spherical, rarely ellipsoid, mostly 3-9 x 1.5-4.5/i. When actively bud- ding, cells are 15-18 x 4-5/i., with the formation of pseudomycelium but no true mycelium on turnip, liver decoction, and glycosuric urine. In other media, especially carrot and potato, cells separate early. On turnip, filamentous forms found, spherical cells about 6/* in diameter, asci abundant. Irregular allantoid forms found, also durable cells. Elongate forms on glycerol arti- choke as well as on turnip. Budding forms abundant in liquid media, espe- cially in the pellicle of old cultures. No trace of copulation before the formation of the ascospores, which are 4 per ascus and measure 2-2. 5/a. Asci 7-8 x 5ft. The ascospores swell and begin budding, still without any trace of copulation. No especial cultural conditions seem necessary for ascospore formation. On Sabouraud agar, colonies white, punctiform, rapidly confluent, forming a white, creamy, shining, flat surface elevated more than 1 mm.; margins crenulate. Colonies on potato grayish white, punctiform, spherical, very much elevated above the surface. As each colony grows rapidly, it becomes acumi- nate in appearance and is confluent with neighboring colonies. When the medium dries, the colonies become chalky white. On carrot, development is more rapid than on potato. Colonies white, rapidly confluent into a varnished, creamj^ surface. On turnip, growth at first similar to that on potato, then colony becomes prominent, surface mammillate, pebbled, remaining grayish white as the medium dries out. On artichoke, growth is slow but otherwise resembles that on carrot. Glycerol, on vegetable media, somewhat inhibits growth. On gelatin, combined with Lasseur's medium, colonies grayish white, develop slowly and are very slowly confluent, if at all. On liver decoction gelatin, grayish white, elevated, punctiform colonies. Development is rapid and abundant. On gelatin, combined with normal Raulin solution, colonies grayish Avhite, rapidly confluent to give the appearance of shagreen. In pepto- glycerol broth, development is slow, slight turbidity at first, with deposit of flocci at the bottom. Lasseur's medium is not especially favorable, develop- ment is as in pepto-glycerol broth. On Courmont's medium with glucose, development is rapid, sediment abundant, pellicle thick; with galactose and lactose, sediment even more abundant ; with sucrose, sediment less marked, pellicle feeble ; with maltose, sediment very marked, pellicle conspicuous ; with inulin growth, slow at first, with pellicle and sediment finall}^ appearing; with starch deposit, development slow. Grown in Sartory's mutton liver decoc- tion, organism causes grayish floccose sediment, leaving liquid clear. In glycosuric urine, containing 10 gm. glucose per liter, abundant deposit ap- peared, liquid clear with some gas evolution, then appearance of pellicle. Ring also after glucose had fermented. In normal Raulin 's solution, a powdery white deposit appeared on the walls and bottom of container. Fermentation occurred with glucose, fructose, galactose, lactose, maltose, and sucrose. Slight 182 MEDICAL MYCOLOGY action on dextrin, none on inulin, starch, or mannite. Milk very slowly coagu- lated, on the tenth day, with the evolution of C0_, gas. Curd not digested. Gelatin not liquefied. BARGELLINIA Bargellmia Borzi, Malpighia 2: 469-476, 1889. Hyphae very slender, hyaline, irregularly branched, septa remote ; asei solitary, terminal, spherical, membrane thickened, minutely tuberculate sca- brous, more or less brownish, indehiscent, spores spherical or subspheric, soli- tary, rarely 2 per ascus, wall thin, content oleaginous. This genus seems not to have been seen since its original isolation. It was not figured. Some characters suggest that it may be based on a misinterpreta- tion of Hemispora, in which the oil globules have been mistaken for ascospores. Until it is found again and more carefully studied, it should be regarded as doubtful. Bargellinia monospora Borzi, ]\Ialpighia 2: 476, 1889. Isolated from the external auditory conduit in catarrhal otitis. Hyphae subequal, 2-4;* in diameter, with distant septa ; asci more or less distant and indehiscent, 8-12/a. Spores spherical or nearly so, solitary or 2 per ascus, smooth, guttulate, 5-7/u, in diameter. HEMISPORA Hemispora Vuillemin, Bull. Soc. Myc. France 22: 125-130, PI. 7, 1906. Trachyiora Saccardo, Syll. Fung. 4: 262-263, 1886 [as subgenus only]. 8pore7idonema Ciferri & Redaelli, Jour. Trop. Med. Hyg. 37: 167-170, 1934; Redaelli & Ciferri, Atti 1st Bot. R. Univ. Pavia IV, 5: 145-198, 1934 not Desmazieres 1827 nor Oudemans 1885. The type species is Hemispora stellata Vuillemin. In tissues, yeastlike cells; in cultures, hyphae hyaline, septate, producing chlamydospores and conidia (blastospores?) ; asci in chains without trace of sexuality, containing a single echinulate spore. The morphologic interpretation of structures in this genus has long been puzzling. Vuillemin considered the structures here called asci as hemispores or deuteroconidia, representing an intermediate phylogenetic stage between arthrospores and true conidia (limited by Vuillemin to the types produced by phialides). More careful cytologic studies by Moore (1934) have shown that in H. coremiformis the asci are borne in chains at the ends of branches. In the early stage they resemble a chain of arthrospores or conidia on the end of a conidiophore. Then a densely staining structure develops within the cell walls which eventually forms an echinulate spore wall. The ascus wall then degenerates, leaving the aseospore free. The abundant formation of coremia in this species suggests that on equally careful cytologic study, Briosia, a saprophytic genus of the Fungi Imperfecti, might belong here. EREMASCACEAE 183 The species placed in this genus by Castellani belong elsewhere. Ciferri & Redaelli (1934) and Redaelli & Ciferri (1934) have attempted a compara- tive study of several organisms referred to this species, but since none of their organisms seem to agree with the morphology originally described by Vuil- lemin, it is doubtful whether they had the same organism as Vuillemin in his original paper. Among others they had a culture which originally came from Vuillemin, but there were no data to show that it was the strain upon which his original description was based. Their only strain which at all resembled H. stellata was isolated along with Aspergillus and may have also been parasitic on the latter. This strain differed more widely from their other strains than the other strains differed among themselves. If the name H. stellata, should be found to apply to a parasite of Aspergillus sp., then the determination by be found to apply to a parasite of Aspergillus sp., then the determination by cold abscesses, would remain in doubt. Hemispora stellata Vuillemin, Bull. Soc. Myc. France 22: 125-130, PI. 7, 1906. Fig-. 36. — Hemispora stellata. (After Vuillemin 1906.) Sporendonema epizoum Ciferri & Redaelli, Jour. Trop. Med. Hyg. 37: 167-170, 1934; Redaelli & Ciferri, Atti 1st. Bot. R. Univ. Pavia IV, 5: 145-198, 5 figs., 1934, excl. syn., quoad "ceppo Pun." Originally described as a parasite of Aspergillus repeiis forming a hyphal mat on the surface of a jar of preserved pears. Subsequently reported from cases of osteoperiosteitis by Gougerot & Caraven (1909, 1910) and from cold abscesses on penis by Beurmann, Clair & Gougerot (1909). Vuillemin identi- fied the organisms in these cases. More recently Fonseca & Area Leao (1927) report it from a sporotrichoid lesion on the arm. Cultures from lesions Avere pathogenic for rabbits, producing periosteitis. Torula epizoa Corda, in Sturm, Deutschl. Fl. Ill, 3: 97-98, PI. 45, 1829, upon which Ciferri & Redaelli base their specific name, was isolated from tallow, and judging from Corda 's figures is not related to the organism under consideration. I find no mention of the species in Corda 's later publications. Arthrospores in chains up to 30 or more, subspheric, 2.6-3.5yu. with a fuliginous granular wall except on the facet of insertion, occasionally elongate and barrel-shaped (Fig. 36). Hyphae 2-3/t in diameter, irregularly septate. 184 MEDICAL MYCOLOGY Colonies white, 0.5-2.5 mm. in diameter covered with conidiophores mak- ing little brown star-shaped spots. On sugar media, colony blackish brown, at first smooth and mammillate or irregular and coarsely convoluted, becom- ing powdered with ochraceous spores. Aerobic, not liquefying gelatin. Fig. 37. — Hemispora coremiformis. 1, hypha showing septation in lactose broth ; i, S, 5, 31, 37, mycelium showing variation on various media ; i, 17, hemispores ; 6, 7, 12-H, SJf, 33, 35, huge terminal cells (chlamydospores [?]) on various media; 8, young filament; 9^ 10, ampuUi- form cells on potato glucose agar; 11, deuteroconidia ; 15, 16, 18-2S, 25, 26, 32, Si, cells on wort agar, showing secretion of a gel ; 27, 28, terminal hemispores ; 29, ascogenous hypha showing- single echinulate ascospores. SO, Adjoining echinulate ascospores formed from hemispore. 3'/. coremium. 56^ hyphae from coremium. 38, Series showing development of echinulate ascospore and disintegration of the ascus. (1-S3, 35-38 X600 ; Si X500.) (After Moore 1935.) Hemispora coremiformis Moore, Ann. Missouri Bot. Gard. 21: 1934 [case of Rotter & Pena Chavarria, Arch. Schiffs Tropenhyg. 38: 414, 415, Fig. 10, 1934] . EREMASCACEAE 185 Isolated from lesions of the skin following scratching a bee sting with soiled hands in Costa Rica. Surface of the lesion is slightly raised and edges irregular, brass red, tissues infiltrated but not painful to touch. Subsequent lesions developed on edge of ear, angle of the jaw and clavicular lesion. The patient was treated with iodine and tartar emetic intravenously and anti- septics locally, with healing in 2 months. In cultures growth wholly of yeastlike cells and arthrospores for the first half year, then hyphae 2-4/a in diameter developed. Intercalary chlamydo- spores spherical 6-10/a or ellipsoid 7-9 x 12-15 fj. ; conidia 4-6/a in diameter. Asci spherical to clavate, at first terminal, in chains, apparently without sexuality; ascospores brown, echinulate 3-6/* in diameter (Fig. 37). On the more acid agars, growth slow, entirely submersed. On wort agar, colony vermiculate, light cinnamon, cells with sheath. On malt extract agar, colony buff to yellowish, vermiculate at center with radial folds and furrows. On Sabouraud agar, colony cerebriform at center surrounded by a ring of coremia, flatter toward the margin, buff to amber. On potato glucose agar, center acuminate surrounded by a cerebriform area from which radiate many furrows, buff. On nutrient agar, colony flat, center slightly elevated, margin irregular giving a stellate appearance. On glycerol agar, colony similar to that on potato glucose but covered with small coremia. On liquid media, no pellicle, flaky sediment. Sugars not fermented, no acid production, milk coagulated, and gelatin liquefied after 12 days. CHAPTER XI ENDOMYCETALES— EREMASCACEAE IMPERFECTAE The species to be discussed may or may not belong in the Endomycetales, since under certain environmental conditions the vegetative stages of many groups may assume a yeastlike appearance. However, when suitable media are employed, it is probable that ascospores will be formed in many species at pres- ent considered as imperfect. Judging from previous cases, we may anticipate that the ascosporic stage will place them definitely in the Eremascaceae. Since cultural studies seem essential in attempting to differentiate the species of a group where the morphology seems variable, the following pro- cedures, advocated by Redaelli and Ciferri (1929), by Talice (1930), and by Langeron & Talice (1932), may be considered as standard until better are pro- duced. They include most of the good features already advocated by Castel- lani during the previous two decades. The fungus is easiest isolated on carrot agar,* or Sabouraud glucose agar.f Spore formation is sought on Gorodkova agar. Cultures are incubated at 20°, 30°, and 37° C. From information gained from these cultures optimum temperature may be determined more accurately if desired. Comparative cultures may be made on the following media : Raulin, acid, or neutral solution, decoctions of carrot or potato, t malt extract solution (without hops), malt agar (2% agar), 2% glucose agar and corn meal agar (recommended by Smith & Sano, 1933) ; malt gelatin, carrot gelatin; glucose meat broth with 0.5% methylene blue, skimmed milk and peptone sugar broth (formula of the Committee of the Society of American Bacteriologists). Descriptions of the colonies on the above-mentioned media should be recorded after 1, 2, 4, 7, 10, and 30 days and even 60 and 90 days are often useful. A microscopic examination should be made on the second, fourth, tenth, thirtieth, sixtieth, and ninetieth days, on the latter days especial search being made for signs of copulation, and ascospore formation. Material should be examined from the bottom growth and pellicle in liquid media, and from the center and edge of the colony on solid media. The material may be mounted in glycerol, lactophenol, or Lugol's solution (water 11 c.c, potassium iodide 2 gm., iodine crystals 1 gm.). They may be stained Avith ZiehFs carbol- fuchsin, Loeffler's alkaline methylene blue, or Ehrlich's anilin methyl [gentian] violet (methods of the Committee of the Soc. Am. Bact.). The cells should *One kilogram of carrots is washed, triturated, and boiled in 1 liter of water for 3 hours. The solution is strained through cheesecloth, cooled. Altered through paper, made up to volume, and 20 gm. agar are added. tAgar 18 gm.. White's [W^itte?] peptone 30 gm., and glucose 40 gm., water 1 liter. JTalice recommends the following: Reduce 20 gm. of potato to pulp, suspend in 1,000 c.c. water, boil for 15 minutes, filter through cotton, replace water lost by evaporation, distribute to tubes, and sterilize at 120° for 20 minutes. It should be noted that this solution is about 1/10 the concentration usually employed in Thaxter's potato agar. The more concentrated solution is said not to give such good results. 186 EREMASCACEAE IMPERFECTAE 187 never be fixed by heat. A smear may be allowed to evaporate the excess water in air, then it should be fixed rapidly in alcohol and stained. Burri's India ink method and Mitsche & Harrison's collargol methods are useful. Since smears usually separate the cells of the filaments, they should be avoided as far as possible and the morphology compared with that obtained in hanging drops, using either liquid media or media containing 0.1% agar. Langeron & Talice suggest a very thin slant of agar, which is streaked all the way to the glass of the tube. For microscopic observation it is held as desired with two lumps of modeling clay on the stage and observed with an 8 mm. objective and an ocular of high magnification. In giant colonies, one must resort to one of the various devices for growing them, so that they may be uncovered for examination. These often give valuable morphologic details. Microcul- tures from hanging blocks of agar are useful. On liquid media, a little of the bottom deposit is lifted by a wide-mouthed pipette, and floated on a drop of liquid on the slide. The excess liquid is removed by blotting or by evapora- tion. It may be stained by the above-mentioned Lugol or by other suitable stains. Hanging drop cultures of dilute potato decoction should also be made. Here, after development has reached a suitable stage, the organism may be allowed to dry to the cover glass and stained if desired. It may be desirable to remove the lanolin which sealed the cover glass to the ring by wiping first with a dry cloth and then with a cloth moistened with toluene. Pigment formation should be noted. It is quite variable, depending on the composition of the medium, its density, the age of the culture, tempera- ture, light, etc. Fermentation should next be studied. Unfortunately, Castellani has em- phasized this to the exclusion of other characters, while others have failed to confirm his results with many strains, perhaps on account of the method used. Redaelli & Ciferri suggest the following list: arabinose, xylose, rhamnose, glucose, mannose, galactose, fructose, sorbite, dulcite, maltose, lactose, meli- biose, sucrose, trehalose, raffinose, starch, soluble dextrin, glycogen, and inulin. However, see remarks on Monilia Castellani, pp. 63, 64. The use of Lendner's microfermentation method is inadequate, unless all doubtful cases are studied more quantitatively. Experiments should be controlled carefully and repeated three or four times. The constancy of fermentative power has been ques- tioned (Bahr 1915), probably as a result of too great reliance on Lendner's method (see p. 63). While occasional cultures on a sugar which the fungus does not ferment will not alter its ability, repeated subcultures on that sugar may induce an irregular increase of abilitj'- to ferment that sugar Avhich is gradually but irregularly lost when again cultivated on the first medium. Mackie & Chitre (1928), in a study of the intestinal Moniliae of India associ- ated with sprue, show that many strains lose their ability to ferment certain sugars in subcultures on laboratory media and may regain this ability on pas- sage through experimental animals. Ability or nonability to ferment maltose was much more constant than that for any other sugars and may be used as a 188 MEDICAL MYCOLOGY character for the separation of species. The power to ferment other sugars is so variable (in their opinion) that it should not be used to separate species. A study of utilization of carbon sources is helpful, using Raulin's neutral solution, replacing the sucrose successively by glucose, maltose, lactose, man- nose, fi-uctose, inulin, starch and soluble dextrin, methyl alcohol, ethyl alcohol, glycerol, formic acid, acetic acid, oxalic acid, tartaric acid, and citric acid. In the case of the above-mentioned acids, the tartaric acid is replaced by the acid in question rather than the sucrose. Similarly sources of nitrogen are studied, substituting 1% potassium nitrate, potassium nitrite, ammonium car- bonate, urea, glycine, asparagine, and White's [Witte?] peptone for the nitro- gen source of the Raulin's neutral solution. Initial and ultimate hydrogen ion concentration should be recorded in these tests. In the liquid media, these authors suggest cultivation in graduated centrifuge tubes. After notes on the character of growth are taken, these tubes are centrifuged for 5 minutes at 2,000 revolutions per minute and the quantity of fungous cells at the bottom is taken as an approximate indication of the amount of growth. In the case of the sugars it is usually better to sterilize separately and add to the sterile Raulin's solution to avoid possible hydrolysis from the hydrogen ions of this solution. For the effect of different sugars on the morphology, see the inter- esting case of Blast odendrion intermedium (Fig. 38). Production of hydrogen sulphide (Kliger's method), hydrolysis of starch (Committee method), indol production (Ehrlich method modified by Gore) may also be tried, but so far have not yielded much information useful in classification. Finally parasitism and pathogenicity should be studied on experimental animals. Needless to say, this elaborate and ideal method has not been carried out for most organisms so far described in this group. It is often impossible to identify recently studied strains with older species in the literature, owing to the total lack of characters used by one author in the description by an- other. This is especially notable in the case of Castellani, who early aban- doned any mention of morphology and relied wholly on fermentation and enzyme reactions. The validity of fermentation reactions has been much dis- cussed in recent years (for general criticisms see pp. 63, 64), often without much apparent realization of the meaning of results or the limitations of the meth- ods employed (e.g., Castellani 1933). Under Castellani 's influence, very little attention has been paid to morphology until very recently, although we have occasional attempts to correlate morphology and fermentation reactions ; e.g., Fineman (1921) Nye, Zerfas & Cornwell (1928), Mackie & Chitre (1928). Re- cently the pendulum seems to be swinging the other direction, and we have Milochevitch (1929), Talice (1930), Shaw (1931), and Langeron & Talice (1932) emphasizing morphology very strongly, and searching for media which will produce normal mycelium rather than sprout mycelium, and in the case of myself and my students (Rewbridge, Dodge & Ayres 1929, Moore 1933- 1935 and much unpublished data) rather successful search for sexual or per- EREMASCACEAE IMPERFECTAB 189 feet stages has removed several organisms from the imperfect stages and placed them in the Eremascaceae. Since some of these researches have much biologic interest for those attempting to determine morphogenetic factors, they may be summarized in more detail. Marantonio (1893) working with a thrush organism found that sprouting occurred almost exclusively on solid media with occasional hyphae in old cul- Fig-. 38. — Blastodendrion intermedium. Showing the effects of various sugars on the morphology. 1, glucose ; 2, starch ; S, galactose ; Ji, maltose ; 5, lactose ; 6, raflflnose ; T, inulin ; 8, erythritol ; 9, mannitol ; 10, asparagin. (After Ciferri & Ashford 1929.) tures. On liquid media he found mycelium either in the pellicle or in the granular deposit that appeared without turbidity. The lower the pH, the greater the quantity of mycelium found. These observations Avere confirmed and extended to a large number of media by Concetti (1900). 190 MEDICAL MYCOLOGY Fineman (1921) working' with 17 strains isolated from cases of thrush and supposed to be Monilia albicans, finds the fermentation reactions constant. Mycelium develops in liquid media, in complex carbohydrate media, in media under low oxygen tension, and with low surface tension, Avhile the yeast form predominates on solid media, simple carbohydrates, abundant oxygen and high surface tension. Milochevitch (1929, also working with strains isolated from cases of thrush and supposed to be Monilia albicans, reports mycelium formed on media with higher surface tension and that the hydrogen ion concentration does not influence mycelium formation. He used a large number of animal tissues and extracts, and reported good growth on liver agar and blood, kidney and spleen agar, broth, kidney and lung broth. Growth was poor on peptone solution, brain, thyroid agar, urine, thyroid broth, ox gall and Raulin's solution. Talice (1930) undertook an extensive study of the media and conditions favoring the formation of hyphae, using a very wide variety of media and 30 strains of various species of Monilia. On solid media hyphae were produced in the first day or two; the yeast forms predominate afterward, hyphae being formed only in contact with the agar. In species Avhich seldom form hyphae, dextrin peptone media or glucose media give short periods of hyphal production, as also to a less extent do protein media. He found no advantage in semisolid media (0.1% agar) over liquid media in the production of hyphae. Hyphae develop best in liquid media, at least at first. Trying a large number of de- coctions, he concluded that he obtained the best growth with dilute potato decoction. In cultures which have been grown for a long time on solid media, as many as three transplants may be necessary to secure hyphae. Tessier (1890) reported that relatively high acidities favor hyphal production, but Talice states that this varies greatly with the species, probably accounting for some of the conflicting results by earlier workers. Lowered oxygen tension favors hyphal production to a certain point. Higher temperatures, as 37° C, produce the same results. Surface tension is important, as reported by Hahn & Junker and by Milochevitch, but again this varies with the species. The dictum of Roux & Linossier in the case of their strains of Monilia albica^is that complexity of morphologic structure increases with molecular weight of the sub- stances in the culture medium does not hold in this group. Talice regards the yeast form as senescent. Shaw (1931) suggests the morphology on dextrose ag-ar and gelatin stabs (i.e., the diameter of the hyphae, length of cells, size and position of monili- form clusters, and shapes of spores) is important in separating species and species groups. Pijper had previously noted that the creamy or membranous character of the giant colony is correlated with other characters, but Langeron & Talice first emphasized its fundamental importance. The most complete consideration of morphology so far produced is that of Langeron & Talice (1932). They emphasize the distinction between creamy and membranous colonies, the former producing abundant sprout mycelium while the latter do not, although the hyphae easily break apart in plane sec- EREMASCACEAE IMPERPECTAE 191 tions into arthrospores whose ends never become rounded. The creamy cul- tures are moist and shining at least in the first weeks, while the membranous colonies are dry and dull. The characters of these tAvo major groups may be distinguished as follows: CREAMY TYPE Earely folded, only when the growth is very rapid. Consistency of thick paste, easily adhering to the needle but never viscid, easily separating from the substrate. Yellowish or Ijrightly colored. Forming flocculent deposits in potato decoc- tion but no pellicle. Giant colony thick, convex, surface smooth, shining humid, uniform or with slight furrows, center often conic, margins lobulate. MEMBRANOUS TYPE Surface soon folded, soon velvety or studded with coremia. Consistency viscid, not adhering to needle, or if adhering drawing out in a long thread, more adherent to the medium. Dull grayish white. Forming less coherent flocculent deposits on potato decoction and usually a thick highly developed pellicle. Giant colony thick, dull, flat folded, fur- rowed, with coremia, margin not lobed. Two intermediate groups may be characterized in the creamy type. In Mycocandida, the thickness of the colony is variable, surface smooth or curdled, shining, or even iridescent, often transparent when young ; surface may be somewhat folded, j'-ellowish white, growth slower and colony diameter less than in the typical creamy type. In Blast odendrion colony thin and with deep radial furrows with a central eminence, surface smooth and dull. The sprout cell or blastospore is the fundamental element of the creamy group. In general, the shape is characteristic of the genus, but one may often find many variations in a given culture (Fig. 39). They may be characterized as spherical, short ellipsoid, long ellipsoid, ovoid, or long ovoid. (In examina- tion of material one should be sure that the cells have their longest axis ap- proximately at right angles to the line of vision, or a long ellipsoid cell may appear short ellipsoid or even subspheric.) Then we have an asjanmetrical form in Geotrichoides, an intermediate genus with membranous colonies. The pyri- form type (stalagmoide) often suggesting drops or tears is characteristic of Blast odendrion. Of the elongate types, cylindric and clavate are common. Sometimes they are somewhat irregular in development, producing allantoid and other irregular shapes. The various stages in the development of the cell have been clearly de- scribed by Shrewsbury for Hansennla (Willia) , and probably his obseiwations might be extended to the groups covered bj^ Langeron & Talice. The young cell is small, ovoid, spherical, or allantoid with a thin wall and a refractile, homogene- ous cytoplasm (Fig. 40, 1). Sprouting is active, the sprout cells being exactly like miniature mother cells. In each a small refractile corpuscle is visible near the center of the cell. The nature of this body is uncertain, as it could not be identified in fixed preparations and was not stained by vital stains. As growth progresses, the young (adolescent) cell enlarges and the cyto- plasm becomes more granular (Fig. 40, 2). Vacuoles appear, generally only 192 MEDICAL MYCOLOGY one per cell, but more may form in elongate cells. The vacuole soon enlarges to occupy about one-half the cell volume. Small, highly refractile bodies ex- hibiting active Brownian movement appear within the vacuoles. These are probably metachromatic corpuscles. Sprouting is still active and the sprout cells may or may not show vacuoles before separation from the parent cells. These adolescent (mote cells of Shrewsbury) cells correspond to the phase of maximum growth ; thereafter the cells begin to store up glycogen preparatory to production of sexual processes or of hypnospores. A B^ ■J©©©© 13O00f iilfii^wiiBM 10 11 Fig. 39. — Types of blastospores. 1, Structure: A, thick-walled; B, stained for glycogen; C, uniguttulate blastospores ; D, biguttulate blastospores ; E, showing refringent granules ; F, degenerating senescent blastospore. S, Spherical blastospores ; S, ellipsoid, short ; h long ellipsoid ; 5, ovoid types ; 6, asymmetric ; 7, stalagmoid or lacrimiform types ; 8, elongate cylindric types; 9, bacilliform types; 10, irregular bacilliform types; 11, still more irregular types from membranous cultures; 12, truncate types; 13, pyriform types; U, arthrospores. (After Lan- geron & Talice 1932.) As the culture ages, the adult cell (durable cell of Shrewsbury, Fig. 40, 3) appears and may persist unchanged for long periods. It is larger than the preceding types. It contains a large vacuole, usually empty but occasion- ally containing a single fat globule. Fat is stored generally in a single large globule at one of the poles. This globule is usually surrounded by a layer of I EREMASCACEAE IMPERPECTAE 193 protein. Some of these adult cells are transformed into hypnospores. The cell appears dark in color, often slightly larger than the other cells, the wall may not be thickened, but is usually darker in color. The cells often contain fat globules and small dark granules. Finally the degenerate, senescent, or dead cells (shadow cells of Shrews- bury, Fig. 40, 4) appear to be empty of contents, often with numerous fat globules in the vicinity of the ruptured cell. Other cells seem to be filled with fat globules. Perhaps the accumulation of fat reaches a stage where it cannot Fig. 40. -Showing' Shrewsbury's cell types in Hansenula. 1, young cell ; Z, S, adolescent cells ; 4, 5, adult cells ; 6-9, senescent cells ; 10, pseudomycelium. be utilized and causes the rupture of the wall. Occasionally these shadow cells may be artefacts caused by the mechanical rupture of young thin-walled cells which are filled with small fat globules. Sprouting may be from any portion of the cell in the true yeasts, but even among them it is often bipolar as it is in all members of the group under consideration. In very young, thin-walled cells before polarity is well estab- lished, sprouting may occur at other points. In thick-walled cells sprouting is almost always unipolar. In some genera verticils of sprouts may develop 194 MEDICAL MYCOLOGY from one pole, which by proliferation, prodnce dichotomously or polychot- omously branched chains of cells. When the branching is repeated in each cell, we have dense bushy masses forming the arhuscules of Ota. After a period of sprouting, some of the mature thick-walled cells begin to develop mycelium, very much as if a spore had been formed. The cytologic changes accompanying this process are unknown. A definite slender cylindric germ tube develops instead of a subspheric sprout cell. Sometimes the septation of the hypha follows promptly on its formation, at other times the septation lags until the hypha is very long and often multinucleate. The septa may be close or distant. After a time sprouting from the mycelial cells begins, producing the sprout conidia or blastospores. Sometimes these blastospores are borne in verticils, as in Mycotorula, or some of the members may be rudimentaiy and transformed into a hyphal branch, as in Mycocandida. The terminal portion of the hypha furnishes an important character. In Candida, the hyphae, instead of ending in a verticil as in Mycotorula, terminate in a chain of blastospores, which in turn may be branched but never verticillate. In Mycocandida and Blast odendrion each hypha ends in a single cell of variable length. In Mycotorida the hypha ends in a verticil or a dense tuft of blasto- spores. In Mycotoruloides the hyphal termination is a dense compound verticil. Hypnospores are often terminal. Thej^ appear on liquid media and in microcultures beginning to dry up. The contents of one or more cells migrate into the terminal cell where the cytoplasm appears dense and stains deeply with Lugol's solution. The hypnospores always germinate with a germ tube. Coremia are common in the group with membranous colonies (Fig. 3). Here the hyphae are collected into thick flexuous cords which rise perpendicu- lar to the surface and fray out at the top. They appear on all media, even on 2% glucose. Occasionally they are seen in some of the other groups where the blastospores are long and relatively slender, but in this case they are rare on 2% glucose. Besides coremia, on malt gelatin where the colony comes in contact with the glass, one often sees long pointed strands. These are also characteristic of the group with membranous colonies. The classification of this group presents exceedingly difficult problems. The earlier workers had very poor optical equipment and did not grow their organisms in culture. For the most part their descriptions are so brief and vague that it is very difficult to apply any of their names to organisms encountered at the present time. Since the same name early came to be used for entirely unrelated groups of organisms, we often have two or three distinct traditions for the application of a given name, the followers of each tradition claiming all the advantages of priority. To make the confusion worse, many authors have quoted incorrectly or cited dates from secondary sources. Frequently when one attempts to verify an original description, it is so different from that quoted that one can only conclude that the original description was not seen by the modern author. In the following discussions, I have attempted to present the various names in chronologic order, quoting from their original description, and tracing the various applications to various groups. It will thus be seen that practically none of the names published in the eighteenth and nineteenth centuries are legitimately available for members of this group, although many such names are in common use. There are only two alternatives, either we must abandon them altogether as has been done by Langeron & Talice, or else adopt by legislation in our code of nomenclature certain EREMASCACEAE IMPERFECTAE 195 new standard species whicli will conserve a name in one of its traditional uses and rename all the species which do not conform to the tradition selected. By either alternative the outlook is not bright for the medical man. To adopt the first would make a break with the past and involve a renaming of many of the species, and discarding the majority as unidentifiable on account of poor description. Unless this were formally legalized by an International Congress of Botanists, there would always be trouble from the legalist, the historian, and the publicity seeker by their puerile attempts to overturn existing nomenclature in favor of their own interpretation of some older name. If the second alternative is adopted, it must also be secured through the action of an International Congress of Botanists in which each faction would vote for the particular tradi- tion in the application of a name to which they were accustomed, with the deciding vote held by the systematists dealing with flowering plants who v.ould have no interest in, or knowledge of, the matter, and would decide it on national lines. By either horn of the dilemma, action by an International Botanical Congress is neces- sary and one is confronted by the practical problem as to which method to adopt, pending action by such a congress, which is apt to postpone resolutions for a generation; e.g., the action on bacterial nomenclature laid on the table at Brussels in 1910 for action at the next congress has not been acted upon yet. In view of the action of the last congress at Cambridge, England, in 19.30, in adopting the principle of the type species determination of the name, I have attempted to apply this principle strictly, and if the type species belongs in another genus with an older valid name, the genus name to which that type species belongs becomes a synonym of the earlier name. Where no type species can be definitely decided upon, I have adopted the view that it should be applied to the species which would produce the fewer new combinations by such applications. MONILIA Monilia Gmelin, Sy.st. Nat. 2: 1487, 1791. Gmelin segregated as Monilia various species previously placed in Mucor and Aspergillihs, iefining the genus as " Fila moniliformia in capitulum congregata." Most of the species belong to the genera Aspergillus and PenicilUum, although it is almost impossible to identify them with current species. Persoon took up the genus in Neues, Mag. Bot. 1: 121, 1794 (Dispositio 40, 1797) practically repeating Gmelin 's diagnosis but confining it to the erect species. He recognized four species, M. aurea, M. rosea, M. glauca, and M. Candida, M. rosea being described and figured by Batsch, the other three by Micheli. The latter belong in Aspergillus, where Micheli originally placed them. M. rosea Batsch is probably Trichothecium roseum. Consequently, we may eliminate these early uses of Monilia as having no nomencla- torial value, unless preventing a later usage. Persoon, in his Syn. Meth. Fung., divides the genus into three groups of which the first two refer to Aspergillus sp. with radiate heads (with a slight admixture of other things) and those with columnar heads; while the third refers to Torula which he had already defined earlier as a separate genus and which he later regarded as separate. In his Myc. Eur. 1822, Persoon uses Monilia as a synonym of Aspergillus. Link used Monilia in the sense of and instead of Torula. This, however, is untenable, since by none of the rules can Monilia in its original usage include the dark-spored species now referred to Torula and Dematium. Fries, in the Systema, uses Monilia as practically a straight synonym of PenicilUum. Bonorden (Eandbuch 1851) defines Monilia in practically the same terms previously used to define Oidium, treating Monilia Candida from rotten wood as the type of the genus, and describing M. cinerea from rotting cherries as new. Saccardo includes the type of Oidium in his genus Monilia which includes both saprophytes and parasites of plants. Therefore none of the usages of Monilia except that of Bonorden in the first century of its history is ac- ceptable in the modern sense. 196 MEDICAL MYCOLOGY Geiger (1910) considered M. Candida Bonorden as the type of tlie genus and separated Pseudomonilia. Vuillemin (1911) correctly renamed Monilia Candida Bonorden non Pers. as M. Bonordeni Vuillemin, since the former name was preoccupied. He proposed to accept the name Monilia in the sense used by Bonorden, which contains two species which are not generally considered congeneric. Berkhout (1923) proposed to retain Monilia fructigena Pers. (closely related to, if not the same as, Bonorden 's M. cinerea) as the type of Monilia, and described Candida as new, to include the saprophytic and human pathogens, taking as her type Monilia Bonordeni Vuillemin or M. Candida Bonorden. In view of the very complicated and varied usage of Monilia, only two courses are open, to disregard the name altogether, which in many ways would be the simplest, or to fix its usage by adopting it as a nomen conservandum at an international congress in one of the senses as used by later authors. This would probably best be done by fixing Monilia Candida Bonorden as the type species. This species was not only considered by Bonorden as the type of the genus to which he added M. fructigena as a new species, considering his M. Candida the same species as that of the earlier authors (probably incorrectly), but also this procedure would conserve the name for very many species widely used in fermentation and medical literature. Some plant pathologists have attempted to typify Monilia by M. fructigena Bonorden, which belongs to a wholly unrelated group of fungi, in utter disregard of all the fundamental principles of nomenclature and, like the prophets of Baal thinking to be heard by their loud cries, have convinced such well-known mycologists as Langeron & Talice. OIDIUM Oidium, Link, Mag. Ges. Naturf. Freunde Berlin 3: 18, 1809. Link characterizes this genus as " Thallus e floccis caespitosis septatis, ramosis, decvm- bentibus; apicihus articulatis; articulis in sporidia secedentibus. Thallus e floccis complicatis, sporidiis inspersis magnis, ovalibus, ita ut Sporothricho aut Geothricho affine credideris genus. Cum vera accurate inspexeris floccos, invenies apices articulatos, articulosque separari et thallo inspergi. Unico species, colore pulchre aureo, Oidium aureum (Trichoderma aure^im Pers.)." The figure shows a habit very similar to Geotrichum, but the arthrospores are ellipsoid. Persoon, Syn. Meth. Fung, had described the fungus ' ' late effusum, villo subalbido, tenuissimo, pulvere obscure flavo. Provenit rariiis in vaporariis ad ligna cariosa, cui immcrsum." Link in his revision of the fungi in Willdenow's edition of Linne, Species Plantarum 6: 121, 1824, recognizes 10 species, of which 0. virescens and 0. Uredinis are described as new, the rest being transfers from other genera, mostly Acrosporiamn and Monilia. There seems no reason to take Oidium monilioides (Acrosporium Nees) on the under- side of grass leaves as the type of Oidium and consider it a plant parasite, as has been done by Jaczewski and others. Fresenius (1851) introduced confusion by his Oidium lactis. GEOTRICHUM Geotrichum, Link, Mag. Ges. Naturf. Freunde Berlin 3: 17, 18, 1809. This genus was first characterized; "Thallus e floccis caespitosis septatis, ramosis, decumbentibus. Sporidia ovalia, utrinque trurtcata, inspersa. Thallus e floccis complexis. Sporidia magna, extremitatibus truncatis genus designant. Affine genus Sporothrico, at sporidiis sat differt." Geotrichum candidum was the only species recognized: " caespitibus effusis, floccis albis, sporidiis concoloribus. Tenuis instar tomenti terram in sylvaticis sterilibus et ericetis obtegit, maculam albam nudo oculo granulosam efficiens. Plantula fugax." The figures show a septate mycelium with cylindric arthrospores. In 1824, in his revision of the fungi of Willdenow 's edition of Linne 's Species Plantarum, Link includes it in Botrytis under the name Botrytis geotricha. Persoon includes Geotrichum EREMASCACEAE IMPERFECTAE 197 in his Mycologia Europaea, 1822, without comment. Saccardo (1886) recognized the genus, placing the earth-inliabiting species in Eugeotrichum and the coprophilous species in Copro- trichum-. Loubiere (1924) has revived it in connection with his studies of organisms in cheese. MYCODERMA Mycoderma Persoon, Mycologia Europaea 1: 96, 97, 1822. The type species is Mycoderma mesentericum Persoon. The genus was first described as " orMculare, coriiforme, primo molle, suhpellucidum dein induratum, sulstantia uhvque aequali (Aspemvum? natura MuccdinumJ." It included four species evidently forming pellicles on various sugar-containing liquids. M. mesentericum-, from a bottle imp of wine, produced a folded white, viscous pellicle; it had been previously described but not named in Persoon, Traite des Champignons ComestiUes 8, 1818. M. lagenae, also from wine, was smooth, obsoletely rugose below, and reddish. M. ollare on a decoction of Bumex acetosella produced a deeply folded, fuscous to bay, rather fragile pellicle. M. pergameneum produced a thin white pellicle with a rough surface. While I have been unable to locate a copy of the work, Demazieres in his Catalogue des plantes omises dans la hotanographie helgique et les flores du nord de la France, Lille 1823, is said to have renamed M. mesentericum and M. lagenae as M. vini (p. 13) and described M. cerevisine (p. 13). Shortly thereafter he issued his Plantes cryptogames du nord de la France [probably a collection of specimens with descriptive labels, but not seen] in which No. 101 was M. cerevisiae, No. 102 was M. malti-juniperini, described as new, and No. 103 was his M. vini. By 1825 he had prepared cultures of this genus by exposing shallow dishes of beer, wine, etc., to the air and produced several cultures with pellicles. These he studied microscopically and finding flagellates as well as fungus filaments, confused the litera- ture for a long time by trying to fit them into a single life cycle. This was published as his Observations hotaniques et soologiques, Rec. Trav. Soc. Amateurs Sci. Agr. Arts Lille 1825: 1826 [the portion on Mycoderma reprinted in Ann. Sci. Nat. I, 10: 42-67, PI. 3 1827]. In this work he recognizes M. cerevisiae, M. nnalti- cerevisiae, M. malti-juniperini, M. vini, M. glutinis-farinulae, and M. vini Vallot (Bibl. phys. econ. aout 1822). A study of Desmazieres' figures shows that he was not working with pure cultures. M. m-alti-juniperini seems to be made up of filaments dissociating into cylindric cells such as are found in the milk organism, Geotrichum lactis. M. vini is mostly bacterial? with some dichotomous hyphae, rather sparingly septate and sterile; M. glutinis-farinulae is a branched moniliform chain of ellipsoid cells, while M. cerevisiae is made up of cells which may belong to as many as four different organ- isms: Saccharomyces cerevisiae, Geotrichum lactis, a species with septate, dichotomous, sterile hyphae, and branched chains of moniliform cells similar to M. glutinis-farinulae but much smaller. From the preceding discussion it seems clear that Mycoderma should be retained for organisms forming a viscous pellicle on the surface of solutions rich in sugars. During the rest of the nineteenth century various heterogeneous elements were included in this genus. According to Jannin (1913) Vuillemin decided to typify the genus by M. malti-juniperini, being either ignorant of, or ignoring, the earlier description of both Persoon and Desmazieres, and thus fixing the name as a synonym of Oidium lactis Fresenius (since shown to belong in Geotrichum). Most French writers have followed Vuillemin blindly. Enlows (1920) would typify the genus by M. ollare, since it is tlie first on the page on which Mycoderma was described. In the brewing and wine industry, the tradition has been strong to make M. vini and M. cerevisiae the typical species of the group, although they have been differently character- ized by different workers. Unfortunately they have been more interested in the products of fermentation than in the agents involved, so that much important information lies buried in a mass of fermentation studies. Leberle (1909) and Will (1910) illustrate this tradition, and 198 MEDICAL MYCOLOGY define the genus as follows: In young cultures, cells cylindric, ends not rounded; in old cultures, elongate cells, spherical and ellipsoid cells often present in groups; few or no chains of sprout cells; cells oily, with air bubbles. Growth rapid and in a short time covering alcoholic solutions with a thick pellicle, with radial ridges running to within 2 mm. of the periphery; giant colonies yellowish, dull; gelatin not liquefied. SPORENDONEIVEA Sporendonema Desmazieres, Ann. Sci. Nat. 11: 246-249, PI. 21, 1827. This genus is apparently closely related to Geotrichum or Mycoderma, both of which antedate it. It was described from colonies on cheese as follows: Hyphae short, simple or branched, not septate, almost hyaline, grouped, about S/j, in diameter, containing very large reddish spores, often crowded and compressed but in a single line so that the hyphae appear closely septate. Dissemination either from the tips or by the destruction of the thin hyphal walls. The free spores are hyaline. The plate shows short filaments apparently of the Mycoderma type breaking up into arthrospores, but some portions of the filament re- main sterile. Duby, Botanicon Gallicwm [CandoUe, Bot. Gall. ed. 2] 925, 1830, recognizes the species and states that specimens were distributed "crypt, exs. n. 161," but I have not yet been able to locate and study these specimens. Corda, Icones Fungorum 2: 8, 1838, was unable to confirm Desmazieres' observations, but it is not clear whether he actually saw a specimen from Desmazieres or whether he was studying some other common organism on Dutch and Swiss cheeses, as he states that the organism is common on these cheeses. Corda certainly figured some species of Mycoderma, although he referred the organism to Torula. Berkeley & Broome, Ann. Mag. Nat. Hist. II, 5: 460, 1850, renamed the organism as Torula Sporendonema, reporting it from rat dung and stating that it was the same as the organism distributed by Mougeot & Nestler, No. 998. Oudemans, Verslag. Mededeel. K. Akad. Wetens., Afd. Natuur. Ill, 2: (115)-(122), 1 pi., 1886, revived the name for another species which fitted the generic description of Desmazieres, although it does not seem related to Desmazieres ' original species. More recently Ciferri & Kedaelli (1934) and Kedaelli & Ciferri (1934) have used the name for wholly unrelated organisms of which Eemispora stellata (p. 183) and Scopulariopsis D'Agatae (p. 649) are probably human pathogens. OOSPORA Oospora Wallroth, Fl. Cryptog. 1: 182-184, 1833. This genus combines Oidium Link, Oideum Schlechtendal, Acrosporium, Nees, Sprengel, and Persoon, Alysidvum Kunze. It includes also the species on rosaceous fruits now commonly referred to Monilia Berkh. non Bonorden. Both light and dark colored species are included so that Torula and Dematium and perhaps Trichoderma belong here as well. Consequently the genus is made up of such diverse elements that it should be dropped altogether. The use by Saccardo is practically synonymous with the original use of Oidium. The usage of several modern French writers is practically synonymous with Actinomyces. GLYCYPHILA Glycyphila Montagne in Payen, C. R. Acad. Sci. 33: 393-397, 1851. This genus was based on two species, G. erythrospora (Champignon rouge du sucre Mirbel & Payen, Mem. Acad. Sci. 22: 6, PI. 1 bis, 1845) and G. elaeospora. These two species were united under the name G. versicolor by Montagne (Bull. Soc. Nat. et Centr. Agr. 462, 1852) and so treated by him in his Syll. Gen. Sp. PI. Cryptog. 307, 1856. The genus was described EREMASCACEAE IMPERFECTAE 199 as follows: Hypliae araclmoid, hyaline radiating from a common center, very much branched, septate, repent, constricted; branches dichotomous, attenuated, including seriate? spores; spores not easily liberated, spherical, at first rose color then olivaceous, conglomerate, held together by a gelatinous sheath when young. While the relationships of this genus are not clear, I think there is little relationship with Eemispora as suggested by Ciferri & Eedaelli (1934). Only a study of a similar organism from a similar habitat or of Montague's micro- scopic preparations, if they still be in existence, can solve its relationship. SYRINGOSPORA Syringospora Quinquaud, Arch. Physiol. Norm. Path. 1: 290-305, PL S, 1868. The type species is Syringospora Eohini Quinquaud, which is based on Oiditt/m albicans Eobin. If this species is to be removed to a segregate of Monilia, Oidium, etc., this genus name must be used rather than Mycotorula, Parasaccharomyces, etc., as has been done by recent authors. For example, since Langeron & Talice give this species as the type of Mycotorula, Syringospora must be used instead of Mycotorula in their classification. Wliether Mycotorida Will is a synonym, must rest on one's decision as to whether his type species is congeneric with Syringospora albicans (Oidium albicans Eobin). Mycelium septate, dicliotomously or trichotomously branched, the spores borne in dense ■.ufts on very short lateral branches. Hyphae 2-5^ in diameter; blastospores ovoid 3-7/* in diameter, germinating with germ tube or by sprouting. The figure shows dense terminal cluster around a short lateral branch no larger than a spore. ENDOBLASTODERMA Endoblastoderma Fischer & Brebeck, Zur Morph. Biol. Syst. Kahmpilze, etc., 1-52, 2 pis., 1894. Type species not mentioned. Three species, with several varieties each are treated: E. amycoides, E. liquefaciens, and E. glucomyces. E. amycoides var. I. is said to be the same as Mycoderma cerevisiae Hansen, isolated from lager beer. This genus was based upon a misconception that the cells were formed endogenously. From the description it seems likely that the large oil globules of senile cells were observed. These are easily liberated from the mother cell by crushing. The authors emphasize that the genus includes only those species in which the pellicle is promptly and regularly developed. No ascospores formed. There seems little to differentiate this genus from Mycoderma. ZYMONEMA Zymonema Beurmann & Gougerot, Tribune Med. 42: 503, 1909. The type species are Zymonema Gilchristi (Blastomyces dermatitidis Gilchrist) and Z. Sahurani. The authors also intended to include Coccidioides immitis, although they do not mention it by name. Later they state that both Z. Sahurani and C. immitis are aberrant and of doubtful position. Hence, after excluding these two species, we must assume Blastomyces dermatitidis is the type of the genus. Since asci have subsequently been found in that species we must transfer Zymonema to the Eremascaceae (see p. 165). Beurmann & Gougerot characterize Zymonema as follows: Thallus a mixture of spherical or ovoid sprout cells, septate, branched hyphae and short chains of sprouting ovoid blasto- spores. Conidia are catenulate and branched. Arthrospores in chains at the ends of hyphae. No asci observed. 200 MEDICAL MYCOLOGY PARENDOMYCES Parendomyces Queyrat So Laroche, Bull. Mem. Soc. Med. Hop. Paris, III, 28: 111-136, 1909. Type species: Parendomyces albus Queyrat & Laroche. Colony creamy, mycelium scanty, limited to short chains in liquid media, cells ellipsoid; chlamydospores abundant; ascospores not seen. Pellicle formation rare, rings more common on liquid media; gelatin not liquefied; milk coagulated. PARASACCHAROMYCES Parasaccharomyces Beurmann & Gougerot, Tribune Med. 42: 502, 1909. The type species is Parasaccharomyces Samhergeri Beurmann «fc Gougerot based upon Pseudosaccharomyces Bussei Bamberger, Sbornik MinicJcy 5: 466-485, PL 6, 1904. Colony creamy, hyphae straight, long; yeast cells ellipsoid thick- walled ; ascospores not seen. No pellicle but ring with aerial hyphae; gelatin liquefied; glucose fermented. PSEUDOMONILIA Pseiodomonilia Geiger, Centralbl. Bakt. II, 27: 97-149, S pis., 4 figs., 1910. No type mentioned, four species described as new. Since Ps. cartilaginosa and Ps. mesenterica are mentioned as differing in some respects, they may not be considered as type. Consequently we have to choose between Ps. albomarginata and Ps. rubescens, both of which are about equally eligible. Since Ps. rubescens is the only distinctly colored species, it seems wise to consider Ps. albomarginata as the type. Cell shape variable in young cultures, sprout cells in old cultures. More or less branched mycelium without true septa develop from the sprout cells. Giant cells common in old cultures. Strong surface growth, very little deposit. Giant colony similar to MonilM Candida, formation of shaggy tufts. No ascospores. No alcoholic fermentation, sugars variously assimilated. Separated from Monilia which he typifies by M. Candida. ENANTIOTHAMNUS Enantiothamnus Pinoy apud Brault & Masselot, Ann. Derm. Syphiligr. V, 2: 592-602, 1911. The type species is Enantiothamnus Braulti. Pinoy has described and figured this genus very well. It has many of the characters which Langeron & Talice ascribe to Blastodendrion, but it is probably a synonym of syringospora (see p. 277). PROTEOMYCES Proteomyces Moses & Vianna, Mem. Inst. Oswaldo Cruz 5: 192, Pis. 14-18, Fig. 2, 1913. The type species is Proteomyces infestans Moses & Vianna. Yeast cells pyriform, germinating by germ tubes which become septate and produce thick-walled chlamydospores [or arthrospores] . These in turn germinate by hyphae which are highly developed; colonies powdery in the center, furrowed; pellicle formed; gelatin liquefied, milk clotted; no fermentation. EREMASCACEAE IMPERPECTAE 201 MYCOTORULA Mycotorula Will, Centralbl. Bakt. II, 46: 226-28], 1916. The type species is Mycotorula craterica Will. M. radioplicata was also described at the same time and neither designated as the type by Will. Sprout mycelium, unbranched or branched, composed of elongate cells, never forming co- enocytic or septate mycelium. Blastospores spherical or ellipsoid, separating from the parent hypha and reproducing by sprouting, forming short unbranched or branched chains, never crowns. Pellicles are usually promptly formed on liquid media, in which sprout cells usually predominate. Giant colonies flat, smooth, with a wavy, more or less regular margin, usually with a crater formation or shallow radial furrows, often with bundles of hyphae penetrating the substrate. Gelatin rapidly liquefied. Sugars fermented. Organic acids easily assimilated, acids formed in most sugar-containing media. Ethyl alcohol assimilated. No pigment forma- tion, fluorescence, or odor. Hydrogen sulphide produced in mineral nutrient solutions contain- ing powdered sulphur. PSEUDOMYCODERMA Fseudomycoderma Will, Centralbl. Bakt. II, 46: 278-281, 1916. The type species is Pseudomycoderma vini Will. Cells fusiform, with 1-3 oil globules, shorter cells citriform suggesting those of Pseiodo- saccharomyces, also small spherical cells with a single oil globule. Pseudomycelium branched, of long slender cells, terminal portions of the branches not forked. Pellicle developing rapidly on malt extract, at first small islets, resembling a drop of fat, then confluent, but still showing the points of union of the separate islets, the upper portions becoming chalk-white, gas bubbles collecting under the pellicle. In old cultures the pellicle is compact and tough, brownish or slightly reddish. On other liquid media, the islets are rapidly confluent, the pellicle is vigorous, smooth, white and glassy, easily sinking to the bottom. Colony folded, center crateriform, upper portions of the fold chalky. Gelatin slowly liquefied; sugars fermented; hydrogen sulphide produced in mineral nutrient solutions to which powdered sulphur is added. CANDIDA Candida Berkhout, De Scliimmelgeslachten Monilia, Oidium, Oospora en Torula 63, 1923. Geotrichoides Langeron & Talice, Ann. Parasitol. Hum. Comp. 10: 62-69, 1932. This genus was based on the group of animal parasites which had previously been placed in Monilia Bonorden. Its type species was Candida vidgaris Berkhout {Monilia Candida Bonorden and Monilia Bonordeni Vuillemin.) Unfortunately Berkhout, while renaming M. Candida Bonorden to avoid a reduplicating binomial (Candida Candida) in accordance with the international rules, overlooked the fact that M. Bonordeni was a synonym and must be used. The correct name, therefore, should be Candida Bonordeni (Berkhout) Dodge n. comb. Langeron & Talice (1932) commit a grave error, when dismembering this genus as con- ceived by Berkhout, in not retaining C. Bonordeni as the type species, but placing this species in Geotrichoides and making Candida tropicalis (Castellani) Langeron &/ Talice the type of this genus. Consequently the only course open is to reduce their Geotrichoides to synonymy with Candida Berkhout, and to find a new name for the group which they call Candida. Such nomenclatorial changes are very unfortunate and should be avoided. The genus was characterized by small, ovoid, or spherical conidia arising by sprouting from the cells of reduced hj-phae; conidia germinating by sprouting or by germ tubes. Sugars fermented. Mostly pathogenic on man. Langeron & Talice characterize their Geotrichoides as follows: Colonies membranous, thick, radially folded or areolate, with tufts of hyphae giving the colony a dull velvety appear- 202 MEDICAL MYCOLOGY ance; margins not lobulate, blastospores often thick-walled, similar to arthrospores, others ovoid or irregular, sometimes pyriform and suggesting conidia, often very large [7-10^]. Pseudomycelium well developed, hyphae little branched, flexuous, not easily breaking apart, ends of cells flattened, cells filled with fine oil globules. Terminal cell of hypha variable, often a chlamydospore of variable form, rarely a chain of chlamydospores ; coremia abundant; verticils more or less regular, composed of a few pyriform blastospores. Langeron & Talice made Monilia Tcrusei Cast, the type of their genus Geotrichoides, but since they claim that the type culture of Candida vulgaris Berkhout is congeneric with this species, Geotrichoides falls into synonymy with Candida Berkli. excl. syn. MYZELOBLASTANON MyseloUastanon Ota, Derm. Woch. 78: 216-237, 260-264, 1924. No type species was designated in the original description which is very difficult to inter- pret. The author states that he is proposing the genus to cover the blastosporic species of Monilia and immediately proceeds to divide it into three subgenera. He then uses specific names with his subgeneric names as if he considered them genera and Myseloblastanon as a tribal designation, making the combinations Blastodendrion Krausi, B. Arzti, Myzelorrhizoides cutaneum, and M. Gruetsi. When he next treated the genus monographically in his Champ. Paras. Homme (1928), he no longer recognized liis subgenera and combined all his species under the name MyzeloMastanon. Should we consider Myzeloblastanon validly published in 1924? A strict interpretation of the International Rules of Nomenclature would deny such publication, as the specific epithets were not formally combined with the new generic name. The intent of the author is not clear. On the other hand, shall we accept Blastodendrion as validly published in 1924? The combinations of specific with generic names are formally made, but the author distinctly states that he is proposing the name as a subgenus. The intent of the author is equally doubtful. If we decide to drop both names as a permanent source of error in 1924, we are troubled equally. In 1925 Ciferri & Eedaelli formally proposed Blastodendrion as a new genus, making the proper new combinations, but in the same year Ota proposed another species, M. Favrei. This latter species differs from the others in show- ing racquet mycelium suggestive of some species of Zymonema and if it be taken as the type of Ota's Myselohlastanon, would exclude all the other species included in it by Ota and other Jap- anese workers subsequently. In 1928 Ota, in his Champ. Paras. Homme, treats the genus mono- graphically, figuring several of the more important species, including M. Krausi, M. Arzti, M. cutaneum, M. Favrei. A study of all these species leads me to place them in Blastodendrion as used by Langeron & Talice (1932) who first thoroughly studied the morphology of the group, although they proposed B. intermedium as the standard species of their genus Blasto- dendrion. Whether one uses Myzeloblastanon or Blastodendrion for this group of organisms, one will probably find equally careful authors using the other name, and putting forth equally sound arguments for its usage. In view of the fact that the more careful morphologic work has been done on Blastodendrion, while even in his later treatments Ota has still included very diverse elements, in this work I shall use Blastodendrion and consider MyzeloMastanon a synonym. For further discussion of type and description of tlie genus see Blastodendrion. BLASTODENDRION Blastodendrion Ota, Derm. Woch. 78: 216-264, 1924; Ciferri & Eedaelli, Atti 1st. Bot. E. Univ. Pavia HI, 2: 129-146, 1925. The type species is Blastodendrion Krausi Ota. Since a culture of B. Krausi was not available, Langeron & Talice (1932) chose B. intermedium Ciferri & Ashford as the type of the genus. EREMASCACEAE IMPERFECTAE 203 Ota defines Blastodendrion as producing a mycelium of elongate cells with dendroid masses of blastospores (Sprossbciwme of Lindner). Langeron & Talice describe it as follows: Colony creamy, thin, beginning from the germination of a thick-walled blastospore, forming a dendroid mass by sprouting, either bipolar or multiple, rarely cruciate; blastospores polymorphous, the lacrimiform or pyriform type predominating; pseudomycelium more or less developed, little branched, less easily dis- sociable than in related genera, forming dendroid masses with ascending branches parallel or suggesting the branching of sporophores in Penicillium, cells mostly elongate, pyriform, hyphae terminated by a chain of lacrimiform blastospores or by a long slender filament; verticils occasionally present, formed of lacrimiform blastospores. REDAELLIA Bedaellia Ciferri, Arch. Protistenk. 71: 424-428, Fig. 3, 1930; Brunetto, Ciferri & Eedaelli, Atti 1st. Bot. E. Univ. Pavia IV, 5: 125-143, 8 figs, 1934. Type species Bedaellia elegans. Colony growth slow, elevated, small, cerebriform, irregularly convoluted, hyphae hyaline, septate, branched with many fusiform blastospores in tufts at the tips. Blastospores germinat- ing either by sprouting or by hyphae (Fig. 41). SCHIZOBLASTOSPORION Schizohlastosporion Ciferri, Arch. Protistenk. 71: 446-448, Fig. 6, 1930. The type species is Torula A Starkey & Henrici (Schisoblastosporion Starlceyi-Henricii Cif.). Starkey & Henrici described their organism as cells variable, predominantly elongated ellipsoid. Eeproduction intermediate between fission and sprouting, as in Saccharomy codes. Cells with numerous small fat globules which increase in numbers but do not coalesce in old cells. Agar colonies smooth, white. In glucose broth, turbidity and sediment. No fermenta- tion of any sugars. Ciferri adds: slight incomplete ring, margins of colony smooth, blastospores spherical, 5m in diameter, to ellipsoid, 5-7 x 2-3/^; mycelial cells elongate, cylindric, 5.5-7.5 x 10-15/*, rarely to 22ai. MYCOTORULOIDES Mycotoniloides Langeron & Talice, Ann. Parasitol. Hum. Comp. 10: 48-54, 1932. The type species is Mycotoruloides triadis Langeron So Talice. Colonies creamy, thick, convex, beginning Ijy bipolar sprouting of a blastospore followed by progressive branching of the pseudomycelium. Blastospores spherical or ovoid, arranged in verticils, arising at the apical portion of the pseudomycelial cells, no terminal chains of cells. Pseudomycelium formed of short cells, each cell often producing a verticil of blasto- spores at its apex. The verticils are less regularly spaced than in Syringospora and are usually compound, producing an ovoid mass of blastospores whose long axis is more or less perpendicular to the main axis of the pseudomycelium. Some branches develop much more than others, giving an irregular appearance. Occasionally a branch grows out and is terminated by short chains from its terminal verticil. Gelatin not Liquefied. MYCOCANDIDA Mycocandida Langeron & Talice, Ann. Parasitol. Hum. Comp. 10: 56-58, 1932. The type species is Candida mortifera Eedaelli. Colonies creamy, sometimes thick, more often thin, fiat, iridescent, transparent at first, then forming a glacis, sometimes coremia present, beginning by bipolar sprouting, blasto- 204 MEDICAL MYCOLOGY spores appearing late, dimorphous, ovoid or elongate, the latter type predominant, much less numerous than in the preceding genera. Pseudomycelium well developed, higlily branched (suggesting the branching of a fir tree), hyphae ending in a group of blastospores, a short chain, or a single elongate cell. Verticils not developed, the apical portion of a pseudomycelial cell not producing more than two opposite blastospores. Gelatin not liquefied. In this work no attempt will be made to cover the pathologic changes in tissues by these organisms, beyond very brief notes in connection with the in- dividual species. The pathogenicity ranges from great severity with a rela- tively high fatality to very mild lesions. Practically all organs may be at- tacked. The majority of the species so far described are evidently saprophytes which have found a suitable substratum for growth and multiplication. This growth may set up a mechanical irritation, often aggravating already existing conditions, or enzymes and toxins secreted may interfere with the normal functions of the organs. The bulk of the species quickly divides into those attacking one or more of the following organs: respiratory tract, alimentary tract, genitalia, and skin. Some of those with weaker parasitism may be found on more than one organ, while on weakened hosts they may migrate and invade organs not originally attacked. In the respiratory tract, many of the symptoms closely simulate pul- monary tuberculosis, from which they may be differentiated by the absence of Mycohacterium tuherculosis in the sputum and the presence of large numbers of blastospores. A specialized form from the Orient, variously known as "tea factory cough" or "tea taster's cough," is a bronchomycosis, supposed to be spread by the dry tea leaves which are sniffed in determining odor while grading tea. These mycoses usually disappear quite promptly following administration of relatively large doses of potassium iodide. In the digestive tract, we find these organisms in the flora of apparently normal persons. "Whether they are the same species as those which are so abundant in the tracts of diseased individuals, is still a disputed point. Too frequently all these organisms have been called Monilia albicans, Soor, sapinho or muguet without a critical study as to specific identities. ' A few morphologic characters, or few fermentation reactions are given, seldom both, usually neither. The two clinical entities which have gradually been differentiated are thrush and sprue. Both these conditions need much further clinical study correlated with a more careful study of the morphology of the organism. Thrush is predominantly a disease of the mouth of infants, occasionally also of senile or very anemic persons. With weakened resistance the organism may spread to the lower digestive tract, the vagina, or even the moister por- tions of the skin. Apparently several different species may cause approxi- mately the same symptoms, giving rise to no end of confusion in the litera- ture, for each worker apparently searches for a case of thrush, isolates the organism, considers it as typical for the organism originally described by Robin, and designates it as Monilia albicans or by one of the numerous syno- nyms. A study of the drawings in the numerous doctoral theses on this dis- ease in the last century, will indicate many different organisms which we EREMASCACEAE IMPERFECTAE 205 would now place in different genera. On the other hand, some authors are still willing: to assign all sorts of organisms from all types of organs and with varying degrees of pathogenicity to this species, very much as some consider various animal pathogens identical with Saccharomtjces cerevisiae, the common beer yeast ! In sprue, the organism flourishes in the lower digestive tract accompany- ing, if not causing, a severe anemia which usually disappears after the or- ganism no longer is found in the stools. This disease, mostly confined to the white migrants to the tropics, has been studied extensively in the Dutch East Indies, India, and Puerto Rico. Its etiology has been ascribed to many condi- tions, from climate and vitamins to fungi. It is usually more severe in the well-to-do white immigrant or native whose food is often not well adapted to a tropical climate than to the poor native. A prolonged diarrhea is followed by anemia which may prove fatal. Ashford has had success with patients fol- lowing a careful diet over a long period, designed to eliminate the organism from the intestines. Since practically no worker has critically studied the disease in all three regions, it seems quite possible that similar clinical symp- toms have been taken for identities. Attempts to extend the findings of sprue to pernicious anemia, which it resembles in many respects, have not been very successful. Ashford has called the common intestinal organism in Puerto Rico Monilia psilosis and provided it with a diagnosis which makes it include several of the species separated by Castellani on the basis of their fermentation. Others have found variants to which they either have or have not given specific names, until the number of species reported from the intestines is quite large. A third substratum where these organisms are commonly found, includes the genitalia. They are rather rare in the urethra or about the glans, but very common about the vagina. Their action is largely a mechanical irritation re- sulting from small creamy colonies scattered over the mucous membranes. However, they are often difficult to eradicate by application of local anti- septics. They are especially likely to be present in diabetics, owing to the sugar in the urine which encourages their growth. Very rarely they penetrate the bladder and set up a gaseous fermentation, causing pneumaturia. Finally, we have the skin as a possible substrate. Many species cause cutaneous or subcutaneous lesions. There are also several species which have been rather inadequately described which cause lesions closely resembling dysidrosis on the palmar and plantar surfaces and interdigital spaces, rarer in other moist situations. Such lesions are rather more common between the fingers than elsewhere and are locally known as "Jewish washerwomen's disease." The greater frequency among Jews is attributed to the nonuse of soap among these women, since they fear that the animals which the fat is taken from were not slaughtered in accordance with kosher rules. Several species have been reported as attacking the nails in chronic paronychia. Mem- bers of this group have frequently been reported from cases of perleche. Except for the infections of nails, few of the species causing the more super- ficial cutaneous lesions have been adequately described or named. 206 MEDICAL MYCOLOGY Undoubtedly many of the species so far described are primarily sapro- phytes and are present in the lesions as secondary invaders. This may ac- count for the relatively large numbers of organisms which have not been reported more than once. On the other hand, there seem to be rapidly ac- cumulating indications that some of the organisms of this group are primarily parasitic, or at least so profoundly modify other factors in the production of disease that they should be considered seriously in the study of the disease. It is to be hoped that with clearer ideas of morphology and physiology of these organisms and a recognition of the ability of many different species to occur in essentially the same clinical entity, we may have tools which will aid us in a more accurate delimitation of species. No really stable clinical differentia- tions, sound animal experimentation, or rational therapeusis can be built up until we have broader and more thorough studies of the many organisms in- volved. In the folloAving systematic discussion, I have attempted to refer organisms to the various genera on the basis of their published descriptions, realizing thoroughly that only a portion of the characters are given, I have also very largely refrained from reducing species to synonymy, since I feel that the burden of proof rests with him who would reduce species rather than with the describer of new species. It is, therefore, quite probable that several species here recognized will eventually be shown to be synonymous with other species, but I feel that at the present time there are few studies sufficiently extensive and thorough upon which to base such action. Key to Genera of Eremascaceae Imperfecta^ Mycelium breaking up into arthrospores, i.e., cells of the filaments cyclindrie or nearly so after separation. Colony cerebriform, villous, irregular, growth slow, little or no growth on liquid media, arthrospores not very abundant. Proteomyces. Colony membranous, folded, dull, usually grayish white. True mycelium not fragile before disarticulation of arthrospores, no blastospores ; thick pellicle on liquid media, sugars not fermented. Gelatin liquefied. Geotrichum. Gelatin not liquefied. Mycoderma. Pseudomycelium fragile, conidia and transitional forms between blastospores and arthrospores present. Gelatin not liquefied, no pellicle on liquid media, although occasionally a partial ring formed. Blastospores verticillate, sugars fermented. Candida (Geotrichaides). Blastospores single, sugars not fermented. Schizoblastosporion. Gelatin liquefied, chalky- white to yellowish pellicle on liquid media, cells long fusiform or apiculate, sugars fermented. Pseudomycoderma. Arthrospores not produced, blastospores abundant and characteristic, colony creamy, yellowish white, shining. Pseudomycelium little branched, blastospores not in verticils nor in regular pairs at the ends of pseudomycelial cells; cells ellipsoid. Gelatin not liquefied, no pellicle, although rings are sometimes produced. Sugars not fermented. Parendomyces. Sugars fermented. Castellania. EREMASCACEAE IMPERPECTAE 207 Gelatin liquefied, sugars fermented. No pellicle. Parasaccharomyces. Pellicle produced. Mycotorula. Pseudomycelium branclied, blastospores in chains, verticils or pairs at the ends of pseudo- mycelial cells. Blastospores apical only. Blastospores isolated or in apical verticils, fusiform, colony cerebriform; sugars not fermented. Redaellia. Blastospores in long chains, rarely ever verticillate, ellipsoid; gelatin not liquefied. Monilia. Blastospores both terminal and lateral, never in long terminal chains, sugars fer- mented. Verticils simple, regular, often in terminal tufts, gelatin liquefied, very thin pellicle on malt extract or none. Syringospora. Verticils either simple or compound, more or less regular, not terminal; gelatin not liquefied. Blastospores lacrimiform or pyriform, mostly in dendroid clusters, colony dull. Blastodendrion. Blastospores spherical or ovoid, verticils compound, elongate. Mycotoruloides. Blastospores elongate-ellipsoid, verticils rudim.entary, reduced to two blasto- spores each, colony folded. Colony moist, shining. Mycocandida. Colony dull, pellicle present, no sediment, no fermentation. PseudomonUia. Artificial Key Based Largely on Biochemical Characters Colony cerebriform, sugars not fermented. Arthrospores produced rarely, gelatin liquefied. Proteomyces, Blastospores fusiform, terminal verticils. Eedaellia. Colony membranous, wrinkled or much folded, grayish white, arthrospores present. Gelatin liquefied. Sugars fermented, pellicle produced. Pseudomy coder ma. Sugars not fermented. Geotrichum. Gelatin not liquefied. Sugars fermented. Candida. Sugars not fermented. Pellicle well developed, thick, on liquid media; no blastospores present. Mycoderma. No pellicle, fragile ring occasionally; blastospores present. Schisoilastosporion. Colony creamy to viscous, smooth, shining, yellowish white, arthrospores rare or absent. Gelatin liquefied. Sugars fermented. Pseudomycelium little branched, blastospore not in dense verticils. No pellicle. Parasaccharomyces. Pellicle produced. Mycotorula. Pseudomycelium branched. Pellicle very thin or none, blastospores in dense verticils. Syringospora. Pellicle well developed, blastospores in terminal chains. Monilia. 208 MEDICAL MYCOLOGY Gelatin not liquefied. Sugars not fermented. Pseudomycelium little branched. Parendomyces. Pseudomycelium branched, colony dull, pellicle present. Pseudomonilia. Sugars fermented. Morphology unknown, no pellicle on liquid media. Castellania. Blastospores pointed at one end, in dendroid clusters. Blastodendrion. Blastospores spherical or ovoid, verticils compound elongate. Mycotoruloides. Blastospores elongate-ellipsoid, verticils rudimentary, reduced to 2 spores each, colony folded. Mycocandida. PROTEOMYCES Proteomyces Moses & Vianna, Mem. Inst. Oswaldo Cruz 5: 192, Pis. 14-18, Fig. 2, 1913. The type species is Proteomyces infestans Moses & Vianna. Yeast cells pyriform, germinating by germ tubes which become septate and produce thick-walled chlamydospores or arthrospores. These in turn ger- minate by highly developed hyphae ; colonies cerebriform, vermiculate or fur- rowed, often appearing powdery in the center ; pellicle formed on liquid media (rarely only a highly developed ring) ; no fermentation. Isolated from abscesses and ulcers in man, the lesions are usually some- what superficial but spread rapidly and widely. In the absence of sufficient morphologic data, I have assembled here those species with cerebriform or vermiculate colonies, showing no fermentation of sugar, whether gelatin is liquefied or not. It seems quite probable that further work will show that these species belong in Zymonema or in some closely related genus. Several of these species were referred to Candida (Geotrichoides) by Langeron & Talice (1932), but they seem better placed here until more is known of their mor- phology. Key to Species Colony rose, P. muris. Colony rose (reddish orange on potato). P. Brocianum. Colony yellowish, becoming brown, remaining white on some media [see also P. asteroides]. Milk coagulated. P. infestans. Milk not coagulated. P. Brocquii. Colony yellowish. Glucose fermented, cells 4-8ai, chlamydospores terminal, 15-18/x. P. cutaneum. Glucose and lactose fermented, chlamydospores up to 30m- P- Faverae. Sugars not fermented. Cells 4-5/01 in diameter, chlamydospores Sfi. P. Balzeri. Cells 2-3 X 1, rarely up to Afi, ellipsoid, chlamydospores rare. P. Perieri. Cells 4-6/u, chlamydospores intercalary. P. Grieivanki. Cells 1.8 X Pj.Gfj., chlamydospores terminal. P. carnealis Proteomyces muris (Ciferri & Kedaelli) Dodge n. comb. Mycotorula muris Ciferri & Redaelli, Atti 1st. Bot. R. Univ. Pavia III, 2: 243-245, 280, 1925. EREMASCACEAE IMPERFECTAB 209 Isolated from spontaneous blastomycosis of a white rat. Cells ovoid, guttulate, or ellipsoid, but often variously shaped, 4.5 x 5/a or 5 X S/A up to 4.5 X 8/A, in malt extract forming a septate mycelium, 2-2.5 x 12.2/A. No ascospores. Giant colonies in must gelatin round, verrucose, rose. No fermentation, no coagulation of milk. The strain referred by Lodder (1934) to Bhodotorula pallida Lodder was evidently a contaminant. Proteomyces brocianum (Pinoy) Dodge, n. comb. Mycoderma brocianum Pinoy apud Anderson, Colas-Belcour & Broc, Arch. Inst. Pasteur Tunis 19: 317-322, 1930. Isolated from ulcerous lesions on the back of the hand. Not pathogenic to experimental animals. Hyphae septate ; arthrospores 4.5 x 3.5//,, thin-walled, no conidia. Colonies on Sabouraud glucose agar at 37° C. creamy white with fine filamentous radiations on the surface with rose color visible on the fifth day and more marked the fourteenth day, becoming somewhat cerebriform in age. Fig. 41. — Proteomyces infestans. (After Moses & Vianna 1913.) Colonies on gelatin similar. On potato, similar but reddish orange, only slightly folded at the center. On carrot, color remains rose. In potato de- coction, sediment, no pellicle, liquid clear. Acid formation with maltose, sucrose, and glucose, none with lactose. Gelatin not liquefied. Proteomyces infestans Moses & Vianna, Mem. Inst. Oswaldo Cruz 5: 192, Pis. 14-18, Fig. 2, 1913. Sporotrichum infestans ? Sartoiy, Champ. Parasit. Homme Anim. 655-656, 1922. Mycoderma infestans Fonseca & Area Leao, Brasil Med. 43: 667, 1929. Isolated from hard, deep-seated abscesses on the extremities Avhich become edematous, and the temperature rises to 39°-40° C. Pathogenic for guinea pigs and rats. Mycelium septate, cells elongate, not easily separating into arthrospores. Blast ospores pyriform, lateral, borne singly, chlamydospores terminal (Fig. 41). Growth on Sabouraud maltose agar good, bright brown, cerebriform sur- face in 3-4 days, later the periphery with radial folds. On Sabouraud glucose, colony cream-colored, elevated, cerebriform. On Sabouraud conser^'ation agar, 210 MEDICAL MYCOLOGY colony smooth and flat with slightly dentate margin. On beet and potato, colony cream-colored with broad convolutions, quickly becoming white and powdery. In agar and Loeffler agar, colony cream-colored with a tough, wrinkled surface. In glycerol solution, both pellicle and sediment formed, the liquid remaining clear. On maltose broth, a thick pellicle with broad con- volutions appears. Gelatin liquefied and milk coagulated. No indol. Survives exposure to a temperature of 80° C, for one-half hour but is killed after one hour at this temperature. Proteomyces Brocquii (Beintema) Dodge, n. comb. Parendomyces Brocquii Beintema, Ann. Derm. Syphiligr. VII, 4: 399-423, 14 figs., 1933. GeotricJium Brocqi Castellani & Jacono, Jour. Trop. Med. Hyg. 36: 317-318, Figs. 48-50, 1933. Isolated from the blood of a patient with papules and pustules forming extensive confluent ulcerated areas, originally described clinically as ''pseudo- bromuride" by Brocq, Pautrier and Fernet. Probably the lungs were also involved, as this organism was regularly found in the sputum. Healed promptly on treatment with potassium iodide. Cells cylindric, 10-15 x 4-5/x, hyphae little branched, forming arthrospores. Chlamydospores ovoid, fusiform or pyriform, walls thick, yellowish. On Sabouraud agar, colony at first a viscous mass, then firmer, shining, reticulate wrinkled to vermiculate, at first white, then cream-colored, yellow, brown or chocolate. On Sabouraud conservation agar, growth very slow, colo- nies not over 3 cm. On potato, colony brown, soft. On carrot, growth better, lighter, drier, with more aerial mycelium. On serum, colonies white, flat, poor gi'owth, no change in medium. On blood agar, growth slow. No growth on ascitic fluid. Growth slight with slight acidity on litmus milk. On peptone maltose broth and other liquids, no pellicle but the ring is highly developed. Sugars not fermented. Gelatin slowly liquefied after some time. It is possible that this species is synonymous with the following which antedates it. However, P. asteroides has been so briefly described that I dislike to reduce P. Brocquii to synonymy without further study. Proteomyces asteroides (Rischin) Dodge, n, comb. Parendomyces asteroides Rischin, Arch. Derm. Syphilis 134: 232-242, 9 figs., 1921, Trichosporium asteroides Bolognesi & Chiurco, Micosi Chirurg. 597, 1927 ; Ota, Jap. Jour. Derm. Urol. 28: [3], 1928. Geotrichoides asteroides Langeron & Talice, Ann. Parasitol. Hum. Comp. 10: 68, 1932. Lesions (probably contracted from a calf) appearing on beard and neck of man. Lesions infiltrated, elevated 1-1.5 mm., round or oval, confluent, swollen, soft, exuding brownish yellow secretion. Pathogenic to guinea pigs, rats, and mice. EREMASCACEAE IMPERFECTAE 211 Colony dull white, circular or oval, with sharp contours, becoming radially and spirally folded, turning yellowish and finally brownish. Growth best on sugar, potato, and gelatin media, poor on serum and ordinary agar. No fer- mentation. Yeast cells gram-positive (Fig. 42). Near P. Balzeri, considered by Ota, Ann. Parasitol Hum. Comp. (1926) as close to P. cutcmeus, later considered distinct. Proteomyces cutaneus (Beurmann, Gougerot & Vaucher) Dodge, n. comb. Oidium cutaneiim Beui-mann, Gougerot & Vaucher, Bull. Mem. Soc. Med. Hop. Paris 28: 256-259, Figs. 54-93, 1909; Tribune Med. 42: 505, Figs. 54-93, 1909 ; Les mycoses nouvelles 52, Fig. 4, 1909. Monilia cutanea Castellani & Chalmers, Man. Trop. Med. ed 2, 830, 1913. Trichosporum cutaneum Ota, Ann. Parasitol. Hum. Comp. 4: 2-13, 1926. Fig-. 42. — Proteomyces asteroides. (After Ota 1926.) Mycoderma cutaneum Brumpt, Precis Parasitol. ed. 4, 1212, 1927. Geotrichoides cutaneus Langeron & Talice, Ann. Parasitol Hum. Comp. 10: 67, 1932. Geotrichum cutaneum Almeida, Annaes Fac. Med. Sao Paulo 9: 12, 1933. Produced hypodermal and dermal gummatous infiltrations, later becom- ing ulcerated. On glycerol agar are formed yeast cells, spherical or ovoid, 4-8/x, elongate forms 3-4 x 20/t, sometimes agglomerating. Cultures smooth, shining, viscous, growth good (Fig. 43). On peptone glucose agar, these yeast cells give rise to pseudofilamentous cells 4-5 X 8-10/x and filamentous forms of cylindric cells with terminal chlamy- dospores 8-12^ in diameter. In a purely filamentous stage, cells are 2-4 x 20/i, 212 MEDICAL MYCOLOGY conidiopliores erect, simple or double, forming chains of cylindric spores 4 x 8/1.. Chlamydospores terminal, 15-18/a. Colonies light yellow, reticulate, crumpled with aureole. No trace of ascospores, even in old cultures. Colonies on agar or potato light yellow, crumpled, mammillate, aureoled, powdery, and whitening in age. Yeast stage ferments glucose, filamentous stage does not. In animal (pus, etc.), yeast form only appears. Repeated subcultures lose virulence. Froteomyces Balzeri (Gougerot & Burnier) Dodge, n. comb. Parendomyces Balzeri Gougerot & Burnier in Balzer, Gougerot & Burnier, Ann. Derm. Syphiligr. V, 3: 282-295, 4 figs., 1912. Saccharomyces Balzeri Castellani & Chalmers, Man. Trop. Med. ed 3, 982, 1919. Fig. 43. — Proteomyces cutaneus. (After Ota 1926.) Monilia Balzeri Brumpt, Precis Parasitol. ed. 3, 1922 (cited by Vuillemin, Champ. Parasit. Homme 86, 1931) ; Sartory, Champ. Parasit. Homme Anim. 708, 1923. Trichosporum Balzeri Bolognesi & Chiurco, Micosi Chirurg. 597, 1927. Trichosporon Balzeri Ota, Jap. Jour. Derm. Urol. 28: [4], 1928. Geotrichoides Balzeri Langeron & Talice, Ann. Parasitol. Hum. Comp. 10: 68, 1932. Candida Balzeri Almeida, Annaes Fac. Med. Sao Paulo 9: 11, 1933. Is.olated from hypodermic gummata in crural region. Cured by medica- tion with KI. Pathogenic to rabbit and guinea pig. Reported by Motta from mycosis of the pharynx. Yeast cells spherical, 4-5(-6)/x. in diameter, granular, vacuolate, thick-walled, reproduce by sprouting. Sprouting pseudomycelial forms 3 x 10-16/i,. Hyphae EREMASCACEAE IMPERPECTAE 213 in old cultures occasionally up to SAfi in diameter, cells mostly 2-2.5 x 14-28/t. Chlamydospores 8/t in diameter, yellowish. No asci found (Fig. 44). On peptone glucose agar, growth whitish yellow, becoming more or less dark, elevated, cerebriform. Moist at first, then waxy and dull. Gelatin not liquefied; sugars not fermented [Ota]. Proteomyces Perieri (Matruchot & Antoine) Dodge, n. comb. Oospora Perieri Matruchot & Antoine, Soc. Path. Comp. Nov. 13, 1917; Ann. Inst. Pasteur 32: 202-214, 1918. Monilia Perieri Castellani & Chalmers, Man. Trop. Med. ed. 3, 1092, 1919. Trichosporon Perieri Ota, Jap. Jour. Derm. Urol. 28: [4], 1928. Candida Perieri Almeida, Annaes Fac. Med. Sao Paulo 9: 11, 1933. Found in war wounds in France. Pathogenic to rabbit and guinea pig. Fig. 44. — Proteomyces Balzeri. (After Ota 1926.) On beet, pseudomycelial cells, elongate, 1-2 x 6-7/x, blastospores 2-3 x 1/x.. On Sabouraud agar, yeast cells predominate, 1-2/a sometimes up to 4 or 5/n in diameter. Division either by sprouting or by equal fission. Oil droplets abun- dant in old cells. On vegetable decoctions with sugar, mycelial forms pre- dominate. In the deposit are small white masses or spheres of mycelium which are sterile or show rare chlamydospores. On coagulated serum hj^phae l-3yu. in diameter, blastospores 1-2/x. Optimum temperature for growth 15-18° C, but growth still good at 37° and slight at 45° C. Antiseptics rich in oxygen (e.g., KMn04, H0O2) do not check growth but produce gigantism (cells of double size). On carrot, glycerol, beet, or gelatin at 16-18°, colonies begin as small, rounded excrescences with small crater, opaline. Later colonies confluent, folded, vermiculate, and finally cerebriform. On carrot it is dry or waxy. 214 MEDICAL MYCOLOGY Color seems to vaiy with medium, being yellow ochraceous on carrot, rose on beet, and whitish, turning ochraceous, on gelatin. On coagulated serum a thick white colony forms. In vegetable decoctions with sugar, a whitish pellicle and a white sediment form. Acid media less favorable than neutral or slightly alkaline. Gelatin not liquefied. Proteomyces Griewanki (Neveu-Lemaire) Dodge, n. comb. Mycoderma sp. (sensu Verdun) Griewanki & Laveau, Bull. Soc. Path. Exot. 12: 478-482, 1919. Mycoderma Griewanki Neveu-Lemaire, Precis Parasitol. 70, 1921. First lesion appeared as tumor at base of big toe. In the course of five years extended dorsally and internally in foot, which finally became globose and like a bear's paw. Ulcerous, but was painful only on pressure, not sensi- tive to pricking. Finally the foot was amputated. A study of the tissues dis- closed red grains. Yeast cells 4-6/a in diameter, end to end, yellow brownish with numerous rounded, cocciform spores in chains or clusters. Filaments long, flexuous, sep- tate, white, refractive, 2-2.5//, x 15/a, sometimes piled up, curved, or branched. Blastospores rounded, arranged in lines. Chlamydospores usually intercalary, rarely terminal. Arthrospores cut off squarely. Staining by Ziehl method good. By Gram method, walls stained, some cells staining deeply. Proto- plasm granular, chlamydospores, arthrospores and budding elements rose. Some arranged in spirals and helices. Cultivated on straw infusion (1.5% agar), on banana, potato + glycerol, and Sabouraud maltose agar, the latter one being unsuccessful. On the former in six days appear small cerebroid masses, gelatinous, yellowish, becoming size of small pea. Finally a rose-colored efflorescence over surface of media. Proteomyces Faverae Dodge, n. sp. Oidium sp. Favera, Giorn. Ital. Mai. Ven. Pelle 55: 650-729, 10 pis., 1914. On man produces cutaneous ulcers similar to those of Z. dermatitidis. Path- ogenic to laboratory animals. In 5-day-old cultures, the cells are spherical, 2-30/* in diameter, thick -walled, sprouting. Hyphae slender, straight or slightly curved, homogeneous, usually 1-2 sprouting from each sprout cell; septa not visible, terminal cell swollen. After 15 days, hyphae are septate, branched, unequal ; yeast cells elongate in chains of 5-8 cells. Four-to-six-week-old cultures show only septate hyphae of long cylindric cells and chlamydospores ; chains of ovoid spores 2-3 cells long on lateral branches. Optimum growth at 34°-38° C, growth very slow at 18°-20° C. On Sabouraud glucose agar, colony is thick, yellowish, white, humid, creamy, margin festooned, surface irregular, rugose becoming cerebriform. After 2 weeks, shining white mycelium develops which rapidly covers the yeast colony and spreads beyond it, suggesting the pleomorphism of the Trichophytoneae. The colony becomes folded and furrowed with the bottom of the furrows moist and yollowish. On Sabouraud maltose, colony is similar, but with a slight rose tint. On common agar, thick, whitish, moist and rugose, mycelium very slow EREMASCACEAE IMPERFECTAE 215 in appearing. Similar on glycerol agar, serum agar, and coagulated serum. On potato the colony is thick, moist, yellowish, tending to grayish, sur- face convoluted, margin elevated, mycelium appearing after a month. On beets, colony reddish but otherwise similar to that on potato. On gelatin, colony elevated, hemispheric, smooth, later becoming Avrinkled. On beef serum, abundant whitish, floccose deposit. On glucose, maltose, or glycerol broth, white pellicle appears with floccose white sediment, liquid remains clear. Glucose and lactose slowly fermented ; gelatin not liquefied ; mik coagu- lated in 9-10 days. Proteomyces cornealis (Nannizzi) Dodge, n. comb. Monilia cornealis Nannizzi in Bencini & Federici, Atti R. Accad. Fisiocrit. Siena X, 3: 748-766, 8 figs., 1928; Ibid. 4: 92-94, 1929. Isolated from corneal lesions. Pathogenic for experimental animals. Cells 3.6 X 1.8/x or spherical 3.5-4/x, in diameter; hyphae septate, cells S/x x 2/* with chains terminal on lateral branches of blastospores. Hyphae of vari- able diameter 1.5-2.6/*, chlamydospores thick-walled, terminal. On Pollacci agar, colonies irregular, margin erose, denticulate, smooth at first, later velvety, center folded and cerebriform vermiculate. On carrot agar, colony smooth, mammillate, folded, dirty white, moist, margin smooth. On Sabouraud agar, growth slow, irregular, small whitish colonies. On po- tato, colonies caf e-au-lait, diffuse, moist, plane then folded ; on carrot, similar but yellowish. On coagulated egg albumen, colony yellowish white, medium partially liquefied; on coagulated serum, colony grayish, humid. On glucose broth, a slender ring and white sediment. Acid in glucose, sucrose, and salicin broths, slight acidity in fructose, lactose, mannite, dextrin, and galactose. Gelatin liquefied, milk and serum coagulated, methylene blue not reduced in glucose broth. Proteomyces goensis Dodge n. sp, Monilia sp. Froilano de Mello, Antiseptic 26: 427-432, 2 pis., 1929; Bull. Soc. Path. Exot. 22: 142-147, 2 figs., 1929. Isolated in Nova Goa from multiple verrucose dermatitis with lesions sug- gesting those caused by Zymonema dermatitidis. Pathogenic to rabbits and white rats. Cells up to 8/x in diameter, mycelium giving rise to groups of blastospores (or conidia). On Sabouraud 's maltose or glucose agar, colony whitish. On simple agar, and Gorodkova agar colony yellowish. On potato glycerol, colony dry, waxy, white. On broth, slight pellicle, turbidity, and white sediment. Milk coagu- lated and slowly digested. No fermentation of usual sugars, acid with maltose. GEOTRICHUM Geotrichum Link, Mag. Ges. Naturf. Freunde Berlin 3: 17, 18, 1809. The type species is Geotrichum candidum Link. Colonies membranous, adherent to the substratum, dull, velvety due to abundant coremia, margins uneven, forming a pellicle on liquid media, gelatin 216 MEDICAL MY(/0LOGY liquefied, and sugars not fermented. Often a filament grows some distance from its colony and by repeated branching starts a new colony. Mycelium al- ways growing by septation, never by sprouting; germination either unipolar or bipolar ; the cells may become very long and even slightly branched before septation occurs. As the hypha matures the walls and septa become thickened, and the cells finally separate, forming arthrospores. Since thickening of the wall occurs before disarticulation, the spores remain strictly cylindric, the ends not round, while in some species the wall undergoes a gelification, and Fig. 45. — Geotrichuni. 1, germination of arthrospores ; 2, fragments of filaments ; S, formation of arthrospores ; i, chains of young arthrospores ; 5, young arthrospores showing cylindric appearance ; 6, mature arthrospores with rounded ends ; 7j chains of arthrospores. (After Langeron & Talice 1932.) the arthrospores tend to become ellipsoid. In the latter case, it is often diffi- cult to separate species from Monilia (sensu strictiore) or Candida Berkh. excl. syn. Chlamydospores have also been reported (Fig. 45). In Geotrichum versiforme, the only species whose cytology has been care- fully investigated (Moore 1934), the young germ tube arising from the uni- nucleate arthrospore is 2-3-nucleate, rarely up to 8-nucleate. As the hypha EREMASCACEAE IMPERFECTAE 217 elongates, septum foiination proceeds rapidly until the resulting cells, even- tually arthrospores, are uninucleate. In this process often the protoplasmic contents of certain cells disappear, perhaps aiding in the disintegration of the chains of arthrospores. Chlamydospores large and thick-walled spores are multinucleate. On certain media some blastospores are produced having 1-3-nuelei, often uninucleate. The so-called conidia of this species seem to be cells densely filled with a granular protoplasm and no trace of a nucleus. Obviously they are not conidia in the sense in which this term is usually used in other groups of fungi. The genus is quite large, the species being mostly saprophytic on earth and decaying organic matter. Twelve have been reported on man, eight from infections of the respiratory tract, three from stools (parasitism unknown) and two from blastomycosis. O. hostonense differs in some respects from the other members of the genus and perhaps should be placed in a separate genus. Key to Species Colony red, milk coagulated. G. pulmonalis. Colony amber yellow to light yellow. Milk not coagulated. Acid in lactose. G. asteroides. No acid in lactose. G. famatum. Action on lactose not reported. G. rugosum. Milk coagulated. G. immite. Isolated from case of enteritis. G. rotundatum. Isolated from case of bronchitis. G. Muisa. Colony cream-buff to grayish white. No acid on sugars, hyphae 2-4/,i. G. louisianoideum. Acid on fructose, maltose, and galactose. G. versiforme. Colony greenish. G. hostonense. Geotrichum pulmoneum (Bennett) Basgal, Contr. Estudo Blastomycoses pulmonares 49, 1931; Langeron & Talice, Ann. Parasitol. Hum. Comp. 10: 74, 1932. Oidium pulmoneum Bennett, Trans. R. Soc. Edinburgh 15: 2:277-294, 1842. Oospora pulmonea Saccardo, Syll. Fung. 4: 16, 1886. Mycoderma pidmoneum Vuillemin, 1891; Brumpt, Precis Parasitol. ed. 4, 1214, 1927. Monilia pulmonea Castellani & Chalmers, Man. Trop. Med. ed.- 2, 829-830, 1913. Pound in sputum at autopsy of case of pseudotuberculosis. Hyphae septate, dichotomous, 5-10/x. in diameter, spores ovoid, smooth, 10-14/A, borne in chains [Robin's (1853) figures almost suggest Scopulariopsis]. Geotrichum pulmonale (Ciferri & Redaelli) Dodge, n. comb. MycotoriUa pidmonalis Ciferri & Redaelli, Atti 1st. Bot. R. Univ. Pavia III, 2: 205-212, 281-282, 1925. Ehodotorida mucilaginosa var. sanguinea Lodder, Anaskosporogenen Hefen 1: 98-99, 1934. 218 MEDICAL MYCOLOGY Isolated from sputum of a case of pulmonary abcesses. Pathogenic for guinea pigs and rats. Cells rounded, slightly ovoid, often cylindric and catenulate, forming mycelium, hyaline or guttulate, 3-4//. in diameter or 2 x 5-ll^t. No ascospores. Colonies on carrot agar, shining, peach-blossom red, finally cinnabar red, pellucid, slightly verrucose with smooth margin which finally becomes sinuous. On malt agar, colonies red with a suggestion of orange, moist, smooth, slimy. On gelatin, colony small, thick, smooth with ragged margin, red. On malt extract, floating islets and carmine red ring, with thick rose to red sediment and turbid liquid. No fermentation, milk curdled, gelatin slowly liquefying after 100 days. The relationship of this species is not altogether clear. It seems to be rather aberrant in Geotrichiim and perhaps belongs in Torulopsis, but accord- ing to descriptions, it has too much mycelium for that genus. Unfortunately, Lodder did not choose her media to encourage the formation of mycelium, and she examined her cultures altogether too soon to observe it if it did occur. Geotrichum asteroides (Castellani) Basgal, Contr. Estudo Blastomycoses Pulmonares. 48, 1931. Monilicb asteroides Castellani, Jour. Trop. Med. Hyg. 17: 307-309, 1 fig., 1914. Oidium asteroides Castellani & Chalmers, Man. Trop. Med. ed. 3, 1095, 1919. Mycoderma asteroides Brumpt, Precis Parasitol. ed. 4, 1212, 1927. Isolated from stools in cases of pseudosprue, also reported from cases of chronic bronchitis. Colonies on glucose agar have characteristic vermiculate, more or less radiating appearance. Yellowish white to amber. Organism does not clot milk, grows badly or not at all on coagulated serum which is not liquefied. Gelatin liquefied very slowly. No fermentation of sugars ; acid on glucose, fructose, maltose, sucrose, galactose, and lactose. Castellani & Chalmers (1919) state that it produces acid and clots in litmus milk. Geotrichum famatum (Harrison) Dodge, n. comb. Mycotonda famata Harrison, Trans. R. Soe. Canada, Biol. Sci. 22: 216-217, 1928. Isolated from a wound in the hand by Dr. Rasch (No. 1136 London Col- lection). Prom young malt extract and malt agar cultures cells are spherical, el- lipsoid and cj^lindric with abrupt ends and rounded corners, sprouting from ends and sides, with groups of 5-6 cells resulting. Spherical cells 1.7-3.5/a in diameter, ellipsoid 3.5 x 2.0/i, in diameter. In malt extract, cells are somewhat larger, attaining 4.2 x S.S/x, cylindric cells, 4.5 x 1.7/i,. From 145-day-old cul- tures cells ellipsoid or cylindric with short hyphae, occasional large cells 6.6 X 5.0/A, occasional oil droplets. Good growth between 25° and 37° C. On malt agar, growth white, shiny, slightly raised, spreading. On B.P.B. agar, growth similar but pale blue. On potato, growth is pure white, slightly EREMASCACEAE IMPERFECTAE 219 waxy, and spreading. On malt gelatin surface growth is white, slightly raised and shining, gradually sinking into gelatin, forming a funnel-shaped depres- sion. Giant colony on malt gelatin round, shiny with raised margin showing concentric ring halfway from center to edge, slight radiate markings. In malt extract, there is formed a thin film and ring with flocculent turbidity in the liquid and heavy sediment. Very slight growth under olive oil. No fermentation of any sugar. Acid with ring formation and turbidity, clearing about the tenth day, in glucose, mannose, galactose, fructose, and sucrose (no ring in glucose). In other sugars growth similar, but no aciditj'" and no ring formation in glycerol and inulin. Sucrose inverted. Milk not coagulated, slight alkalinity in 115 days. Gelatin slightly liquefied in 29 days, completely in 116 days. Geotrichum rug-osum (Castellani) Dodge, n. comb. Endomyces rugosus Castellani, Brit. Med. Jour. 2: 1208-1212, 1912. Monilia rugosa Castellani & Chalmers, Man. Trop. Med. ed. 2, 827, Fig. 414, 1913. Hemispora rugosa Castellani & Chalmers, Man. Trop. Med. ed. 3, 1108, 1919. Parendomyces rugosus Ota, Denn. Woch. 78: 236, 1924. Trichosporum rugosum Ota, Jap. Jour. Derm. Urol 28: [4], 1928. Originally isolated from cases of bronchitis and tonsillitis in Ceylon. ?Isolated from cases of thrush by Pijper (1917) in South Africa. Colony yellowish amber, or brownish, surface crinkled, almost vermicu- late. No fermentation, slight acid produced on glucose, fructose, maltose, and galactose. Castellani states no action on milk, gelatin slowly liquefied. Pijper states that gelatin and serum were not liquefied by his strain, milk rendered acid, and slightly clotted and peptonized. It is possible that Pijper 's organism was Mycoderma pararugosum instead of this species. Geotrichum iimnite (Castellani) Agostini, Jour. Trop. Med. Hyg. 35: 266- 269, 3 figs., 1932. Blast omycoides immiiis Castellani, Amer. Med. 23: 290-291, 1928; Amer. Jour. Trop. Med. 8: 385, 386, 1928. Isolated by Castellani from a ease of "blastomycosis,'' authentic cultures studied and described by Agostini. Mycelium slender, hyaline, 2-2. 5/a in diameter, not branching, often uniting in fascicles, later mycelium thicker, 3-5/t, with cylindric arthrospores 6-9/a, racquet mycelium present. Large chlamydospores rich in fat globules, present. Colonies on Pollacci agar, white, fluffy, powdery, adherent to the surface, becoming slightly yellowish brown. Growth similar on carrot and potato. On mannitol agar, colonies dark brown to black with pigment diffusing into the medium. Glucose and maltose not fermented, milk not coagulated. No growth on blood agar. Gelatin and serum rapidly liquefied. Optimum tem- perature 22°-27° C. From the description of the organism given by Agostini, it is quite evi- dent that this organism is not related to Coccidioides immitis and that the synonymy quoted by her is incorrect. While Castellani does not state, it is 220 MEDICAL MYCOLOGY probable that the organism was isolated in Louisiana, perhaps the atypical case mentioned by Castellani (1933, p. 297), while Coccidioides is practically confined to California, most of the patients elsewhere giving a history of a recent sojourn in that state. It is also quite probable that Agostini is wholly unfamiliar with the references she quotes, or she would not suggest that the ascospores reported by American workers in Coccidioides are oil globules. Geotrichmn rotundatum (Castellani) Almeida, Annaes Fac. Med. Sao Paulo 9: 12, 1933. Endomyces rotundatus Castellani, Brit. Med. Jour. 2: 1208-1212, 1912; Arch, de Parasitol. 16: 184-186, 1913. Moyiilid rotunda Castellani & Chalmers, Man. Trop. Med. ed. 2, 828, 1913. Monilia rotundata Castellani, Jour. Trop. Med. Hyg. 17: 307, 1914. Oidium rotundatum Castellani & Chalmers, Man. Trop. Med. ed. 3, 1095, 1919. Mycoderma rotundatum Brumpt, Precis Parasitol. ed. 4, 1213, 1927. Mycelohlastanon rotundatum Ota, Jap. Jour. Derm. Urol. 28: [4], 1928. Candida rotundata Basgal, Contr. Estudo Blastomycoses Pulmonares 49, 1931. Isolated from saliva and stools from cases of sprue and enteritis. Growth on glucose agar wrinkled and yellowish. Acid formation with glucose, fructose, maltose, galactose, and lactose. Milk strongly acidified and coagulated. Coagulated serum not liquefied. Gelatin liquefied, sometimes slowly. Geotrichum Muisa (Mattlet) Dodge, n. comb. Torula Muisa Mattlet, Ann. Soc. Beige Med. Trop. 6: 25, 26, 1926. Patient a chief in the Belgian Congo who had a bronchitis of long stand- ing. Mucous sputum containing well-defined, greenish purulent islets which on examination showed cells of above organism. Medication with potassium iodide and sodium cacodylate caused amelioration of symptoms. Patient dis- appeared. In sputum, spherical cells, 4-5/a in diameter, with clear double membrane, uniformly colored content with some large refringent granules. No sprout- ing forms. Optimum temperature for growth 37° C. In potato decoction after three days at 37°, only hyphal forms. Branched hyphae of variable diameter, 1.5-3.5/a, septa distant, chlamdospores spherical, 3-7(U, in diameter. On Sabouraud agar at 37° C, colony amber yellow, coherent, with dull surface ; later wrinkled and furrowed, with abundant submerged hyphae. In potato decoction, it forms a voluminous sediment of yellowish flakes, no pel- licle. No fermentation of sugars. Coagulation and acid formation with milk. Gelatin liquefied. Geotrichum louisianoideum Castellani, Med. Press Circular 136: 439, 1933; Jour. Trop. Med. Hyg. 36: 304, 1933. Isolated from cases of primary pulmonary blastomycosis, one case from Louisiana, two subsequent cases from Southern Italy. EREMASCACEAE IMPERPECTAE 221 Hyphae undulating, 2-4/x in diameter, arthrospores abundant and very short; chlamydospores also produced. Asei [?] in tissues of experimental animals. On glucose agar, colony fluffy, white or grayish white. No pigmentation on mannitol agar. No fermentation or acidity on any carbohydrate. Milk neither coagulated nor digested, but may become alkaline. Gelatin and serum liquefied rapidly. Geotrichum versiforme Moore, Ann. Missouri Bot. Gard. 21 : 349-366, 1 pi. 1934. Isolated from thick, greenish, mucopurulent, at times bloody tenacious sputum from a patient with bronchiectasis and pulmonary infiltration of sev- eral years' duration. Organism varies in size and proportion on various standard media, at- taining the largest cells on malt extract agar. Hyphae 3-8fi in diameter with young cells approximately 6-40/x, long; arthrospores with abrupt or rounded ends 4-9 x 6-18/t; round, thick-walled chlamydospores 4-18/x in diameter and elongated, 6-8 x 20-30/a; small round cells 4-6/t in diameter, possibly blasto- spores; conidium-like cells, round 4-6/a in diameter, pyriform 3-4 x 4-6/x. On malt extract agar, cells spherical approximately 15/a in diameter ; arthrospores 6-9 X 12-18/x, also barrel-shaped cells and smaller spherical cells. Old cultures show many arthrospores and fewer hyphae (Fig. 46). Colonies vary from dull grayish white, on Raulin's, Richard's, and Czapek's agar, becoming light cream with age, to a dull cream to creamy-buff on Sabouraud's, nutrient, glycerol, and lactose agar. On potato-dextrose, growth grayish white to light cream, showing a plushlike whirl. Colony on malt extract vermiculate or pebbled. Saboui'aud colony velvety, while cultures on nutrient, lactose, and Endo's agar moist. Mycelium partially submerged in Richard's, Czapek's and oatmeal agar. In broth, pellicle and a fairly heavy sediment. No fermentation of sugars. Acid on maltose, galactose, d-mannose, 1-xylose, and fructose. No acid on lactose, sucrose, glucose, raffinose, 1-arabinose, rhamnose, and inulin. Milk slowly acidified and coagulated. Plain gelatin liquefied on surface after 12 days, beef-extract gelatin after 14 days. It seems probable that the following unnamed organism may belong here. Oospora sp. Bachmann, Centralbl. Bakt. I, 86: 129-132, 1921. This species is nearest to 0. lactis of Rabenhorst, Kryptogfl. Deutschl. ed. 2. Metachromatic granules lacking near vacuoles. Oidia 7-10//, in diameter, not multinucleate. Cultures, on maltose agar, gray white with seedlike surface. After 3 days, colony becomes yeastlike near periphery, margin showing mycelium, with cylindric or ovoid cells, 5-7/t long, occasionally chlamydospores up to lOfi with highly refractive wall. Longer filaments in young agar colonies. On gelatin plates, colonies as above, except that no filaments appear in 4-6 days. Myce- lium 2/A thick. Fermentation slight except with maltose. Malt gelatin is liquefied. 222 MEDICAL MYCOLOGY 1162- Geotrichum membranogenes (Martins) Dodge, n. comb. Zymonema pulmonalis memlranogenes Martins, C. R. Soc. Biol. 98; 1164, 1928. Isolated from a case of ill-defined pneumonopathy, frequent and abun- dant hemoptyses, temperature oscillating between 37° and 39° C. Portugal, Not markedly pathogenic for laboratory animals. I I Fig. 46. — Geotrichum versiforme. 1, 2, i, coenocytic cells, probably chlamydospores, on Sabouraud agar ; 3, submersed mycelium showing branching, in Sabouraud agar ; 5, aerial mycelium on same medium ; 6-8, cells stained to show volutin ; 9-13, cells showing glycogen as dark heavily stained material, lipoids as small hyaline granules, and small dark chondriosomes ; li-SS, cells showing volutin granules in the vacuoles ; 2S-S'i, living cells stained to show fat content. (After Moore 1935.) Mycelium present, septate, forming arthrospores and terminal chlamydo- spores. Growth at room temperature, but optimum at 37° C. On Sabouraud glucose and maltose agar, growth in 24 hours, colony cover- ing the medium and climbing the tubes, yellowish white, tough, easily separat- EREMASCACEAE IMPERPECTAE 223 ing from the medium, surface moist and finely mottled. On carrot, the same, except slight velvet on surface of colony. No spores on Gorodkova agar. On gelatin, growth slow, with liquefaction of medium. No development on several other media tried. Geotrichum bostonense Dodge, n. sp. Parasaccharomyces G Nye, Zerfas & Cornwell, Amer. Jour. Med. Sci. 175 : 163-164, 1928. Isolated from stools in case of carcinoma of stomach. Cells ovoid, 4-5/x, in diameter, few elongate forms and many septate hyphae. Cytoplasm slightly granular and contains vacuoles without refractive granules. No nucleated cells. On agar, colonies elevated, regular, shiny, and dirty green in color. In peptone water, growth at bottom of tube, with clear supernatant liquid and no pellicle. In gelatin stab marked mycelial production, but no lateral out- growths. Acid, without fermentation, with all sugars except sucrose and lactose. Gelatin liquefied. MYCODERMA Mycoderma Persoon, Myc. Europaea 96, 1822. The type species is Mycoderma mesentericum Pers. Morphology, in general, as in Geotrichum but gelification of the walls more complete, so that in old cultures spherical or ellipsoid cells are abundant; cells contain abundant oil globules and grow rapidly, soon covering liquid media with a thick folded pellicle with radial folds running to within about 2 mm. of the periphery; giant colonies sometimes yellowish, surface dull; gelatin not liquefied, sugars not fermented. While the majority of species in this genus are saprophytic, some species have been found on man, mostly confined to the respiratory and digestive tract, in the latter merely isolated from stools and not known to be etiologic agents of disease. A few species have been described as producing abscesses in the skin, all but one so poorly described that the reference here is not cer- tain. The one well-described species, Mycoderma nohile, differs from all the other members of the genus in the presence of raquet mycelium and in fer- menting sugars, and perhaps belongs in another genus. When its perfect stage is found, quite likely it will appear to belong to Zymonema. Key to Species Sugars fermented. Glucose and starch fermented, colony white, raquet mycelium present. M, nohile. Glucose, sucrose, and lactose fermented, colony becoming sulphur yellow, no raquet mycelium present. M. sulfureum. Sugars not fermented. Colony white. Colony zonate velvety. M. Nyahisi. Colony with elevated wrinkled yellowish center. M. virulens. 224 MEDICAL MYCOLOGY Colony yellowish. Colony elevated granular, yellow. M. Caoi. Colony wrinkled, yellowish amber, or brownish. M. paranigosum. Colony zonate. Arthrospores nearly all spherical. M. Muyaga. Arthrospores strictly cylindric, not abundant. M. Kieta. Arthrospores elongate ellipsoid [cylindric with rounded ends]. M. Issari. Mycoderma Nyabisi Mattlet, Ann. Soc. Beige Med. Trop. 6: 30, 31, 1926. Geotrichum Nyabisi Basgal, Contr. Estudo Blastomycoses Pulmonares 49, 1931. Isolated in Belgian Congo from a case of chronic bronchitis, showing regular presence of cylindric cells, 3-5 x 6-8/x. In potato decoction, close to Mycoderma Kieta but arthrospores rare, elongate, 1.5-2/a, chlamydospores barrel-shaped, 8 x S/u. On Sabouraud agar at 37° C, colony is white, velvety, with concentric zones of alternating long and short velvet. At 25° C. the velvet is thicker and the zones are less marked. The zone of propagation is moist, as in M. Issavi. Filaments abundant in the medium. Gelatin stab, surface white velvety, margin evenly indented. Acid formation on the second day with glucose, fructose, galactose, slight with inulin, solutions becoming alkaline on nearly all media on the 20th day, at least in the upper layer of the liquid. More strongly acid at the bottom of the tube in glucose, fructose, galactose. Inulin more strongly acid both at the bottom and immediately under the pellicle. No sugars fermented. Litmus milk turns slightly acid, but does not coagulate. Gelatin not liquefied. Mycoderma ? virulens (Norris) Dodge, n. comb. Saccharomycete pleomorphus virulens Norris, Southern Med. Jour. 24: 482- 489, 16 figs., 1931. Isolated from sputum in a ease of bronchomoniliasis. Toxins produced, causing anemia, leucopenia, and neutrophilic depression in experimental animals. Cells spherical to ovoid, 2-10>, hyphae up to 50/a in length. Cells usually uninucleate, though ovoid ones sometimes have 4 nuclei in old cultures. Hy- phae composed of clavate cells, with large terminal cells. No true lateral conidia. Arthrospores present, blastospores in clusters. Sprout cells divide by amitosis, hyphal cells by karyokinesis. Organism killed after 15 minutes at 58° C. Direct sunlight killed in two hours, inhibited growth in one hour. ''Ultraviolet light not harmful." Killed by metaphen 1:64,000; iodine, 7% solution in 1:120 dilution; half saturated KI ; gentian violet 1 :25,600. Colony white, center elevated, wrinkled, slightly yellow; margin white, very irregular and feathery. Growth on glucose agar white with yellow brown pigment on exposure to ultraviolet light. Yellow pigment on lead agar. On potato, growth slow, confluent, smooth, fine, white, not elevated. On Kracke & Teasley broth, a heavy, tenacious wrinkled pellicle forms, with heavy chalky EREMASCACEAE IMPERFECTAE 225 sediment. Alkali fonned on Endo's medium. Granular soapy pellicle on arabi- nose. No fermentation of sugars but slight acid formation with maltose, rhamnose, arabinose. No action on inulin, lactose, and mannite. Milk not coagulated and no acid formed. Gelatin not liquefied. Mycoderma Caoi Jannin, Les Mycoderma 186-187, 1913. Monilia Caoi Castellani & Chalmers, Man. Trop. Med. ed. 2, 831, 1913. Geotrichum Caoi Basgal, Contr. Estudo Blastomycoses Pulmonares 49, 1913. Oidium 22 Cao, Zeitschr. Hyg. 34: 317-318, 1900. Isolated from a case of chronic bronchitis in an old man. Pathogenic to rabbits. Cells large, hyphae long and little branched. Colonies on agar yellowish, granular, elevated with radial strands at margin. On potato, growth abundant Avith irregular margins, granular, yel- low with surface of potato browned. Star-shaped colonies on gelatin surface, inverted fir tree growth along the stab. No fermentation of sugars. No change in milk. Mycoderma paranigosum (Castellani & Douglas) Dodge, n. comb. Hemispora pararugosa Castellani & Douglas, Jour. Trop. Med. Hyg. 24: 149, pi. 1, 1921. Isolated from sputum, no fever, cough with mucopurulent expectoration, with no blood in mild type of cases. In severe type, cases resemble phthisis, patient emaciated, hectic fever with blood in expectoration; often develop- ing after severe tonsillitis, with yellowish gray patches. Physical examina- tion revealed patches of dullness, fine crepitation, and pleural rubbing. No fermentation or acidity with sugars, no action on milk, gelatin not liquefied. It is quite possible that the organism described by Pijper (1917) from South Africa under the name Monilia rugosa (p. 219) should be referred here, although it differs in minor characters. Mycoderma Muyag-a Mattlet, Ann. Soc. Beige Med. Trop. 6: 28-29, 1926. Isolated in cases presenting symptoms of true dysentery and enteritis in natives of Belgian Congo. No amebas or cysts present. Medication with intramuscular injections of emetine and administration of mild laxatives gave amelioration of symptoms. In potato decoction, hyphae septate, branched, 2.5-7^, narrowed at the septa. The young, slender hyphae are rich in protoplasm and contain few fat droplets, few or no septa. Secondary hyphae arise near, but not at, transverse walls. In older hyphae the positions of rupture are forecast by the condensa- tion of protoplasm about the fat droplets without thickening of membrane. Arthrospores nearly spherical, not thick-walled. Optimum temperature 22°- 25° C. On Sabouraud agar at 37°, colony creamy yellow, margin of radiating hyphae, adherent to the medium with radial folds. At 25°, colony covered 226 ME3)ICAL MYCOLOGY by a short, thin, tufted velvet, white with concentric zones. On gelatin stab, surface colony white, velvety, with fine radiations. In potato decoction, dense deposit of yellowish, filamentous flakes. Liquid turbid, of mucus consistency. Acid production with glucose, fructose, and galactose before pellicle forma- tion. When pellicle covers the surface, alkali production starts, but ceases when the pellicle is immersed. With maltose, sucrose, lactose, mannite, dex- trin, and inulin, the trend is definitely alkaline. No fermentation. Milk turned slightly acid, no coagulation. No liquefaction of gelatin. Mycoderma Kieta Mattlet, Ann. Soc. Beige Med. Trop. 6: 29-30, 1926. One of the fungi isolated from a case showing symptoms of enteritis in Belgian Congo. Treatment with mild laxatives and intramuscular emetine caused amelioration of symptoms. In potato decoction after 3 days, hyphae branched, 2.5-7/* in diameter, narrowed at the septa. The young, slender hyphae are rich in protoplasm and contain few fat droplets, few or no septa, and abjoint the arthrospores. Secondary hyhpae arise near, but not at, the septa. The older filaments form series of three or four chlamydospores each, square or slightly oblong. After 30 days there still remains a large quantity of mycelium. Optimum tempera- ture for growth 22°-25° C. On Sabouraud agar at 37°, colony dull, yellow cream, with concentric zones. At 25°, colony yellow, powdery; later the center becomes humid and zones, alternately dull and dusty, are formed. On gelatin stab, colony at surface white, velvety, mammillate at center, with fine rays toward the mar- gin. In potato decoction, sediment white coalescent tufts, the liquid remain- ing clear. Slight acid formation at first with glucose, fructose, and galactose. After the pellicle forms, the upper portion becomes alkaline, but alkali forma- tion ceases when the pellicle is immersed. No action on lactose and inulin. Maltose, sucrose, mannite, and dextrin become slightly alkaline. Milk slightly acidified, not coagulated. Gelatin not liquefied. Mycoderma Issavi Mattlet, Ann. Soc. Beige Med. Trop. 6: 26-28, 1926. Isolated from cases presenting symptoms of true dysentery and enteritis in natives of Belgian Congo. No amebas or cysts present. Medication with intramuscular injections of emetine and administration of mild laxatives gave amelioration of symptoms. In potato decoction after 3 days at 25° C, well-developed hyphae, branched, 2.5-7fi, narrowed at septa. The young, slender hyphae are rich in protoplasm and contain few fat droplets, few or no septa. Secondary hyphae arise near, but not at, the septa. Arthrospores cylindric when first ab jointed, ends rounding later. Short, spherical arthrospores rare. After 30 days most of the hyphae are changed into arthrospores. Spores of variable volume as hyphae of all sizes segment. The greatest measure 30 x Ifi. No yeast forms seen. Optimum temperature for growth 22°-25° C. On Sabouraud agar at 37°, colony cream yellow, humid, margin of radiat- ing hyphae, strongly adherent to medium, later little elevations at center. EREMASCACEAE IMPERFECTAE 227 When growth reaches edge of tube, hyphae push up along the glass. Below, abundant development of hyphae. At 25° on same medium, colony covered by short white velvet which forms concentric zones of different thicknesses with wrinkled center and humid zone of growth. On gelatin, velvety, white growth at surface with large rays from center toward edge. In potato decoc- tion, dense deposit of yellowish, filamentous flakes. Liquid turbid and of mucus consistency. Acid production with fructose, and galactose before pel- licle develops. With pellicle on surface, upper portion of liquid becomes alkaline but alkali production ceases if pellicle is immersed. No action on glucose. In sucrose and lactose slight acid formation at first but giving place to alkali by twentieth day. Trend strictly alkaline with maltose, mannite, dextrin, inulin. No fermentation. Slight acidification of milk but no coagula- tion. Gelatin not liquefied. Doubtful Position The following species are referred here either on account of the lack of adequate descriptions or because of characters which make their presence in this genus anomalous. Mycoderma nobile A. Sartory, R,. Sartorj?-, Meyer & Charles, Ann. Myc. 29: 325-338, 6 figs., 1931. Following a wound, organism produces a soft tumor suggesting a gumma on the forearm, with vertebral metastasis. Besides the Mycoderma, cultures on alkaline media showed colonies of Staphylococcus citreus, especially in coagulated serum. Pathogenic for guinea pigs. Cells in the tissue spherical, 5/a in diameter, or ovoid, 3.5 x Ifj., no mycelium observed. On carrot, long tufts of hyphae, up to 5fi in diameter, branches long and slender, l-2ja in diameter. Raquet mycelium present. Sometimes the branches are strongly curved, sometimes chlamydospores and arthrospores are formed. On agar and gelatin media growth velvety at first, finally becoming powdery or chalky or even moist, adherent, and viscous. On glucose media there is a blackening of the medium. On glycerol agar growth luxuriant, zonate, of glabrous and silky zones. On carrot, colonies whitish velvety, covering the medium. Growth similar on potato but a little slower. Growth only at 37° C, not at room temperature, ceasing at about the twelfth day. In liquid media tough pellicle, folded, forms a ring, falls to bottom after 15-20 days, and a new pellicle is formed. Acid and fermentation of glucose and starch. Acid with sucrose and lactose, slight fermentation with galactose and man- nose, no action or assimilation of fructose or maltose. Milk coagulated and digested. Gelatin not liquefied. Mycoderma sulfureum (Beauverie & Lesieur) Dodge, n. comb. Cryptococcus sulfureus Beauverie & Lesieur, Jour. Physiol. Path. Gen. 14: 996-998, 1912. 228 MEDICAL MYCOLOGY Monilia sidplinrea Vuillemin, Champ. Parasit. Homme 84, 1931, non Persoon. Isolated from exudate of pharynx of a woman suffering from a severe ease of typhoid fever. Inoculations negative. On carrot, after 5 days at 25° C, cells spherical, occasionally slightly oblong, 2-8/t in diameter. In malt extract (40 days) mycelium branched, septate, 1-4/^ in diameter, often terminating in a swollen budding head. No spores seen on plaster block. In Kaulin's liquid at 26° after 6 days, cells 2.5- 5.5/i,; after 36 days, cells 2-4. 5/i, in sediment. Grows at 25° and 37° C. On potato, growth good, margin with round denticulations, dirty white, becoming sulphur yellow, the upper portions of colony remaining dirty white and pulverulent. On malt gelatin, colony much wrinkled and prominent. In malt extract after 40 days, appear fioccose masses of mycelium floating and immersed. In carrot juice, sediment in 17 hours. In Raulin's liquid, after 53 hours, ring several centimeters high. In malt extract, a very small ring in 23 hours and pellicle over surface in 73 hours. Glucose, sucrose, and lactose fermented. Maltose not fermented. Gelatin not liquefied. Mycoderma multifermentans (Castellani) Dodge, n. comb. Geotrichum multifermentans Castellani, Med. Press Circular 136: 439, 1933; Jour. Trop. Med. Hyg. 36: 303, 304, 1933. Isolated from a case of "blastomycosis." Hyphae 2-4/a in diameter, arthrospores and chlamydospores present. On glucose agar, growth fluffy, white, grayish white, or grayish brown. No pigmentation of medium on mannitol agar. No fermentation, acid pro- duced on glucose, fructose, galactose, maltose, lactose, mannitol, glycerol, inulin, rhamnose, inositol, arabinose, xylose, and dextrin, and after 21 days in all the carbohydrates tried. Litmus milk alkaline, gelatin and serum not liquefied. Mycoderma subtile (Blanchard) Verdun, Precis Parasitol , 1912; Jannin, Les Mycoderma, 232-235, 1913. Oidium suhtile-cutis Babes, Termeszettuduomanyi Kozlony 14: 192-195, 1882; Biol. Centralbl. 2: 569-571, 1882 [DeBoyer & D'Antin, Prog. Med. 9: 1025, 1881]. Oidium subtile R. Blanchard, in Bouchard, Traite Path. Gen. 2: 839, 6, 1895. Endomyces subtilis Castellani & Chalmers, Man. Trop. ]\Ied. ed. 1, 601, 1910. Monilia suhtilis Castellani & Chalmers, Man. Trop. Med. ed. 2, 829, 1913. Parasaccharomyces suhtilis Froilano de Mello & Gonzaga Fernandes, Arq. Hig. Pat. Exot. 6: 273, 278, 1918. Geotrichum suhtile Almeida, Annaes Fac. Med. Sao Paulo 9: 12, 1933. Isolated from abscess on thigh and on abdomen. Round ulcers the size of a pea up to four times as large and going even deeper than the corium. Borders sharp, surrounded by a mulberry-red zone. Ulcer covered with crust 2-7 mm. thick. Radulescu & Babes inoculated rabbits subcutaneously. Typical ulcerations of disease developed in 3-5 days and were transmitted to other animals. Fungus recovered. EREMASCACEAE IMPERFECTAE 229 Hyphae straight at the surface of cultures, parallel, dicliotomous, partly homogeneous, partly septate, 6/x in diameter. In the middle third the hyphae turn outward, so that they end at the surface. Arthrospores ovoid, cylindric or elongate, ellipsoid, even biscuit-formed, abjointed from ends. No asci or chlamy- dospores. Mycoderma pseudoalbicans (Neveu-Lemaire) Dodge, n. comb. Monilia pseudoalhicans Nevevi-Lemaire, Precis Parasitol. Hum. 77, 1921. Monilia albicans Magrou, Montpellier Med. 39: 278-282, 2 figs., 1918. Mycelohlastanon pseudoalbicans Ota, Jap. Jour. Derm. Urol. 28: [4], 1928. Isolated from sputum of bronchitis patient. No trace of stomatitis or other symptoms of M. albicans infection. Subcutaneous inoculation of yeast cells into guinea pig formed an abscess, which was resorbed. Intravenous in- jection into ear of rabbit caused death. Organism recovered from blood; kidneys, especially, hypertrophied and widely invaded by parasite. In sputum, hyphae 2-4ju, in diameter, dissociating into short, rectangular arthrospores. Lateral and terminal yeast cells. On carrot and Sabouraud glucose, yeast forms develop first. Later, as medium dries, mycelium of septate, branched hyphae with lateral or terminal spherical chlamydospores, 10-11/A in diameter. CANDIDA Candida Berkhout, De Schimmelgeslachten Monilia, Oidiuni, Oospora en Torula 63, 1923, excl. syn. The type species is Candida vidgaris Berkhout based on a culture isolated by Kloecker under the name Monilia Candida but evidently not that species. Colonies membranous, thick, radially folded or areolate, with tufts of hyphae giving the colony a velvet}^ appearance ; margins not lobulate ; blasto- spores often thick-walled, similar to arthrospores, others ovoid or irregular, sometimes pyriform and suggesting conidia, often very large, 7-10/a. Pseudo- mycelium well developed, hyphae little branched, flexuous, not easily breaking apart, ends of cells flattened, cells filled with fine oil globules. Terminal cell of hypha variable, often being a chlamydospore of variable form, rarely a chain of chlamydospores ; coremia abundant, verticils more or less regular, composed of a few pyriform blastospores (Fig. 47, 9). All but one of the species of this genus so far reported attack the respira- tory tract. One is suspicious that the extremely slender mycelium of C. bethaliensis belongs in Actinomyces. Key to Species Only glucose and fructose fermented. No pellicle on liquid media. C. urinae. Pellicle on liquid media. C. Krusei. Maltose also fermented, no pellicle. Cells very slender, 0.75-1.5,11 in diameter, no pellicle. C. hethaliensis. Cells normal, 3-4ti in diameter. C. tumefaciens. Sucrose also fermented but not maltose, cells 2-3. 5/i. in diameter. C. rhai. 230 MEDICAL MYCOLOGY Candida urinae Dodge, n. sp. Oidiiim Stamm I Gentzsch, tJber patliogene Sprosspilze bei Diabetes, Inaug. Diss. Med. Fak. Univ. Jena, 30 pp., 1908. Isolated from urine of a diabetic, drawn from the bladder under sterile conditions. Pathogenic for rabbits and mice. Fig. 47. — Candida (Geotnchoides Langeron & Talice). I, irregular blastospores ; 8, giant blastospores ; 5, clavate tips ; i. Inverted clavate tips ; 5, chains of arthrospores : 6, typical bits of mycelium ; 7, sinuous hyphae ; 8, pyrif orm blastospores ; 9, hyphae with, verticils of blasto- spores. (After Langeron & Talice 1932.) Yeast cells spherical or ovoid, occasionally some filaments seen in old cultures. On agar, colonies white, confluent, dull, the deeper ones somewhat yellow- ish. On gelatin, colonies similar to agar but growth slower. On potato, growth good, similar to that on agar. On alkaline broth and glucose broth, I EREMASCACEAE IMPERFECTAE 281 a granular precipitate on the walls of the tube but no pellicle, flocculent sedi- ment. On malt extract, a ring but no pellicle. Glucose and fructose fermented after 3 days; acid produced with inulin, maltose, sucrose, lactose, and gal- actose. No action on milk or gelatin. Candida Krusei (Castellani) Basgal, Contr. Estudo Blastomycoses Pul- monares 50, 1931, Saccharomyces Krusei Castellani, Philippine Jour. Sci. B 5: 197-203, 1910. Endomyces parairopicalis B Castellani, Centralbl. Bakt. I, 58: 236-238, 1911. Endomyces Krusei Castellani, Brit. Med. Jour. 2: 1208-1212, 1912; Arch, de Parasitol. 16: 184-186, 1913. Moiiilia Krusei Castellani & Chalmers, Man. Trop. Med. ed. 2, 826, 1913. Mycelohlastanon Krusei Ota, Jap. Jour. Derm. Urol. 28: [4], 1928. Geotrichoides Krusei Langeron & Talice, Ann. Parasitol. Hum. Comp. 10: 67, 1932. fMycoderma Krusei Stovall & Bubolz, Jour. Infect. Dis. 50 : 73-88, 3 figs., 1932. Geotrichoides tumefaciens Talice & Mackinnon, Reunion Soc. Argentina Patol. Reg. Norte 8: 162, 1933. Found in sputum of a convict suffering from a severe cough, other symp- toms obscure. Case under observation for two months. Monilia iropicalis also present. Castellani also reports this organism in a case of urethritis with yellow discharge in a Serbian officer. On glucose agar, very thick growth (Fig. 48). On other agars, delicate, whitish growth. On serum, white growth. Broth and other liquid media re- main clear with pellicle of variable thickness and light sediment. Glucose and fructose litmus broth fermented and acid produced, no change with other sugars. No acidification or coagulation of milk. Coagulated serum not liquefied. Gram-positive. Creamy white, dull surface. Cells oval. — ^Zepponi (1931). Candida bethaliensis (Pijper) Dodge, n. comb. Monilia sp. Pijper, Med. Jour. S. Africa 12: 129-130, 1917. Monilia dethaliensis Pijper in Castellani & Chalmers, Man. Trop. Med. ed. 3, 1091, 1919. MyceloMastanon hethaliense Ota, Jap. Jour. Derm. Urol. 28: [4], 1928. Isolated from sputum of patient suffering from cough, periods of fever- ishness, bronchitis, hyperleueocytosis. Sputum whitish yellow. No tubercle bacilli found. Vaccine treatment improved but did not cure cough. Inocula- tion to rabbit proved fatal in 48 hours. Hemorrhages in lungs. Organism not recovered. Cells 0.75-1.5ju in diameter. Septate hyphae up to 80/x, in length. On Sabouraud agar, growth in a uniform whitish layer with yellow tinge. No filament formation. On potato, growth slightly verrucose and yellower, with filament formation. In gelatin, a fir-tree arborescence formed. Sedi- ment forms in broth, liquid remaining clear. Acid formation and fermenta- tion of maltose and glucose, acid only with sucrose, no action with lactose, dextrin, or mannite. 232 MEDICAL MYCOLOGY Candida tumefaciens (Pollacci & Nannizzi) Dodge, n. comb. Saccharomyces tumefaciens alhus Foulerton, Jour. Path. Bact. 6: 37-63, PI. 4, 1900. Monilia tumefacieiis alba Ota apud Motta, Atti R. Accad. Fisiocrit. Siena IX, 17: 639-654, 4 figs., 1926. Monilia tumefaciens Pollacci & Nannizzi, I Miceti Patog. No. 55, 1927. Myceloblastanon tumefaciens album Ota, Jap. Jour. Derm. Urol. 28: [4], 1928. Pig. 48. — Candida Krusei showing intercalary chains of blastospores or arthrospores (?). (After Langeron & Talice 1932.) Geotrichoides tumefaciens Langeron & Talice, Ann. Parasitol. Hum. Comp. 10: 68, 1932. Isolated from two cases of pharyngitis along with Corynehacterium diph- theriae. Pathogenic to rabbit, producing tumors at point of inoculation and small nodules elsewhere. Cells 3-4vu. in diameter. No mycelium or spores observed. Case of Motta shows chains of cells, some elongate, terminal cell swollen, hyphae simple or EREMASCACEAE IMPERFECTAE 233 branched, cells cylindric, 18-30 x S-^fi, ending in chains of ovoid blastospores. Grows at 22°, better at 37° C. Gram-positive. On nutrient agar and nutrient glucose agar, growth spreading, thick white, moist. On potato, growth dry and white becoming brownish. On coagulated serum, growth scanty, dry, thin, white. Growth in peptone broth scanty, liquid clear, no surface growth, sediment in bottom of tube. In alka- line litmus milk, growth good, no coagulation, no acid. Glucose rapidly fer- mented, maltose slowly, no action on lactose. Candida rhoi (Castellani & Chalmers) Almeida, Annaes Fac. Med. Sao Paulo 9: 11, 1933. Endomyces rhoi Castellani & Chalmers, Man. Trop. Med. ed. 1, 601, 1910. Monilia rhoi Castellani & Chalmers, Man. Trop. Med. ed. 2, 828, 1913. Parasaccharomyces rohii Froilano de Mello & Gonzaga Fernandes, Arq. Hig. Pat. Exot. 6: 272-273, 277, 1918. My celohl a station rhoi Ota, Jap. Jour. Derm. Urol. 28: [4], 1928. Isolated in case of otomycosis, also from tea samples. Mycelium septate, 2-3. 5/t in diameter, and spherical free "spores,'' 3.5-5/x in diameter, also arthrospores and intercalary chlamydospores. Acid formation and fermentation with glucose, fructose, and sucrose, slight with galactose. Very slight acid with maltose. No coagulation of milk. No liquefaction of gelatin or coagulated serum. Doubtful Position The following species are too poorly described to refer here with cer- tainty but probably belong here. Monilia Fioccoi Pollacci & Nannizzi, I Miceti Pat. Uomo Anim. 8 ; No. 77, 1928. Isolated from a man in Venice suffering from diffuse dermatosis with numerous ulcerous pustules, chiefly on the upper part of the body and arms (for additional case, see Zoon 1930, 1931). Cells ovoid, ellipsoid or spherical, minutely guttulate (one or many), occurring singly, sometimes together in short chains 7.3-9.7 x 3.5 or 7-8/* in diameter; hyphal cells cylindric, elongate, rounded at each end, easily separat- ing, not or rarely branched, producing terminal chains of ovoid blastospores 7.3-7.5 X 4.8-5/A. Chlamydospores terminal, spherical, 7-8yu in diameter, unigut- tulate, granular. On Pollacci agar, colonies punctiform, small, 0.5-2 mm. in diameter, smooth, dirty white, confluent, moist. On Sabouraud agar, colonies remain isolated and become yellowish in age. Glucose, fructose and sucrose fermented. Trichosporum Asahii Jap. Jour. Derm. Urol. 29: 733-773, 1929. Lesions appear as red nodules growing to 5 cm. in diameter, circular, crateriform, eventually producing thin yellow crusts, accompanied by pruritus. Blasto-, arthro- and chlamydospores present. Gram-positive. Cultures unfortunateh' not described in the summary. 234 MEDICAL MYCOLOGY SCHIZOBLASTOSPORION Schizdblastosporion Ciferri, Arch. Protistenk. 71: 446-448, Fig. 6, 1930. The type species is ScMzohlastosporion Starkeyi-Henricii Cif. (Torula A Starkey & Henrici 1930.) Colony creamy, white, or slightly yellowish. Turbidity and sediment on liquid media, sometimes a trace of a ring but no pellicle present. Blastospores spherical or slightly ellipsoid ; mycelial cells elongate, ellipsoid ; pseudomyce- lium somewhat branched. No fermentation of sugars; gelatin liquefied. This genus is not well known, but three organisms of skin and nails seem to belong here. The genus seems intermediate between Geotrichum and Parendomyces. Key to Species Margin smooth; colony on malt agar smooth with central elevation; from nails, Austria. S. gracile. Margin festooned or dentate. Colony on malt agar smooth; from nails, Austria. S. glohosum. Colony on malt agar umbilicate, surface wrinkled ; from vesicular lesions of the feet, Japan. S. toTcioense. Schizoblastosporion gracile (Zach) Dodge, n. comb. Blast odendrion gracile Zach in Wolfram & Zach, Arch. Derm. Syphilis 169: 95-104, 5 figs., 1933. Isolated from a case of onychomycosis in Austria. Cells spherical to ovoid, 2.7-3.2 x 6.5, mostly 5.4 x 6.5/a. Mycelial cells elongate, ends rounded, 2.5-3.2 x 19.5/i, thin-walled with one small oil globule when young, later with a gelified wall, a large central vacuole, and 1-2 oil globules. Sprouting only at the poles. No ascospores. Colonies on maltose agar, dull, whitish, smooth with a smooth margin, later yellowish with a central elevation but without folds, margins sometimes showing pseudomycelial outgrowths. On gelatin, colony thin, white dull, margin wavy, slightly sinking into the medium. On acid potato, colony thicker, smoother more shining, grayish yellow. On carrot, colonies similar but white. In maltose broth and in yeast water, sediment slightly flocculent, no turbidity, no pellicle. In yeast water sometimes a slight turbidity. Growth good at 37° C. No fermentation, gelatin liquefied in 3 weeks. Schizoblastosporion globosum (Zach) Dodge, n. comb. Blast odendrion glohosum Zach in Wolfram & Zach, Arch, Derm. Syphilis 169: 95-104, 5 figs., 1933. Isolated from a case of onychomycosis in Austria. Cells spherical, mostly 5-5.5/a, giant cells up to 10.8-12. 8ju,. Pseudomycelial cells elongate up to 30/t long, slender, producing short ellipsoid blastospores, sometimes almost lacrimiform, in depauperate whorls. Sprouting not confined to the poles. No ascospores. Colonies on maltose agar dull, smooth, white, later slightly yellowish, margin wavy to dentate, sometimes showing pseudomycelial outgrowths. On gelatin, colony similar but weaker, somewhat striate and verrucose, margin EREMASCACEAE IMPERFECTAE 235 wavy, slowly sinking into the medium. On acid potato and carrot, colony thick, grayish yellow, more or less moist and shining. In maltose broth and in yeast water, sediment developed in 4 days, no turbidity and no pellicle. Growth good at 37° C, no fermentation, gelatin liquefied slowly after 1 month, becoming complete in 2^ months. Schizoblastosporion tokioense (Fujii) Dodge, n. comb. Mycelohlastanon iohioense Fujii, Jap. Jour. Derm. Urol. 31: 959-983, 15 figs. [71-72], 1931. Isolated from lesions on toes of young Japanese woman. Lesions began as a depression on the top of the foot near the toe, in time becoming whitish and more visible, progressing between the toes as erythematous vesicles which ruptured. Ulcerations became dry and formed a crust all over the toes. Vesicles and pustules abundant, reddish, covered by crust. Pruritus present. Intraperitoneal injection into mice caused enlargement of the abdomen, then death. In crust, organism shows spherical, ovoid, or angular cells, 10-7. 5-5. 5ju, in diameter, mostly thick-walled, some thin-walled and 2.7-3/* in diameter, not colored. Hyphae flexuous. On malt glycerol agar, cells long-ovoid, 10 x 7.5/i,, showing distinct vacuoles and chromatin granules, in chains or dendroid, dichotomous groups, slender, delicate, swollen in places. No spores on Gor- odkova agar. On Sabouraud agar in ten days, colony round, with margin sunk into medium and center umbilicate, becoming elevated with radiating wrinkles which start at center or in between but never reach margin, yellow gray above, reverse not colored. Growth best in drier parts. Hyphae seldom penetrate medium. After 20 days, center slightly depressed, radiations reach the margin which also is slightly depressed, surface otherwise flat, colony moist yellowish brown, reverse uncolored, medium not colored. On malt agar, colony 2.5-3 cm. in diameter, center slightly depressed, surface furrowed, part wet, part dry, margin comparatively transparent, partly sunk in medium. At the optimum temperature, colony finally 6 cm. in diameter, yellowish white with denticulate margin, center moist, umbilicate, wrinkled, slightly elevated, with short furrows, margin sunken with filaments projecting into medium. On malt glycerol agar, colony round, 2 cm. in diameter, center dry, irregularly wrinkled, elevated with radiating wrinkles. Margin moist. In malt extract, liquid is cloudy with a flocculent sediment. No fermentation of sugars, acid formation with glucose, xylose, and rhamnose. PSEUDOMYCODERMA Pseudomycoderma Will, Centralbl. Bakt. II, 46: 278-281, 1916. The type species is Pseudomycoderma vini Will. Cells fusiform, with 1-3 oil globules, shorter cells citriform suggesting those of Pseudosacckaromyces, also small spherical cells with a single oil globule. Pseudomycelium branched, of long slender cells, terminal portions of the branches not forked. Pellicle developing rapidly on malt extract, at 236 MEDICAL MYCOLOGY first small islets, resembling a drop of fat, then conflvient but still showing the points of union of the separate islets, the upper portions becoming chalk- white, gas bubbles collecting under the pellicle. In old cultures, the pellicle is compact and tough, brownish or slightly reddish. On other liquid media, the islets are rapidly confluent, the pellicle is vigorous, smooth, white, and glassy, easily sinking to the bottom. Colony folded, center crateriform, upper portions of the fold chalky. Gelatin slowlj' liquefied; sugars fermented; hydrogen sulphide produced in mineral nutrient solutions to which powdered sulphur is added. Key to Species Glucose fermented, colony white or brownish. P. Bahesalama. Glucose, galactose, and sucrose fermented, colony white. P. vesica. Glucose and lactose fermented, colony yellow to brick red. P. Tanakae. Glucose and mannite fermented, colony grayish. P. fecale. Glucose, mannite, lactose, and sucrose fermented. P. Mazsae. Pseudomyco derma Rabesalama (Fontoynont & Boucher) Dodge n. comb. Mycoderma Bahesalama Fontoynont & Boucher, Ann. Derm. Syphiligr. VI, 4: 330-338, Figs. 7, 8, 1922. Geotrichuni Bahesalama Basgal, Contr. Estudo Blastomicoses pulmonares 50, 1931. Isolated from abscess of lung with hemoptysis. Abscesses on rabbit heal spontaneously. Retroculture negative. Fungus produces rods with square ends, 20-25 x 4-4.5/a, filaments up to IGOyx in length, 3/a in diameter, terminal cells sometimes long and pointed, forming greatly branched holdfasts in contact with glass of culture vessel. On second day floating rods detach from filaments. On glucose agar, colony shows on third day, center 3-4 mm., elevated, with white rays. "Poor" agar as above, but rays raised from surface of agar. Maltose agar, same as glucose, but base gray "star" brown. On potato, cul- ture is light, velvety, same color as medium. Potato glycerol, white, dry lumpy. Gelatin glucose, colony white powdery with very slow liquefaction after 5 months. Glucose broth, uniformly turbid with eventual formation of thick pellicle, turbidity settles, finally also pellicle; on shaking, gas is evolved (fermentation of glucose). Pseudomycoderma vesica (Harrison) Dodge, n. comb. Mycotorula vesica Harrison, Trans. R. Soc. Canada, Biol. Sci. 22: 219, 1928. Isolated from the urinary bladder, Rigshospitalet. From young malt agar colonies, cells ellipsoid, cylindric or irregular, bud- ding at the ends and septa, ellipsoid cells 5 x 3.3)U,, cylindric cells 10 x 2fi. From pellicle on 56-day-old malt extract cultures, cells more variable in shape, oil globules more abundant, spherical cells 4/x in diameter, cylindric cells 15 X 1.8/A, ellipsoid cells 6 x 4/a, some hypliae present. On B. P. B. agar, colony spreading, shining, bluish with white margin. On malt gelatin, colony slightly crateriform with radial markings, margin slightly Avrinkled, white waxy. On potato, colony wrinkled to vermiculate. EREMASCACEAE IMPERFECTAE 237 On malt extract, a pellicle produced, liquid turbid and remains so, sediment heavy. Sucrose inverted, g'lucose, mannose, fiiictose, galactose, and sucrose fermented with acid production, acid production in arabinose and maltose. In xylose, rhamnose, mannite, maltose, lactose, and raffinose a pellicle is pro- duced without acidity; in giycerol, sorbite, dulcite, salicin, inulin, and dextrin only a ring is formed. Milk slightly akaline without coagulation or digestion. Gelatin liquefaction slow, becoming complete in 56 days. Anaerobic growth (under olive oil) slight, growth good between 25° and 37° C. Pseudomycoderma Mazzae Dodge, n. sp. Monilia sp. Mazza & Nino, Bol. Inst. Clin. Quirurg. Univ. Buenos Aires 4: 545-558, 15 figs., 1927. Isolated from hemoptoic expectorations of a patient suffering with cough and pains in the thorax. Pathog-enic to guinea pig, rat, and mouse, producing generalized infection Avith lesions at a distance from the point of inoculation. Filamentous forms present on most media along with abundant yeast cells, few large, spherical, thick-walled cells. Hypliae of variable diameter, about 1.5-2/jL, flexuous, rarely branched. Yeast cells singly or in groups. Hyphae rare on solid media. On carrot glycerol, yeast cells and broad bacilli- form cells. On potato glycerol, abundant yeast cells of small dimensions. Gram stain inconstant, not acid-fast. Growth good at 37° and 20° C. On Sabouraud agar, good growth, colony much folded and contorted. Drigalski medium changes color in 24 hours, slight gas in 4 days, then de- colorized. Litmus sucrose agar color change in 24 hours. Neutral red color change in 72 hours. On mannite, lactose, or glucose + litmus agar, color change in 24 hours. In broth, slight pellicle in 24 hours, with uniform tur- bidity and sediment. In acid Raulin solution, slight development, with sediment only. Ferments sucrose, levulose, lactose, arabinose, maltose, mannite, raffinose, slightly galactose, glucose, dulcite, inulin. No indol formation. Growth is strictly aerobic. Coagulated serum not liquefied. Milk coagulation begins in 24 hours and is complete in 4 days. Gelatin liquefied, with gas formation. Pseudomycoderma Tanakae Dodge, sp. nov. Saccharomyces sp. Tanaka, Mitt. Med. Ges, Tokyo 28: 1914; Jour. Path. Bact. 25: 350-355, PI. 18, 1920. Found in swollen mesenteric glands in 12 cases of cancer of stomach or intestines. Not pathogenic to mice, guinea pigs, or rabbits. Cells ellipsoid to fusiform, 2-6.6 x 1.5-3.7^, yellowish, not easily stain- ing. Spores 1-10/i in diameter, sometimes 16/a. [Spores here described are probably only granules since author states that in rapidly growing cells spores become smaller and very numerous. "Spores" stained by Moeller's and Klein's method.] Optimum temperature for growth 25° C. Grown on several different media, colonies vary between yellow and brick red in color. Pigment best developed on potato, less at 37° C, than at room temperature. Fermentation of lactose and glucose. Pseudomycoderma fecale Dodge n. sp. "Mycoderme" LeDantec, C.R. Soc. Biol. 74: 412-413, 1913 [Gaz. Hebdom. Sci. Med. Bordeaux 34: 83, 1913]. 238 MEDICAL MYCOLOGY Monilia de LeDantec Sartory, Champ. Paras. Homme Anim. 709, 710, 1922. Isolated from the feces of two sailors ill with nautical beriberi. Not pathogenic on hypodermic injection to mouse, guinea pig, or hen. Intraperi- toneal injection in guinea pig produced peritonitis. Growth on glucose agar, thick, creamy, grayish, center thickest, two marginal zones thinner. Surface moist, not velvety but rough with points, suggesting the surface of a stormy sea. Hyphae penetrate agar. No color change on ageing. In Sabouraud media, colonies appear in 24 hours and in 48 hours grow onto walls of tube. On potato, colony is gray ; on carrot, colony is white, creamy. On gelatin stab, growth resembles inverted fir tree. In broth, a silky pellicle forms and sinks to bottom, liquid is cloudy. In glucose broth, there develop an ethereal odor and acid reaction which in about 8 weeks give way to fetid odor and alkaline reaction. In peptone broth, the fragile pellicle soon sinks while another forms, wide ring, liquid turbid and of fetid odor. In milk, a pellicle forms, soft curd, acid at first, and then alkaline and fetid. Glucose and mannite fermented, lactose slightly, maltose and sucrose not at all. Milk coagulated. Gelatin liquefied in 25-30 days. Growth slow under anaerobic conditions. Doubtful Position The following species, perhaps saprophytes, isolated from cases of lingua nigra pilosa, may belong here but are too poorly described for certain deter- mination. Oospora catenata Schaede, Derm. Woch. 98: 521-527, 1934. Isolated from lingua nigra pilosa. Mycelium G-l/x in diameter, cells ellipsoid, 6-8.5 x 8-15/i, or spherical 8-10/x in diameter, clinging together in short chains. Colonies on malt agar dull, glassy white with slight grayish tinge, some- what rough. On carrots, colonies thick white, also on liquid media. Oospora fragilis Schaede, Derm. Woch. 98: 521-527, 1934. Isolated from lingua nigra pilosa. Mycelium 6/^ in diameter, easily separating into single cells. Ellipsoid cells 5-8 X 7-13. 5/x, spherical cells 5-7ix in diameter. On malt agar, colonies white, powdery, smooth; colonies thinner on car- rots and on liquid media. PARENDOMYCES Parendomyces Queyrat & Laroche, Bull. Mem. Soc. Med. Hop. Paris III, 28: 111-136, 1909. The type species is Parendomyces alhus Queyrat & Laroche. Colony creamy, mycelium scanty, limited to short chains in liquid media, cells ellipsoid; chlamydospores abundant; no ascospores. Pellicle formation rare, rings more common on liquid media, gelatin not liquefied, sugars not fermented. EREMASCACEAE IMPERFECTAE 239 Species which have been very poorly described as to morphology have been referred here if their biochemical reactions warranted, on the ground that if the morphology were distinctive, it would have been described. The genus as here treated is probably heterogeneous. It is found principally on mucous membranes, producing vaginitis, bronchitis, and enteritis, not produc- ing subcutaneous abscesses as far as known. Milk clotted. Key to Species Colonies white. Serum liquefied, from perionychia. Serum not liquefied, from bronchitis. Colonies yellowish. Growth on broth cloudy; from vaginitis. Growth on broth clear; isolated from sputum. Milk not clotted. Colonies yellowish, isolated from sputum. Colonies white. No acid formed in sugars. Cells 4-5ya; from macroglossia. Cells 6-6.5m; from cutaneous abscesses. Acid with glucose and maltose. Acid with maltose and galactose. Acid with maltose, galactose, and sucrose. Eeaction with sugars unknown. Isolated from vaginitis. Isolated from intestinal tract in sprue. Reaction with milk unknown. Eeaction with sugars unknown. Acid with glucose, sucrose, and maltose. Acid with glucose, sucrose, maltose, and dextrin. Acid with glucose, sucrose, maltose, dextrin, and galactose. No acid with sugars. P. periunguealis. P. zeylanoides. P. albus. P. seylanicus. P. zeylanicus. P. macroglossiae. P. Hessleris. P. inexorabilis. P. Perryi. P. Blanchardi. P. vaginalis. P. Vanderiurgii. P. Krausi. P. butantanensis. P. Vuillemini. P. communis. P. enterocola. Parendomyces periungiiealis (Nino) Dodge, n. comb. Monilia periunguealis Nino, Bol. Inst. Clin. Quirurg. Univ. Buenos Aires 5: 270-283, 1930. Monilia sp. Swartz, Arch. Derm. Syphilol. 18: 74-78, 5 figs., 1928. Mycotorula albicans Talice & Mackinnon, Reunion Soc. Patol. Reg. Norte Santiago del Estero 8: 161, 1933. Isolated from two cases of perionychomycosis in Argentine women. In one case, the finger nail was deformed, rough and dull, with longitudinal striations. Cuticle and skin of both cases loose and thickened at edge, exuding greenish yellow pus on pressure. Yeast isolated from pus. Amelioration by treatment with KI.Io and alkaline washes. Final cure effected by irradiation with x-rays. Pathogenic to rabbit, guinea pig, and rat. In pus, yeast cells mostly spherical, some sprouting, more or less refrin- gent. In culture on either solid or liquid media, at first only sprouting forms appear. Yeast cells which are variable in size, 2-4/t, spherical or ovoid, form 240 MEDICAL MYCOLOGY chains of 3-4 cells when the sprouting is bipolar, or in groups when sprout- ing is multipolar. These cells have a thin wall, more or less granular content, and a rather large vacuole which is almost always central. After a while hyphal forms appear, composed of mother cells and blastospores, rich in proto- plasm and of very variable size. Others are made up of several long cells. Blastospores lateral, spherical, or ovoid in form. Growth good both at room temperature and at 37° C. Mostly gram-positive, though in part negative. With Gueguen stain colored an intense blue. On solid media, colonies at first hemispheric, creamy, humid, not adherent to medium. With age the center flattens while the edge curves gradually downward to the medium. Good growth on Sabouraud glucose agar, potato or carrot and glycerol, or gelatin. Scant growth on agar in 4 days. In simple broth or Sabouraud glucose broth, good growth in 48 hours with the forma- tion of a pulverulent sediment on the sides of the tube. Starch paste not lique- fied. Probably no fermentation of sugars, although the author reports slight fermentation of arabinose and maltose, more of inulin. Milk coagulated in 48 hours with acid formation and partial digestion of clot. Coagulated human serum digested. Gelatin not liquefied. Parendomyces zeylanoides (Castellani, Douglas & Thompson) Dodge, n. comb. Monilia zeylanoides Castellani, Douglas & Thompson, Jour. Trop. Med. Hyg. 28: 257-258, 1925. Isolated from a case of bronchial infection, but case history not given. Colonies white, lead agar not darkened. No fermentation. Litmus milk clotted. No liquefaction of gelatin. Parendomyces albus Queyrat & Laroche, Bull. IMed. Soc. Med. Hop. Paris III, 28: 111-136, 1909. Cryptococcus alhiis Castellani & Chalmers, Man. Trop. Med. ed. 3, 1080, 1919. Monilia alba Sartorv\ Champ. Paras. Homme Anim. 707, 1923, not Castel- lani, 1911. Candida alba Almeida, Annaes Fac. Med. Sclo Paulo 9: 11, 1933. Isolated from little white spots on surface of the vagina. Spots easily washed off but recurrent within 12 hours. Patient experienced burning sen- sation and was unable to sit or sleep. Irrigation with a variety of antiseptics twice a day gave relief, but no cure. Finally arrested by a dressing of creo- sote and olive oil (20 gm. to 60 gm.). Rabbits, rats, mice, and guinea pigs found susceptible. Pigeons, monkeys, and kittens not susceptible. In the yeastlike form, cells are ovoid, slightly j^ointed at each extremity, sometimes pyriform, more rounded in age. Length 3.4/a, width 2.3/*, with extreme variations 6-2.5 x 4.5-1/x. Easily stained Avith aniline dyes. Gram- positive. Filamentous forms appear only in liquid media and are rarely 8-10 cells long. Reproduction is by budding and chlamydospores which are from EREMASCACEAB IMPERFECTAE 241 two to three times the size of ordinary cells, thick-walled and not staining well. These are abundant in Naegeli liquid to which 2% sucrose has been added and are 12-15/x, in diameter. Ascospores not seen. On ordinary agar, and sugar agars colony is yellowish white, thick, glisten- ing, creamy. Growth on potato plug slow, potato becomes brown, colonies somewhat dry and in rounded groups, not bands. Best growth on carrot plug placed in glycerol-glucose solution (100 c.c. water, 10 e.c. glycerol, 5 gm. glucose). After 12 hours at 38° C, the white colonies are thick at the center, confluent. Liquid cloudy at first, then clear with whitish deposit. Pellicle forms between carrot and glass. Growth stops in 3 days. At lower tempera- tures, growth slower. Color of carrot not modified, as the growth is strictly aerobic. On potato, in glycerol-glucose solution, growth is similar to that on carrot, but more rapid. On gelatin stab, abundant growth appears at stab on surface, then scattered colonies but no growth below. On gelatin streak, mar- gin finely toothed, thicker in lower part. No mycelial forms on gelatin. On coagulated serum, growth is good, colonies chalky white. In broth, ordinary or slightly acid, growth shows as cloudiness with abundant yellowish white sediment, sometimes a slight ring but no pellicle. In glycerol broth, growth is rapid with yeast cells only. In alkaline broth, much less abundant. In broth and alcohol (6 drops of alcohol to 6 c.c. broth), growth is poor at first, then good with occasional filaments of 3-4 cells appearing. In malt extract, growth rapid and abundant, white, \8yringospora albicans grows very poorly in this.] In red wine, growth is poor, yeast cells only. On media, colored with methylene blue or sky blue, organism grows well and assumes a greenish blue hue. Milk coagulated in 3-7 days. Gelatin not liquefied. Parendomyces zeylanicus (Castellani) Dodge n. comb. Endomyces zeylanicus Castellani, Arch, de Parasitol. 16: 184-186, 1913. Mo7nlia zeylanica Castellani & Chalmers, Man. Trop. Med. ed. 2, 826, 1913. Parasaccharomyces zeyla^iicus Froilano de Mello & Gonzaga Fernandes, Arq. Hig. Pat. Exot. 6: 273-274, 278, 1918. Myceloblastanon zeylanicum Ota, Jap. Jour. Derm. Urol. 28: [4], 1928. Candida zeylanica Basgal, Contr. Estudo Blastomycoses Pulmonares 50, 1931. Castellani isolated this organism from sputum. Froilano de Mello & Gonzaga Fernandes found it as a laboratoiy contaminant. Colony yellowish, acid with glucose, fructose, maltose, galactose, sucrose, dextrin; slight acid with lactose; very slight acid with inulin and raffinose. Litmus milk acid with slight coagulation. Gelatin and serum not liquefied. Pajrendomyces macrog-lossiae (Castellani) Dodge n. comb. Monilia macroglossiae Castellani, Jour. Trop. Med. Hj^g. 28: 219-221, 1925; Am. Jour. Trop. Med. 8: 389, 1928; Re, Jour. Trop. Med. Hyg. 28: 317-319, 1925. Torulopsis macroglossiae Castellani & Jacono, Jour. Trop. Med. Hyg. 36: 314-315, Figs. 40-42, 1933. 242 MEIDICAL MYCOLOGY Isolated from a case of macroglossia. Tongue enlarged, occasionally pain- ful, without the usual verrucoid patches of blastomycosis. Cells ovoid, 4-5/^ in diameter, gram-positive, not acid-fast. On ordinary agar and glucose agar, growth good, smooth, white. A little mycelium in liquid cultures. No fermentation of any sugar-peptone medium. Sometimes a slight acidity is produced with glucose. No change in litmus milk. Serum and gelatin not liquefied. Parendomyces Hessleris (Rettger) Dodge, n. comb. Blastomyces Hessleris Rettger, Centralbl. Bakt. I, 36: 519-527, 2 pis., 1904. Cryptococcus Hessleri Castellani & Chalmers, Man. Trop. Med. ed. 2, 771, 1913. Torulopsis Hessleri Almeida, Annaes Fac. Med. Sao Paulo 9: 10, 1933. Isolated from papules which slowly developed into abscesses following a cut while shaving. Pathogenic to rabbits, white mice, and guinea pigs [case of Hessler, 1898]. Cells spherical or slightly ovoid, 6-6.5/x, in malt extract becoming 9-lOju,. In animal tissue cells form mycelium ; cells often sprout as many as 7-8 daughter cells, mostly in a group at one end of the parent cell. No capsule or asco- spores noted. Hessler (1898) noted mycelium but did not describe its mor- phology. On agar, colonies white, opaque, circular. On gelatin, colonies small, white, circular, slightly elevated. In blood serum colonies light cream color. On potato, colony light cream color, texture cheesy. On beet, colonies coarsely granular, growth better. In broth, a fine sediment, liquid clear, no pellicle, hyphae occasionally seen. In malt extract, sediment heavy, grayish white with a thin grayish pellicle. No acid or fermentation with sugars. No action on milk. No liquefaction of gelatin. Syringospora inexorabilis (]\Iazza & Palamedi) Dodge, n. comb. Monilia inexorabilis Mazza & Palamedi, Reunion Soc. Argentina Patol. Reg. del Norte en Tucuman 7: 424-467, 1 pi., 50 figs., 1932. Mycotorula Candida Talice & Mackinnon, Reunion Soc. Argentina Patol. Reg. del Norte en Santiago del Estero 8: 165, 166, 1933. Lesion began in the commissural surface of the lips, edematous, spreading to the gingival mucosa, tongue, and interior of the mouth generally, producing ulcers. Thence it spread to the lungs (x-ray) the organism being isolated from sputum. Ulcers then appeared on the arms and foot. The disease proved fatal, autopsy showing the lungs to be the principal internal focus of infection. Yeast cells in tissues and pus spherical, somewhat suggesting the picture with Zymonema dermatitidis. Yeast cells predominate in cultures, but hyphae seen in potato decoction and serum. On Sabouraud glucose, colony white, creamy, slightly drier at the periph- ery, margin festooned, only yeast cells seen in the first 22 days. On simple agar and 3% glycerol agar, potato and potato-glycerol, growth similar but some hyphae noted in 14 days at 37° C. On liquid media, broth remains clear with EREMASCACEAE IMPERFECTAE 243 a thick sediment at the bottom of the tube. No fermentation, producing acid with glucose and maltose only. Produces an inverted pine tree in gelatin stab, without liquefaction ; egg albumen not digested, milk not coagulated. Data secured in my laboratory while this book is in page proof, indicate that this species belongs in 8yringospora (see p. 272). Parendomyces Perry! (Castellani) Dodge, n. comb. Endomyces Perryi Castellani, Arch, de Parasitol. 16: 184-186, 1913. Monilia Perryi Castellani & Chalmers, Man. Trop. Med. ed. 2, 829, 1913. Isolated from samples of tea dust. Colony white, slightly acid and fermentation vv^ith fructose and sucrose, acid only with glucose, galactose, maltose; slightly acid with raffinose and inulin. Litmus milk not coagulated, decolorized, slightly acid then alkaline; no action on serum or gelatin. Parendomyces Blanchardi (Castellani) Dodge, n. comb. Endomyces Blanchardi Castellani, Arch, de Parasitol. 16: 184-186, 1913. Monilia Blanchardi Castellani & Chalmers, Man. Trop. Med. ed. 2, 829, 1913. Isolated from tea dust. "White, smooth growth on maltose, glucose, and other sugar media. Slight fermentation of glucose, none of other sugars. Acid formed with fructose, maltose, galactose, sucrose; very slightly acid with raffinose and inulin. Milk at first slightly acid, then alkaline. Neither coagulated serum nor gelatin liquefied. Parendomyces vaginalis (Mazza & Los Rios) Dodge, n. comb. Monilia vaginalis Mazza & Los Rios. Bol. Inst. Clm. Quirurg. Univ. Buenos Aires, 6: 216-225, 13 figs., 1931, not Escomel. Isolated from viscous secretion of vagina in an Argentine woman suffer- ing from vaginitis. Authors unable to follow case to end, although case con- siderably improved by applications of KI.Io, and of Lugol solution and glycerol in equal parts alternating with alkaline washes. Intravenous injection killed rabbit in 50 hours. All of the internal organs showed this species. Cells elongate, forming septate mycelium. Occasional blastospores soli- tary at septa. On Sabouraud glucose agar, colonies round, of appearance of spots of stearine, viscous, confluent, not very adherent to medium and composed almost exclusively of yeast forms. On Drigalski medium, by puncture, growth on the surface with slight color change. Organism grows well on potato and glyc- erol, producing a white colony without pigmenting the medium. Similar on carrot and glycerol. In plain broth or with glucose, slight turbidity and sediment. Indol test negative. No fermentation with glucose, maltose, mannite, sucrose, dextrin, fructose, dulcite, arabinose, galactose, racemose, raffinose, lactose, or amygdalin. Coagulated human serum slowly liquefied, milk not coagulated, gelatin not liquefied. 244 MEDICAL MYCOLOGY Parendomyces Vanderburgii (Kohlbrugge) Dodge, n. comb. Oidiiim Vanderh^irgii Kohlbrugge, Arch. Schift's-Tropenhyg. 5: 394-397, 1901 [name and case history] ; Nederl. Tijdschr. Geneesk. 37: 2: 881-890, 1901 [description without name] . Isolated from a case of sprue in the Dutch East Indies. No discoloration of potato. No fermentation of sugars. Milk not coagu- lated but growth good, colonies milky white, not slimy, easily emulsifiable in water. Gelatin not liquefied. Parendomyces Krausi (Ota) Dodge, n. comb. Cryptococcus Krausi Ota, Derm. Woch. 78: 229, 1924. In 24 hours on malt agar, cells ellipsoid or ovoid, rarely spherical, aver- age cells 6-7 X 10/A with several small oil drops, in branched chains which break up into individual cells. In a month, the chains have all disappeared and the oil drops are larger. No true mycelium formation. No fermentation. Parendomyces butantanensis (Gomes) Dodge, n. comb, Monilia butantanensis Gomes, Annaes Paulistas Med. Cirurg. 12 : 1924 ; Mem. Inst. Butantan, 1924. Candida butantanensis Langeron & Talice, Ann. Parasitol. Hum. Comp. 10: 54, 1932. Isolated from sputum of a case clinically resembling tuberculosis. Patho- genic for laboratory animals. Cells 4-6/A in diameter, in pairs or short chains of 3-4 cells, spherical or ovoid, chains rather longer on carrot. No ascospores on Gorodkova agar. On Sabouraud maltose, potato or carrot, colonies Avhite, soft, and shining. In glucose broth, pellicle and sediment developed. No fermentation of sugars, acid with glucose, fructose, maltose, sucrose and arabinose, very slight with galactose and dextrin, no acid with inulin, mannite, and lactose. No action on coagulated serum or gelatin. Parendomyces minor (Pollacci & Nannizzi) Dodge, n. comb. Cryptococcus minor Pollacci & Nannizzi, apud Nannizzi, Tratt. Micopat. Umana [Pollacci] 4: 317, 1934. Torulopsis minor Lodder, Anaskosporogenen Hefen 1: 178-179, 1934. Cryptococcus dermatitidis Benedek in Lodder, Anaskosporogenen Hefen 1: 156-158, 1934 excl. all syn. Isolated from the scales of a case of psoriasis? [psionasi Lodder] by Spicca & Tarentelli. Source of Benedek 's culture unknown, probably from lesions clinically suggesting those of Atelosaccharomyces hominis or perhaps even Zymonema dermatitidis, as Lodder has confused it with these organisms. Cells small, short ovoid, 2.5-4 x 3.5-5 (-6. 5)/a clinging together in short chains of 4-6 cells. On malt agar, colony almost white, dull, center verrucose, margin almost smooth. On malt gelatin, yellowish, dull, center slightly elevated and of deeper color, margin slightly ragged. In malt extract ring white, well-developed sedi- ment, and occasional floating islets. Good growth on alcohol; no fermenta- tion, no liquefaction of gelatin. EREMASCACEAE IMPERFECTAE 245 Parendomyces Vuillemini (Froilano de jNIello & Gonzaga Fernandes) Dodge, n. comb. Endomifces V uillemini Froilano de Mello & Gonzaga Fernandes, Arq. Hig. Pat. Exot. 6: 230-234. 1918, excl. syn. Isolated five times from sputum in respiratory diseases. Yeast cells, elongate cells, and septate, branched mycelium present. On Sabouraud glucose agar, colonies dry, white, with margins indented, separable from the medium. Asci abundant, mycelial elements rare, chlamy- dospores terminal or intercalary. On neutral agar, colonies circular, 5-7 mm. in diameter, white, elevated, waxy, surface moist and shining. Asci ovoid, ellipsoid 1-4-spored, of variable dimensions [ ?were these oil droplets]. Some arthrospore formation. On alkaline agar, colonies whitish, humid suggesting bacterial colonies. No mycelium, asci 4-spored. In alkaline and neutral broth, turbidity, no pellicle, floccose deposit, yeast cells, ascospores 1-2 per ascus, no mj^celium. In acid broth, gelified sediment and turbidity with yeast cells, pseudomycelium, and asci. Acid only, with glucose, sucrose, maltose, dextrin, no action with lactose, mannite, or fructose. No fermentation. From the description, it seems quite likely that the large refringent oil globules of the older cells have been mistaken for ascospores and that this species as described by Froilano de Mello & Gonzaga Fernandes really belongs in Parendomyces. Parendomyces communis (Castellani) Dodge, n. comb. Monilia communis Castellani, Ann. Inst. Pasteur 30: 152-153, 1916. Broth and peptone water clear. Very slight acid formation and fermen- tation with fructose. Acid only with glucose, maltose, galactose, sucrose or dextrin. No action with others. Gelatin not liquefied. Parendomyces enterocola (Macfie) Dodge, n. comb. Monilia enterocola Macfie, Ann. Trop. Med. Parasitol. 15: 275-279, 1921. Mycelohlastanon enterocola Ota, Jap. Jour. Derm. Urol. 28: [4], 1928. Isolated from feces of a patient suffering from persistent diarrhea accom- panied by ascites and edema of the face, especially the left parotid area, the lumbar region, and the penis. Feces semifluid, pale canary yellow in color, frothy, containing numerous yeastlike cells. Grown on neutral-red lactose bile-salt agar. Hyphae often predominate in liquid media, yeast forms on solid media with occasional branched, septate hyphae. Gram-positive, not acid-fast. Colonies grow slowly, appear only after 48 hours. On glucose agar, growth abundant, white, cells somewhat elongate. Under anaerobic condi- tions growth somewhat slower. On potato, growth abundant and whitish, medium not stained. On gelatin, growth is slow. In broth and peptone water, whitish sediment but no pellicle is formed, medium remains clear. No fer- mentation or acid formation with glucose, fructose, maltose, or galactose. Gelatin not liquefied. 246 MEDICAL MYCOLOGY CASTELLANIA Coloniis albidis, humidis, nitidis, pseudomycelio non bene evoluto vel morphologia ignota ; gelatina non liquef acta ; f ermentatio adest. The type species is Monilia hronchialis Castellani. Colonies white, humid, usually shining, pseudomycelium not well developed, morphology unknown; gelatin not liquefied, sugars fermented. After all the species of the imperfect filamentous yeasts have been dis- tributed to existing genera as far as possible, one is confronted with many species which have been described without a sufficient characterization of their morphology. They have mostly been based on a minute description of their reaction with sugars and the coagulation of milk. They constitute the bulk of Monilia as described by Castellani & Chalmers (1913). While it is pos- sible that some species have been included here which will be found to belong elsewhere, it is felt that they make a natural group. They are mostly sapro- phytic or weakly parasitic in lungs, mouth, and intestinal tract. It is with great pleasure that I dedicate this genus to Castellani, who has done so much work on this group. There are a few species which differ from the greater portion of the genus in having a thin pellicle on broth. The position of this group of species is uncertain since they have strong resemblance to Pseudomonilia, but it has been thought best to place them here until more is known of their morphology, especially as they differ from Pseudomonilia by their fermentation of sugars. Key to Species No pellicle on broth. Only glucose fermented. Not growing on ordinary media, colonies yellowish to brown, isolated from trachom- atous tissue. C. Nogami. Growing on ordinary media. Colony yellowish or darker, brown floccose sediment in liquid media, from onychomycosis. C. unguium. Colony white. Eing or floating islets forming on liquid media, also sediment, from ulcero- membranous stomatitis C. Lesieuri. No ring or sediment in liquid media. Milk not coagulated, fiom sputum. C. halcanica. Milk coagulated. C. parahalcanica. Eeaction with milk unknown, from thrush. C. MetchniJcoffl. Colony color unknown. C. hominis. Glucose and fructose fermented. Colony yellowish or brownish, lesions resembling furunculosis. C. Castellanii. Colony white with center yellowish. Center folded, from mouth. C. dissocians. Center not folded, from skin. C. epidermica. \ EREMASCACEAE IMPERFECTAE 247 Colony white. Ring on liquid media, from tear ducts of ass. No ring on liquid media. Milk coagulated. C. Mirandei. C. Copellii. C. londinensis. Negrii. intestinalis. macedonensoides. Milk not coagulated. Acid in maltose. Acid in galactose Glucose, fructose, and galactose fermented. Glucose, fructose, and maltose fermented. Milk coagulated. No acid on other sugars. Acid on galactose, sucrose, and dextrin. Milk not coagulated. On coagulated serum a brownish pigment which diffuses into the medium. C. faecalis. On coagulated serum not forming a pigment. Glucose, fructose, maltose, and sucrose fermented. Milk coagulated. Curd digested. Curd not digested. Milk not coagulated. Acid with raffinose. No acid with raffinose. Glucose, fructose, maltose, and galactose fermented. Milk coagulated. Acid with sucrose. Inulin fermented. Dextrin fermented. Milk not coagulated. Dextrin fermented. Acid with pentoses, mannite fermented. No acid with pentoses. Dextrin not fermented. Eaffinose fermented. Eaffinose not fermented. C. Nabarroi. 0. decolorans. C. bronchialis. C. Bogerii. C. aegyptiaca. C. pulmonalis. C. 'burgessi. C. pseudometalondinensis. C. pseudolondinoides. C. mannitofermentans. C. pseudolondinensis. C. nivea. C. metalondinensis. C. Bichmondi. Glucose, fructose and sucrose fermented. Milk coagulated, clot digested. C. Guilliermondi. Milk not coagulated. No acidity on other sugars. C. Muhira. Galactose acidified. C. Guilliermondi. Maltose acidified. C. Chalmersi. Maltose not acidified. C. Lustigi. Glucose, fructose, maltose, galactose, and sucrose fermented. Milk coagulated. C. metatropicalis. Milk not coagulated. Dark pigment diffusing into serum medium, milk becoming alkaline. C. insolita. Dark pigment not diffusing into medium, milk becoming acid. C. tropicalis. 248 MEDICAL MYCOLOGY Lactose fermented. Milk coagulated, colonies white. Sucrose fermented. C. Sucrose not fermented. C. Keaction of milk unknown, colonies gray or even brown. Sucrose fermented. Sucrose not fermented. Thin pellicle on broth. Glucose and fructose fermented. Milk not coagulated. Milk coagulated. No acid on other sugars. Acid on oilier sugars. Glucose, fructose, and sucrose fermented. Glucose, fructose, and dextrin fermented. Glucose and maltose fermented. Glucose, maltose, and galactose fermented. Sucrose not fermented. Sucrose fermented. Slight or no fermentation with maltose. Maltose fermented. Isolated from bronchomycosis. Isolated from stools. Valeriana, pseudotropicalis. Kartulisi. Hashimotoi. C. linguae-pilosae. C. C. C. C. C. pa raknisei. nitida. Orticoni. africana. platensis. C. alba. C. accraensis. paratropicalis. enterica. Castellania Nogami Dodge, n. sp. Crijptococcus sp. Oclii, Klin. Montatsbl. Augenlieilk. 86: 309-313, 1931; Nogami, Ihkl. 86: 313-316, 1931. Isolated from trachomatous tissue in 19 out of 32 cases. Optimum tem- perature 22°-25° C. Grown on kojisenim agar. [This is made up of 3 parts "kojiwasser" and 1 part human serum with 3% agar. Kojiwasser is rice fermented by Aspergillus oryzae.] The organism may also be grown on a modified Loeffler medium. [This is made as follows: to horse meat broth con- taining 2% glucose or maltose and 1% glycerol, add horse or human serum, 1 part to 3 of the broth. Sterilize at 65° C. for 2 hours on each of 4 successive days, raising temperature on last day to 90° C. to produce semicoagulation.] Colonies at first colorless, then white, yellowish, browiiish to reddish, and, finally, dark brown. Colonies hard, deep hyphae thicker in Loeffler than in koji medium. On koji serum agar, colonies more superficial, mycelium less abundant, spores slightly smaller. Growth either aerobic or anaerobic. Glu- cose fermented. Gelatin not liquefied. Castellania unguium (Bourgeois) Dodge, n. comb. Saccharomyces unguium Bourgeois, Derm. Zeitschr. 22: 411-420, 1915. Found in cases of onychomycosis of the hands with nails yellowing and breaking, causing some pain. It is not virulent for animals. Intraperitoneal injection results in the formation of grayish white nodules in the liver from which the organism may easily be recovered. Cells 2-5 X l-3yu,, solitary or in chains of 3-4 cells. Branched hyphae appear in old liquid cultures. EREMASCACEAE IMPERFECTAE 249 Colonies flat or hemispheric, ivory white on neutral agar, yellowish on maltose agar, dirty grayish red on potato, becoming flattened, convex, 1 cm. in diameter. On gh^cerol or glucose agar, colony smooth or finely folded, bright brown to whitish. On gelatin, growth along stab is slow but surface growth is abundant. In beef and maltose broth, there is a brown floccose sediment but no cloudiness. Glucose fermented. Litmus milk turns alkaline. Castellania Lesieuri (Beauverie) Dodge, n. comb. Cryptococcus Lesieuri Beauverie, Jour. Physiol. Path. Gen. 14: 994-996, 1912. Cryptococcus Lesieurii Castellani & Clialmers, Man. Trop. Med. ed. 2, 771, 1913. Isolated in a case of ulceromembranous stomatitis in a young woman with typhoid. Not pathogenic for experimental animals. After 4 days on malt agar at 26° C, cells are ovoid, 2-3/*, becoming 8x3 or 12 X 2fx, although mostly 5 x 2ix. Both sprouting forms and hyphae present. No spores formed on plaster blocks. On malt agar, growth cream white, surface finely folded, vermiculate to the edge. On potato at 26° C, growth is less rapid and less abundant, dirty white, the upper part appearing dry, pulverulent and chalky white. On car- rot, after 3 days at the same temperature, development is abundant, white with the surface viscid and brilliant, formed of small connivent mounds. Growth on malt gelatin, at the laboratory temperature of 15°-18° C., is the same as on the agar. In malt extract, after 9 hours a ring and islets appear, a light sediment in 3 days with white ring well developed. Islets white and dry, 0.5-1.5 mm. in diameter. In carrot juice, development is rapid. After 8-9 hours the surface is covered with small, white islets, smaller than in malt, about 0.5 mm. in diameter. No ring forms although a fcAv of these islets are drawn up onto the sides by capillarity. Sediment present. In Raulin's liquid, development is slow, a sediment appearing in 4 days. Glucose fer- mented, not fructose, sucrose, maltose, or levulose. Gelatin not liquefied. Castellania balcanica (Castellani & Chalmers) Dodge, n. comb. Monilia halcanica Castellani & Chalmers, Man. Trop. Med. ed. 3, 1090, 1919. Mycelohlastanon lalcanicum Ota, Jap. Jour. Derm. Urol. 28: [4], 1928. Found in sputum. Hoffstadt & Lingenfelter (1929) describe in detail a case of pulmonary infection due to this organism. They found it pathogenic for rabbits. Cells 4 X 7/A. Hyphae composed of chains up to 6 cells. Hoffstadt & Lingenfelter give cultural characters of their strain which had no action on fructose. Organism originally described as fermenting glucose only, with a slight acid formation in fructose and arabinose (due, per- haps, to impurities in the sugars). Castellania parabalcanica (Castellani & Chalmers) Dodge, n. comb. Monilia parahalcanica Castellani & Chalmers. Man. Trop. Med. ed. 3, 1090, 1919. 250 MEDICAL MYCOLOGY Myceloblastanon parahalcanicum Ota, Jap. Jour. Derm. Urol. 28: [4], 1928. Glucose only fermented, with acid formation. Milk coagulated. Castellania Metchnikofla (Castellani) Dodge, n. comb. Monilia Metchnikoffi Castellani, Ann. Inst. Pasteur 30: 149-154, 1916. Isolated from a ease of thrush in London. Peptone broth remains clear. Acid formation and fermentation with glucose, slight with fructose and maltose. Slight acid formation with galac- tose. Neither coagulated serum nor gelatin liquefied. Castellania hominis (Castellani & Chalmers) Dodge, n. comb. SaccJiaromyces sp. Klein & Gordon, Centralbl. Bakt. I, 35: 138, 139, 1904. Saccharomyces hominis Castellani & Chalmers, Man. Trop. Med. ed. 2, 769, 1913. Not Busse, 1895. Isolated as a contamination during an epidemic of tonsillitis which re- sembled diphtheria. Not pathogenic to rabbit and guinea pig. Yeast cells spherical or ovoid, 5-7 x 1-3/x, with pseudomycelium formed only in media containing glucose. On sugar media, colonies red, round, margins pellucid. On sugar broth, sediment but no ring or pellicle. On milk, red pellicle curd slowly digested. Glucose not fermented. Gelatin slowly liquefied. Castellania Castellanii (Re) Dodge, n. comb. Cryptococcus sp. Castellani, Jour. Trop. Med. Hyg. 27: 326-328, 2 figs., 1924. Monilia sp. Castellani, Jour. Trop. Med. Hyg. 28: 317-319, 1925. Monilia Castellanii Re. Jour. Trop. Med. Hyg. 28: 217-223, 1925. Cryptococcus Castellanii Castellani, Am. Jour. Trop. Med. 8: 390-392, 1929; Mallardo, Jour. Trop. Med. Hyg. 32: 145-147, 1929. Torulopsis Castellanii Castellani & Jacono, Jour. Trop. Med. Hyg. 36: 312-313, Figs. 38-39, 1933. Isolated from atypical cases of furunculosis not affected by staphylococcus vaccine. Cells gram-positive, not acid-fast, spherical 3-4;u, in diameter, occasionally a few hyphae on liquid media. No ascospores. Colony white then yellowish or brownish on glucose agar, similar with a reddish or purplish tinge on potato. No fermentation when first isolated, but after a few transplants ferments glucose and fructose ; also produces slight acidity in galactose. Milk not affected. Gelatin and serum not liquefied. Castellania dissocians (Mattlet) Dodge, n. comb. Monilia dissocians Mattlet, Ann. Soc. Beige Med. Trop. 6: 24-25, 1926. Isolated from a grayish white coating of the base of the tongue in a native of the Belgian Congo. In one ease the tonsils and pharynx were also involved. Microscopic examination of scrapings showed filaments 1.5-2 x 6-8/a. Treat- ment with KI.Io in glycerol cured the lesions slowly. After 3 days in potato water at 37° C. this shows a large variety of forms between the spherical cell and the hyphal form. Long cells about 20 x 2/i, EREMASCACEAE IMPERFECTAE 251 occasionally separated by spherical cells. By the thirtieth day these long- celled filaments are much like those of Casiellania Muhira. No chlamydospores. On Sabouraud agar, at 37° C., culture dull, white, circular. Later the center yellows with radial striations, margin lobed with hyphal tufts pene- trating the medium. On gelatin stab, appearance of inverted fir tree. In potato decoction, white flaky sediment, no pellicle. Fermentation with glu- cose and fructose, no fermentation of maltose, galactose, sucrose, lactose, man- nite, dextrin, or inulin. Litmus milk slightly acid. Gelatin not liquefied. Castellania epidermica (Ciferri & Alfonseca) Dodge, n. comb. Blastodendrion intesiinale var. epidermicum Ciferri & Alfonseca, Centralbl. Bakt. II, 83: 273-276, 1 pi., 1931. Isolated from a cutaneous mycosis of the face. Cells spherical in the center of the colony and on unfavorable media, 5fx in diameter, occasionally 6 x 4/t, giant cells 7-8/x. Somewhat branched pseudo- mycelium at the margins of the colony, cells elongate in short chains with thick cell walls (1-1. 5/i). On liquid media up to 20 cells in chains. No ascospores. Optimum temperature 36°-38° C. On Sabouraud agar (pH 6.2), colony white then yellowish, dense, creamy, thick, opaque with porcelain luster in refracted light, later almost cloudy, center level and uniform, margins thinner and grossly lobate, sometimes plumelike ; obscurely zonate in old cultures. On grape must gelatin (pH 5.6), similar but more zonate. On Difco malt agar (pH 6.8), similar, but growth more abundant and irregular. On Difco prune agar (pH 6.8), growth poor, transparent, zones not evident, margins irregular but nearly continuous. On potato and carrot, colony irregular, indefinite, spreading, growth rapid. On corn meal agar (pH 7.2), surface very brilliant. On Gorodkova agar (pH 6.8), growth poor, colonies small, almost transparent, later whitish and opaque. On peptone glucose broth (pH 6.4), growth poor, no pellicle, slight ring, sediment hyaline, agglomerated, grayish white, viscous, liquid darkens and thickens. On autolyzed yeast (pH 6.4) and malt solution (pH 6.0), growth very poor, no ring or pellicle, little sediment. Giant colony of Will's type I, no crater, periphery thin, transparent, slightly denticulate, odor of fermentation. No inversion of sugars, ferments glucose, fructose, trehalose, assimilates glucose, fructose, maltose, and sucrose, no assimilation of organic acids. Gelatin not liquefied. Castellania Mirandei (Velu) Dodge, n. comb. Cryptococcus Mirandei Velu, Bull. Soc. Path. Exot. 17: 545-547, 1924. Torulopsis Mirandei Almeida, Annaes Fac. Med. Sao Paulo 9: 10, 1933. Cryptococcus lacrymeatus Jeaume, Blastomycose des voies lacrymales de I'ane au Maroc. These Doc. Vet. Avignon, 1926. Found in tear ducts only. Differs from Zymonema farciminosum in spec- ificity for ass. Causes a pseudoinflammatory tumor characterized by a lym- phocyte and plasmatic infiltration without gigantocellular reaction, vasculari- zation, or sclerosis. 252 MEDICAL MYCOLOGY Obtained in pure culture on 5% citric acid agar. Grows well on Sabour- aud agar. Growth poor on potato and potato glycerol. On acid potato, colonies thick, creamy, confluent. Growth on carrot similar to that on acid potato but less. On beet, still less. Organism will grow in 1.5% NaOH glycerol peptone. In liquid media, there is abundant growth at bottom. In malt extract, a ring in 48 hours. In prune decoction, broth, or liver broth, Hay en, Hansen, or Cohn's solutions, growth is good: in carrot decoction, poor. Also poor in Pasteur, Laurent, or Nageli's solutions. Glucose and fructose fermented. Lactose, galactose, mannitol, sucrose, and maltose not fermented. Gelatin not liqueJSed. Castellania londinensis (Castellani & Chalmers) Dodge, n. comb. Monilm londinensis, Castellani & Chalmers, Man. Trop. Med. ed. 3, 1084, 1919. Mycelohlastanon londinense Ota, Jap. Jour. Derm. Urol. 28: [4], 1928. Isolated from a case of thrush, also found in the vagina. Colony white. Acid formation and fermentation with glucose and fruc- tose ; acid only with maltose, galactose, sucrose, and lactose. Milk acidified and clotted. Castellania Copellii (Neveu-Lemaire) Dodge, n. comb. Cryptococcus sp. Copelli, Giorn. Ital. Mai. Ven. Pelle 46: 467-492, 2 pis., 1912. Cryptococcus Copellii Neveu-Lemaire, Precis Parasitol. Hum. 79, 1921 ; Ota, Ann. Parasitol. Hum. Comp. 2: 51-53, Fig. 6, 1924. Hefe Copelli Sasakawa, Centralbl. Bakt. I, 88: 269-285, 1 pi., 1924. Mycelohlastanon Copellii Ota, Jap. Jour. Derai. Urol. 28: [4], 1928. Torulopsis Copellii Almeida, Annaes Fac. Med. Sao Paulo 9: 10, 1933. Referred to Cryptococcus Breweri by Vuillemin, Champ. Parasit. Homme Anim. 96, 1931. Isolated from lesions on the tongue. Pathogenic to rabbits and guinea pigs. Cells spherical, sparingly allantoid, 5-12/^ in diameter in tissues, 3-16yu, in cultures. Elongate cells in older cultures. Beginnings of hyphal formation was observed on certain media, but they Avere not studied long enough to obtain spores. Optimum temperature for growth 33°-38° C. ; no growth at 42°-45° C. Colonies small, white, rounded, elevated at center. On Sabouraud agar, colony circular, crateriform, yellowish white, moist, opaque. On potato, colony thick, an elevated plateau, with margins coarsely festooned, opaque, velvety. On beet, colony similar but rose color. On gelatin, colony shining, glassy white, round, elevated, margin regular. Growth strictly aerobic. On coagulated serum, colony thin, dry, powdery. On liquid media, no pellicle, slight turbidity, abundant sediment of tiny white flocci (glucose, mannite, maltose-peptone, wine, milk, Raulin's solution). Growth on urine or on horse EREMASCACEAE IMPERFECTAE 253 or rabbit broth shows no pellicle or turbidity but small white flocei. No indol formation on peptone. Glucose and fructose fermented. Milk coagulated in 10-15 days. Gelatin not liquefied. Castellania Negrii (Castellani) Dodge, n. comb. Endomyces Negrii Castellani, Brit. Med. Jour. 2: 1208-1212, 1912. Monilia Negrii Castellani & Chalmers, Man. Trop. Med. ed. 2, 822, 1913. Thin pellicle on broth (denied by Castellani & Chalmers, Man. Trop. Med. ed. 3, 1919). Acid and fermentation with glucose and fructose, slight with galactose, sucrose (1913, not 1919), and raffinose (1913, not 1919). Acid only with maltose, sucrose (1919), and lactose; slight with mannite and raffinose (1919). Castellania intestinalis (Castellani) Dodge, n. comb. Endomyces intestinalis Castellani, Brit. Med. Jour. 2: 1208-1212, 1912. Monilia intestinalis Castellani & Chalmers, Man. Trop. Med. ed. 2, 827, 1913. Mycelohlastanon intestiyiale. Ota, Jap. Jour. Derm. Urol. 28: [4], 1928. Isolated from stools and saliva. Acid formation and fermentation with glucose and fructose ; slightly acid with maltose ; acid with galactose and sucrose. Litmus milk first acidified, then decolorized. Castellania macedonensoides (Castellani & Taylor) Dodge, n. comb. Monilia macedonensoides Castellani & Taylor, Jour. Trop. Med. Hyg. 28: 244, 1925; Castellani, Douglas & Thompson, Jour. Trop. Med. Hyg. 28: 258, 1925. Found in vagina? Or bronchomycosis fide Zepponi, 1931. Morphology close to that of M. tropicalis fide Zepponi, 1931. Fermenta- tion of monosaccharides as in M. tropicalis, slight acidification with maltose and potato starch, decoloration of pea starch. Castellania Nabarroi (Castellani & Chalmers) Dodge, n. comb. Monilia Nabarroi Castellani & Chalmers, Man. Trop. Med. ed. 3, 1090, 1919. Mycelohlastanon Nabarroi Ota, Jap. Jour. Derm. Urol. 28: [4], 1928. Isolated from sputum. Colony white. Fermentation of glucose, fructose, and maltose. Milk coagulated. Gelatin not liquefied. Castellania decolorans (Castellani & Low) Dodge, n. comb. Monilia decolorans, Castellani & Low, Jour. Trop. Med. Hyg. 16: 33-35, 1913. Mycelohlastanon decolorans Ota, Jap. Jour. Derm. Urol. 28: [4], 1928. Isolated from scrapings of the tongue in sprue. Growth abundant, slightly acid on solid sugar media, colonies creamy white with smooth surface. Yeastlike forms with some hyphae in water of condensation. Poor growth in alkaline media, fairly good growth on gelatin. Acid and fermentation with glucose, fructose, and maltose. Acid only with galactose, sucrose, and dextrin. 254 MEDICAL MYCOLOGY Castellania faecalis (Castellani) Dodge, n. comb. Endomyces faecalis Castellani, Brit. Med. Jour. 2: 1208-1212, 1912. Monilia faecalis Castellani, Jour. Trop. Med. Hyg. 17: 307, 1914. MyceloUastanon faecale Ota, Jap. Jour. Derm. Urol. 28: [4], 1928. Isolated from feces. On coagulated serum, growth forms a brownish pigment which diffuses into the medium. Acid and fermentation with glucose, fructose, and maltose, slightly acid and fermentation with galactose and sucrose. Litmus milk turns acid, then is decolorized and slightly digested. Neither coagulated serum nor gelatin liquefied. Castellania bronchialis (Castellani) Dodge, n. comb. Endomyces bronchialis Castellani, Brit. Med. Jour. 2: 1208-1212, 1912. Mycelohlastanon bronchiole Ota, Jap. Jour. Derm. Urol. 28: [4], 1928, Candida bronchialis Basgal, Contr. Estudo Blastomycoses Pulmonares, 49, 1931. Isolated from sputum. Probably the organism of Smith & Sano (1933) from a case showing meningeal involvement should be referred here. Colonies white. Acid formation and fermentation with glucose, fructose, and maltose, slight with sucrose. Acid onlj^ with dextrin. No action on milk, coagulated serum, or gelatin. Castellania aegyptiaca (Khouri) Dodge, n. comb. Monilia uegyptiaca Khouri, Bull. Soc. Path. Exot. 26: 7-9, 1933. Isolated from sputum mixed with blood from case of pulmonary blasto- mycosis. No trace of Mycobacterium tuberculosis, no blood parasites, Was- sermann reaction negative. Organism slightly pathogenic for guinea pig. Spherical or ovoid cells with occasional hyphae observed in the sputum. Long hyphae with elongate-ovoid blastospores in cultures. No ascospores. On Sabouraud agar, colony creamy white, confluent, cells 3-8 average bp. in diameter, rarely sprouting. Optimum temperature 36°-37° C. Cells gram- positive. On carrot and potato, colonies creamy white. On liquid media, no pellicle, floccose sediment, no pigment. Litmus milk acidified, coagulated slowly after seventh day and curd slowly digested. Glucose, fructose, maltose, and sucrose fermented and acidified; galactose and raffinose also acidified. Neutral red reduced. Serum and gelatin not liquefied. Castellania Rogerii (Sartory & Demanche) Dodge, n. comb. Cryptococcus sp. Demanche & Sartory, C. R. Soc. Biol. 63: 261-262, 1907, Cryptococcus Rogerii Sartory & Demanche, Bull. Soc. Myc. France 23: 179-185, 1907. Torulopsis Rogerii Almeida, Annaes Fac. Med. Sao Paulo 9: 10, 1933. Isolated from pus in a case of peritonitis following perforation of the stomach. Cells elongate, 8-10 x 2-3/^, sprouting. Pseudomycelium present. No spores. Optimum temperature for growth 30°-35° C, no growth over 41° C, EREMASCACEAE IMPERFECTAE 255 Sucrose inverted. Maltose fermented, no action on galactose or lactose. Milk coagulated in 5 days, curd not digested. C. Bogerii var. (?), Beauverie & Lesieur, Jour. Phys. Path. Gen. 14: 992- 994, 1912, from pharyngeal exudate in case of typhoid, differs only in being nonpathogenic to rabbit. Strain IV of Spiethoff (1904), isolated from diabetic urine may belong here, but was nonpathogenic to laboratory animals. Castellania pulmonalis (Castellani) Dodge, n. comb. Endomyces pulmonalis Castellani, Arch, de Parasitol. 16: 184-186, 1913. Monilia 'pulmonalis Castellani & Chalmers, Man. Trop. Med. ed. 2, 828, 1913. Candida pulmonalis Basgal, Contr. Estudo Blastomycoses Pulmonares 49, 1931. Isolated from sputum and samples of tea. Thin pellicle formed on broth, brown pigment on coagulated serum. Acid formation and fermentation with glucose, fructose, maltose, sucrose, slight with galactose and arabinose, acid only with raffinose, very slight with mannite. Castellania burgessi (Castellani) Dodge, n. comb. Endomyces iurgessi Castellani, Arch, de Parasitol. 16: 184-186, 1913. Monilia hurgessi Castellani & Chalmers, Man. Trop. Med. ed. 2, 828, 1913. Isolated from the air. Colonies generally white and creamy, but black or brown pigment pro- duced on serum. Slight acid formation and fermentation with glucose, mal- tose, sucrose, acid only with fructose and galactose, no action on other sugars. No action on milk, serum, or gelatin. Castellania mannitofermentans (Castellani) Dodge, n. comb. Monilia mannitofermentans Castellani, Proc. Soc. Exp. Biol. Med. 26: 544- 545, 1929. Isolated from sputum in a case of chronic bronchitis. Gram-positive, acid-fast negative. Acid and fermentation with glucose, galactose, maltose, fructose, mannitol, and dextrin. Acid only with arabinose and xylose. Castellania pseudolondinensis (Castellani & Chalmers) Dodge, n. comb. Monilia pseudolondinensis Castellani & Chalmers, Man. Trop. Med. ed. 3, 1082, 1919. Isolated from sputum. Acid formation and fermentation of glucose, fructose, maltose, galactose, and dextrin. Litmus milk not clotted. Castellania pseudolondinoides (Castellani & Chalmers) Dodge, n. comb. Monilia pseudolondinoides Castellani & Chalmers, Man. Trop. Med. ed. 3, 1082, 1919. Differs from C. pseudolondinensis only in clotting litmus milk. Acid and fermentation with glucose, fructose, maltose, galactose, and dextrin. Castellania pseudometalondinensis (Castellani & Chalmers) Dodge, n. comb. 256 MEDICAL MYCOLOGY Monilia pseudonietalondifiensis Castellaiii & Chalmers, Man. Trop. Med. ed. 3, 1082, 1919. Acid and fermentation with glucose, fructose, maltose, galactose, and inulin. Litmus milk clotted. Castellania nivea (Castellani) Dodge, n. comb. Endomyccs niveus Castellani, Lancet 1: 15, 1912; Brit. Med. Jour. 2: 1208- 1212, 1912. Monilia nivea Castellani & Chalmers, Man. Trop. Med. ed. 2, 826, 1913. Mycelohlastanon niveum Ota, Jap. Jour. Derm. Urol. 28: [4], 1928. Candida 7iivea Basgal, Contr. Estudo Blastomycoses Pulmonares, 19, 1931. Reported to cause bronchomycosis, but isolated from sputum not collected in a sterile vessel, and therefore of doubtful pathogenicity. Acid formation and fermentation with glucose, fructose, maltose, galac- tose, and raffinose, slight with sucrose, Castellania metalondinensis (Castellani & Chalmers) Dodge, n. comb. Monilia metalondinensis Castellani & Chalmers, ]\Ian. Trop. Med. ed. 3, 1082, 1919. Mycelohlastanon metalondinense Ota, Jap. Jour. Dei-m. Urol. 28: [1], 1928. Candida metalondinensis Basgal, Contr. Estudo Blastomycoses Pulmonares, 49, 1931. Isolated in eases of thrush and from vaginal discharge. Spaar reports this organism in hard, milky growth on palate which was cured by a potassium chlorate mouth wash. On plain agar and glucose agar, growth is abundant and creamy white. Growth on gelatin fairly abundant, scanty on serunu Fermentation and acid formation with glucose, fructose, maltose, galactose ; none with other sugars. Litmus milk unchanged or slightl.y acid, no gas evolved. Neither coagulated serum nor gelatin liquefied. Castellania Richmondi (Shaw) Dodge, n. comb. Monilia Richmondi Shaw, Sci. 64: 300, 1926. From a case of clinical tuberculosis, with small hard whitish granules in the sputum. Pathogenic to guinea pigs intravenously but not intraperitone- ally, pathogenic to rabbits intraperitoneally. Colonies on dextrose agar, creamy white with a smooth surface, of yeast- like cells. In dextrose broth, both budding forms and mycelium. No pellicle on broth. Milk alkaline in 48 hours. Acid and fermentation with glucose, fructose, maltose, and galactose. No acid or fermentation with sucrose, lac- tose, mannite, duleite, raffinose, arabinose, adonite, dextrin, sorbite, or inulin. Neither gelatin nor coagulated serum liquefied. No indol produced. Castellania pseudogmllermondi (Castellani & Chalmers) Dodge, n. comb. Monilia psendoguillermondi Castellani & Chalmers, Man. Trop. Med. ed. 3, 1082, 1919. Isolated from sputum. EREMASCACEAE IM PERFECT AE 257 Acid formation and fermentation witli glucose, fructose, and sucrose. Milk clotted and digested. Gelatin not liquefied. Castellania Muhira (Mattlet) Dodge, n. comb. Monilia Muhira Mattlet, Ann. Soc. Beige Med. Trop. 6: 23-24, 1926. Isolated from scrapings of a grayish coating of tongue which caused pain ni swallowing and slight elevation of temperature each night. Patient a soldier in Belgian Congo. Treatment with glycerol Kl.Ig brought about com- plete cure. In scrapings, spherical or ovoid cells, 2-3/t, some sprouting. Growth for 8 days in potato decoction at 37° C, only spherical cells and elongate cells, occasionally arranged end to end, with either spherical or elongate cells at the articulations. Spherical cells 6-7/i, in diameter, elongate cells 12 x 2.5/^,. Typical liyphae to be seen in depths of gelatin cultures. These are formed of cells about 20 x 2-2.5ja, spherical or ovoid blastospores or other hyphae at the septa. Free ends of the cells are rounded. Protoplasm of long cells is reduced to a thin layer along the wall. No chlamydospores observed. Optimum temperature 37° C. On Sabouraud agar at 37°, the culture is dull, white, rounded. Later the center yellows with radial striations, margin lobed with outgrowths of large, contorted hyphae, extending down into medium. On gelatin stab, culture white, margin indented at surface. Horizontal hyphae grow out into depth of medium, giving a growth of inverted cone shape. On potato decoction, sediment of small white iflakes. Ferments glucose, fructose, sucrose, not mal- tose, galactose, lactose, mannite, dextrin, or inulin. No acid or coagulation with milk. Gelatin not liquefied. Perhaps this species should be referred to Blastodendrion, near B. Kayongosi. Ca^ellania Guilliermondi (Castellani) Dodge, n. comb. Endomyces Guilliermondi Castellani, Brit. Med. Jour. 2: 1208-1212, 1912. Monilia Guilliermondi Castellani & Chalmers, Man. Trop. Med. ed. 2, 826, 1913. Mycelohlastanon Guilliermondi Ota, Jap. Jour. Derm. Urol. 28: [4], 1928. Isolated from sputum in case of bronchomycosis. Thin pellicle on broth. Acid formation and fermentation with glucose, fructose, sucrose, slight with raffinose. Galactose becomes acid, maltose, slightly acid. Coagulated serum and gelatin not liquefied. Action on milk uncertain, some reporting coagulation which Aichelburg denies. Castellania Chalmersi (Castellani) Dodge, n. comb. Endomyces paratropicalis D Castellani, Centralbl. Bakt. I, 58: 236-238, 1911. Monilia Chalmersi Castellani & Chalmers, Man. Trop. Med. ed. 2, 826, 1913. Endomyces Chalmersi Castellani, Arch, de Parasitol. 16: 184-186, 1913. Mycelohlastanon Chalmersi Ota, Jap. Jour. Derm. Urol. 28: [4], 1928. Candida Chalmersi Basgal, Contr. Estudo Blastomycoses Pulmonares 49, 1931. 258 MEDICAL MYCOLOGY Isolated in case of bronchitis. Colonies Avhite. Acid formation and fermentation with glucose, fructose, and sucrose, slight with galactose and inulin. Slight acid formation only with maltose and raffinose. Milk turned slightly acid, then alkaline. Neither coagulated serum nor gelatin liquefied. Castellania Lustigi (Castellani & Chalmers) Dodge, n. comb. Monilia Lustigi Castellani & Chalmers, Man. Trop. Med. ed. 2, 828, 829, 1913. Isolated from samples of tea. Colonies snow white. On coagulated serum, a black pigment diffuses into the medium. Litmus milk slightly acid then decolorized. Slight acid forma- tion and fermentation with fructose, sucrose, raffinose. Acid with glucose, galactose, and dextrin, very slight with maltose. No liquefaction of coagu- lated serum or gelatin. Castellania metatropicalis (Castellani & Chalmers) Dodge, n. comb. Monilia metairopicalis Castellani & Chalmers, Man. Trop. Med. ed. 3, 1087, 1919. White colony on glucose agar. Acid formation and fermentation with glucose, fructose, maltose, galactose, and sucrose. Milk coagulated. Castellania insolita (Castellani) Dodge, n. comb. Endomyces insolitus Castellani, Brit. Med. Jour. 2: 1208-1212, 1912. Monilia insolita Castellani & Chalmers, Man. Trop. Med. ed. 2, 827, 1913. Mycelohlastanon iyisolitum Ota, Jap. Jour. Derm. Urol. 28: [4], 1928. Candida insolita Basgal, Contr. Estudo Blastomycoses Puliuonares, 49, 1931. Isolated from feces and saliva. In culture, forms a dark pigment which diffuses into the medium. Acid formation and fermentation Avitli glucose, fructose, maltose, galactose, sucrose. Slightly acid with mannite. ]Milk turns slightly acid, then alkaline. No lique- faction of coagulated serum or gelatin. Castellania tropicalis (Castellani) Dodge, n. comb. Oidium tropicals Castellani, Philippine Jour. Sci. 1> 5: 197-202, 1910; Brit. Med. Jour. 1910 2: 686-689, 1910. Endomyces tropicalis Castellani, Centralbl. Bakt. 1, 58: 236-238, 1911. Monilia tropicalis Castellani & Chalmers. Man. Trop. ]\Ied. ed. 2, 824-825, 1913. Atelosaccharomyces tropicalis Froilano de Mello & Gonzaga Fernandes, Arq. Ilig. Pat. Exot. 6: 263-268, 1918. MyceloMastanon tropicale Ota, Jap. Jour. Derm. Urol. 28: [4], 1928. Candida tropicalis Basgal, Contr. Estudo Blastomycoses Pulmonares 49, 1931. Isolated from sputum in 4 cases of coughing, 2 of the patients having been exposed to tea dust. Three cured by medication with potassium iodide. Tu- berculosis absent. Later same organism isolated in cases of tonsillomycoses and thrush. EREMASCACEAE IMPERFECTAE 259 Mycelium 3-4/* in diameter, bearing 4 shorter cells at either end. Blasto- spores ovoid or spherical 4-8/x in diameter. Gram-positive. Growth on agars abundant, thick, roundish, creamy, white. Acid forma- tion and fermentation with glucose, fructose, maltose. In the cases derived from tea dust, galactose and sucrose also fermented. No action on milk. Colonies large, elevated, round, creamy, Avhite, cells spherical, Gram-posi- tive, not acid-fast. Mycelium formed in peptone water. Acid formation and fermentation with glucose, maltose, and sucrose in 24 hours, with galactose and fructose in 5-6 days — Reimann, 1931. Zepponi (1931) reports fermentation and acidification with xylose, slight acidity with arabinose, no change in alcohols, polysaccharides or giucosides of hydroaromatic compounds. Castellania macedonensis (Castellani & Chalmers) Dodge, n. comb. Monilia macedonensis Castellani & Chalmers, Man. Trop. Med. ed. 3, 1087, 1919. Mycelohlasianon macedonense Ota, Jap. Jour. Derm. Urol. 28: [4], 1928. Found in sputum. Colony thick, creamy, whitish, cells ovoid to spherical, solitary or in chains, 3-6)U. in diameter, gram-positive. Sediment produced in broth, no turbidity. [Sanfilippo 1924.] Acid formation and fermentation of glucose, fructose, galactose, sucrose, and inulin. Milk coagulated. Gelatin and serum not liquefied. Castellania Valeriana Dodge, n. sp. Oidium alhicans var. Galli-Valerio, Arch, de Parasitol. 1: 572-582, 13 figs., 1898. Isolated from stools of infant suffering from chronic gasteroenteritis. Virulence slight for laboratory animals but is augmented when inoculated along with Bacterium coli. Mycelium on ascitic gelatin with terminal chlamydospores after one month. On agar, colonies white, shining, pinhead size. On maltose, growth yel- lowish white and of a yeasty odor. On gelatin, small, subspheric, white colo- nies with small, granular colonies along stab. Glucose, maltose, and lactose fermented. Milk coagulated in 5 days and coagulum digested in 3 months. Gelatin not liquefied. Castellania pseudotropicalis (Castellani) Dodge, n. comb. Endomyces pseudotropicalis Castellani, Centralbl. Bakt. I, 58 : 236-238, 1910. Monilia pseudotropicalis Castellani & Chalmers, Man. Trop. Med. ed. 2, 825, 1913. Atelosaccharomyces pseudotropicalis Froilano de Mello & Gonzaga Fer- nandes, Arq. Hig. Pat. Exot. 6: 264, 268, 1918. Mycelohlastanon pseudotropicalis Ota, Jap. Jour. Derm. Urol. 28: [4], 1928. Candida pseudotropicalis Basgal, Contr. Estudo Blastomycoses Pulmonares, 49, 1931. 260 MEDICAL MYCOLOGY Four numbered strains producing bronchomycosis were described by Castellani, Practitioner 124: 69, 1930. Acid formation and fermentation of glucose, fructose, sucrose, lactose ; slight acid formation and fermentation with galactose. Milk coagulated. Gelatin not liquefied. Castellania Kartulisi (Castellani) Dodge, n. comb. " Saccharomyzetaceen" Kartulis, Zeitschr. Hyg. 64: 285-304, 1909. Levure de Kartulis Sasakawa, Centralbl. Bakt. I, 88: 272, 1922; Ota, Ann. Parasitol. Hum. Comp. 2: 55-57, Fig. 11, 1924. Cryptococcus Kartulisi Castellani, Am. Jour. Trop. Med. 8: 413, 1928. MyceloMastanon Kartulis Ota, Jap. Jour. Derm. Urol. 28: [4], 1928. Isolated from chronic fistulas of gluteal region in one hundred cases, all men, mostly Egyptians between forty and sixty, in Alexandria. Three stages of growth observed; knot formation, softening of knots, fistula formation. Pathogenicity to gray mice, rabbits, and guinea pigs unproved. Cells ovoid and allantoid or longer, up to 18;u. in diameter, in chains. Spores (?) 3-4 per ascus, lO/x in diameter. Allantoid cells have enlarged ends (with 12 spores?). Chlamydospores in old cultures. Figures of spores not convincing, although author asserts that they differ from oil droplets. Growth good on sugar agar and potato, not so good on glycerol and peptone agar. Colonies on potato, white to ash-gray, shining. In beef broth there appears a slimy sediment, but no surface growth. Glucose and lactose slowly fermented, sucrose rapidly, with the formation of large bubbles of gas. Castellania Hashimotoi, Dodge, n. sp. Hefe Hashimoto, Jap. Zeitschr. Derm. Urol. 22: 1-34, PI. 1 [1-2], 1922. Lesions on the face in front of the ear (2 cases). Pathogenic for mice and guinea pigs. Yeast cells spherical or ovoid on solid media, more elongate in the pellicle of malt extract. Endospores appear in 20 hours on gypsum block at 22° C Optimum 23°-25° C. Colonies gray white at first, becoming grayish yellow or brownish with shining surface and elevated border, with radiating margins. Fermentation of lactose, maltose, glucose, not sucrose and galactose. The figures are so poorly reproduced that they give little information, but one is inclined to doubt the presence of true ascospores. Castellania linguae-pilosae (Lucet) Dodge, n. comb. Saccharomyces linguae-pilosae Lucet, Arch, de Parasitol. 4: 262-287, 1901. Cryptococcus linguae-pilosae Castellani & Chalmers, Man. Trop. Med. ed. 2, 770, 1913. Monilia sp. Catanei, Arch. Inst. Pasteur Algerie 3: 386-388, 1925. MyceloMastanon linguae-pilosae Ota, Jap. Jour. Derm. Urol. 28: [4], 1928. Torulopsis linguae-pilosae Almeida, Annaes Fac. Med Sao Paulo 9 : 10, 1933. Isolated from black pilose tongue with hypertrophy of papillae and dark discoloration. EREMASCACEAE IMPERFECTAE 261 Pathogenic for mice but not for larger laboratory animals except by injection, which reproduced lesion on tongue of rabbit. Cells spherical, 4-8/* in diameter, or elongate, 12-17 x 6/x, sometimes form- ing chains of up to 10 cells. Spores perhaps present, but not definitely demon- strated. Gueguen reports that, in old cultures, cells are 4-5/a in diameter, with small ovoid protrusions attached to the mother cell by fine pedicels. Some cells have no more than 4-5 such protrusions, while the remainder of the cell content divides into 5 or 6 unequal masses grouped at the periphery, each mass surrounded by its own wall. Best growth at 25°-35° C, growth slow above 40° C., ceases at 42° C. On glucose agar, Sabouraud agar, or glucose gelatin, growth white, creamy, shining, or slightly dull, tomentose. Colonies on potato thin, yellow- ish, brown, dry, dull, potato finally assuming same color as culture ; on potato glycerol, creamy, growth better; on carrot, colonies small, punctiform, white. On gelatin, small yellowish white colonies with arborizations in stabs. Growth slow on meat broths in the absence of glycerol or glucose, very good in malt extract, juice of apples, pears, or grapes, with slight pellicle and ring and a turbidity which settles, leaving a thickened gray pellicle and sediment. Glucose, fructose, galactose, maltose, sucrose, and dextrin fermented. No action with mannite or lactose. No action on milk. Gelatin and coagulated serum not liquefied. Castellania parakrusei (Castellani & Chalmers) Dodge, n. comb. Monilia parakrusei Castellani & Chalmers, Man. Trop. Med. ed. 2, 810, 1913. Mycelohlastanon parakrusei Ota, Jap. Jour. Derm. Urol. 28: [4], 1928. Isolated from sputum. Acid and fermentation with glucose and fructose. No action on other sugars. Milk coagulated. Pellicle on broth. Castellania nitida (Castellani) Dodge, n. comb. Endomyces pseudotropicalis C Castellani, Centralbl. Bakt. I, 58: 236-238, 1911, non al. Endomyces nitidus Castellani, Brit. Med. Jour. 2: 1208-1212, 1912. Monilia nitida Castellani & Chalmers, Man. Trop. Med. ed. 2, 826, 1913. Mycelohlastanon nitidum Ota, Jap. Jour. Derm. Urol. 28: [4], 1928. Candida nitida Basgal, Contr. Estudo Blastomycoses Pulmonares, 50, 1931. Isolated from sputum in ease of bronchomycosis. Colony white, shining, thin pellicle on broth. Acid and fermentation Avith glucose and fructose ; acid only with maltose, galactose, sucrose, lactose, man- nite ; slight acid with dextrin and raffinose. Milk coagulated. Castellania Orticoni Dodge, n. sp. Cryptococcus sp. Sartory & Orticoni, Rev. Path. Comp. 14: 174-176, 1914. Isolated from grayish superficial lesions in the mouth. Pathogenic to rabbit and guinea pig. Cells ovoid, 5-11 x 4/a, sprouting. Optimum temperature for growth on carrot 22°-30° C, no growth above 39° C. On peptoglycerol broth, optimum temperature is 26°-28° C. 262 MEDICAL MYCOLOGY On peptoglycerol broth, pellicle is formed with pseudomycelium and a sediment of spherical or ovoid cells. No aseospores seen on gypsum block. Sucrose inverted. Glucose and fructose fermented, but not maltose, lactose, or galactose. Almost no growth and no action on coagulated serum. Gelatin not liquefied. Castellania africana (Macfie) Dodge, n. comb. Monilia africana Macfie, Ann. Trop. Med. Parasitol. 15: 275-277, 1921. Isolated from feces in a case of obstinate diarrhea. Patient's tongue red, irregularly fissured, indented by teeth and partly eroded. Throat red. Feces canary yellow and frothy. Castellania abundant. Entamoeba coli and Blasto- cystis enterocola also present. Cells ovoid and somewhat elongate, 4.5 x 1.6/^ [average of measurements of 10 cells] . On solid media, yeastlike cells predominate with a few branched, septate hyphae. In liquid media, hyphae most common. Gram-positive. Not acid-fast. On glucose agar, after 24 hours, white, very fluid growth with dull, pel- licle-like surface. Growth slower and less abundant under anaerobic condi- tions. On potato, growth is dull and white with a whitish efflorescence appear- ing later. In broth and peptone water, sediment whitish, medium clear. Surface pellicle in broth only. Glucose, fructose, and dextrin fermented. Gelatin not liquefied. Castellania platensis (Peruchena) Dodge, n. comb. Monilia platensis Peruchena, Semana Med. 36: [1-31] 10 jigs., 1929. Isolated from a patient who suffered from cough, expectoration, loss of strength and weight. Lung involved. Pathogenic for rabbits. Yeast cells predominate at first, finally hyphae ; septate, elongate cells, 15-20 X 2-3/x up to 50 x 4-5/i,. No asci observed. Cells rounded or ovoid, 4-6/x in diameter, increase by sprouting. Gram-positive. On simple agar at 38°, small, rounded, isolated colonies, 2-4 mm., slightly elevated in the center, finely granulose at the periphery, dull white, becoming confluent. On litmus lactose agar, abundant development with a slight nile green color in the substratum. On glucose neutral red agar, good develop- ment, colonies take dye, no color change. On Sabouraud glucose agar, colo- nies dull white, 1-2 mm. Endo medium red with colonies taking dye. On gelatin stab, 18 small colonies, 1-2 mm. in diameter. No pigmentation, colonies superficial. Broth shows no turbidity or odor, milky white sediment formed, also sometimes there is a weak pellicle from floating islets. In bile broth, no turbidity, but there is a precipitate. No indol formation. Ferments fructose and maltose. Acid formation strong with galactose and arabinose, slight with fructose, maltose, and mannose. Glucose not fermented. On coagulated human sernm, there is slight development, no liquefaction or pigmentation. Milk slightly acidified without coagulation. Gelatin not liquefied. Castellania alba (Castellani & Chalmers) Dodge, n. comb. Monilia alha Castellani & Chalmers, Man. Trop. Med. ed. 3, 1089, 1919. EREMASCACEAE IMPERFECTAE 263 Source of organism and pathogenicity not mentioned. Colony white on ghicose agar. Thin pellicle on broth. Acid formation and fermentation with glucose, maltose, and galactose, acid only with sucrose. Milk coagulated. Neither coagulated serum nor gelatin liquefied. Perhaps the organism described by Porgues should be referred to this species. Below is a description based on Forgues' thesis: Parendomyces sp. Forgues, These de Bordeaux 87: 1-100, 1913. Monilia de Forgues Sartorj^ Champ. Parasit. Homme Anim., 710-711, 1922. Isolated from the exudate in the throat and from the sputum of a patient suffering from angina and pleuropneumonia. The soft palate, tonsils, pillars of fauces, and posterior surface of the pharynx Avere all covered with the whitish coating. Pulmonary symptoms appeared later. Fatal to white mice both by hypodermic and intraperitoneal injection. Organism recoverable. Patho- genic, though not always fatal, to guinea pigs and rabbits. Yeast cells spherical, except where flattened by pressure, in groups of 10-20. Dimensions vary with medium, the diameter being 2-4/x in serum gelatin and 6-8;u in Kaulin's liquid. On 15- to 20-day-old cultures on carrot, di- ameter sometimes as large as 15-20^. Protoplasm surrounded by a cell wall and showed metachromatic granules and vacuoles. liypliae appear on older, solid cultures, measuring l-2fji. in diameter and 80-lOOya in length. At intervals of 30-40/i, there is a gap of 2-3/a in length, which is not colored by the dyes which stain the remainder of the protoplasm. Hyphae rarely branch but yeast cells may be attached anywhere. Pseudomj^celium present, 5-20 x 5/*. Reproduction by sprouting. On agar, growth very poor, very tiny colonies. On malt agar, colony abundant, not folded, with regular pointed projections from border. On Sabouraud medium, uniform colonies, irregularly folded, often mammillate or cracked, rarely reaching the Myalls of the tube. On potato and glycerol, colony abundant in 48 hours, grayish white rapidly developing, thick, uniform with occasional elevations, covering whole carrot surface in a few days. Grayish white sediment at the bottom of the tube, with some membranous clots above. On carrot and glycerol and on carrot and 5% glucose in 12 hours grayish white, creamy colonies which are confluent in 48 hours, covering whole surface of carrot with thick coating. In 20 days, culture has dried and cracked transversely and in circles, powdery whitish sediment at bottom of the tube. On gelatin, s^all punctiform colonies in 15-20 days. On stab, growth similar both at surface and in depths. Colonies similar but confluent, on serum gelatin Avith simple syrup. Growth similar on serum gelatin with gly- cerol, acid, or alkali. In plain broth, slight sediment in 48 hours, liquid clouds with formation of clots Avhich sink to bottom, no pellicle, liquid clears in 20 daj^s. GroAvth similar in broth Avith tartaric acid, 5% glycerol, or 5% glucose. In 20% glucose broth, a slight pellicle appears to form, but this is composed of clots Avhich finally sink to the bottom. GroAvth similar in Raulin's medium. GroAvth absent or negligible in the presence of peptone. 264 MEDICAL, MYCOLOGY Acidification and fermentation with glucose, fructose, galactose, maltose, lactose, and dextrin; none with sucrose, starch, inulin, glycerol or mannite. Coagulated serum and gelatin not liquefied. Castellania accraensis (Macfie & Ingram) Dodge, n. comb. Monilia accraensis Macfie & Ingram, Ann. Trop. Med. Parasitol. 15: 274, 1921, non al. Monilia nivea ? Macfie & Ingram. Ann. Trop. Med. Parasitol. 15: 53-58, 1921, non al. Candida acraensis Basgal. Contr. Estudo Blastomycoses Pulmonares 49, 1931. A complicating organism in fatal cases of tuberculosis of the lungs. Colonies diffuse and creamy white on most media. Thick white colony on potato. White deposit in broth and peptone water, slight pellicle on pep- tone water. Hyphae only on liquid media. Acid formation and fermentation with glucose, galactose, and sucrose, slight also with maltose, none with other sugars. Coagulated serum and gelatin not liquefied. Differs from Castellania nivea by not fermenting raffinose. Castellania paratropicalis (Castellani) Dodge, n. comb. Endomyces paratropicalis Castellani, Centralbl. Bakt. I, 58: 236-238, 1911. Monilia paratropicalis Castellani, Jour. Trop. Med. Hyg. 17: 307, 1914. Atelosaccharomyces paratropicalis Froilano de Mello & Gonzaga Fernandes, Arq. Hig. Pat. Exot. 6: 264-268, 1918. Mycelohlastanon paratropicale Ota, Jap. Jour. Derm. Urol. 28: [4], 1928. Candida paratropicalis Basgal, Contr. Estudo Blastomycoses Pulmonares, 49, 1931. Isolated from case of bronchomycosis, also from tAvo cases of blastomycetic dermatitis in Ceylon. Thin pellicle formed on broth. Acid formation and fermentation with glucose, fructose, galactose, sucrose, and maltose. Very slight acid with dextrin. Milk not coagulated. Castellania enterica (Castellani) Dodge, n. comb. Endomyces enterica Castellani, Brit. Med. Jour. 2: 1208-1212, 1912. Monilia enterica Castellani & Chalmers, Man. Trop. Med. ed. 2, 827, 1913 ; Delamare, Bull. Mem. Soc. Hop. Paris 43: 572-575, 1919. Mycelohlastanon entericum Ota, Jap. Jour. Derm. Urol. 28: [4], 1928. Candida enterica Almeida, Annaes Fac. Med. Sao Paulo 9: 11, 1933. Isolated by Delamare from the stools of a patient with chronic diarrhea in Senegal. Produced lesions in rabbit ear, slight symptoms in guinea pig and, administered per os, produced typical diarrhea in cat. On glucose and lactose agars, growth is rapid, and white granular colony forms. On maltose, mannite, and sucrose agars, growth is in projecting tufts, umbilicate, surrounded by a striate zone and by an almost transparent aureole. Ivory tint on ageing. On fructose agar, meager growth. On stab in ordinary agar, colonies are bright, white, rounded, elevated with umbilicus EREMASCACEAE IMPERFECTAE 265 or central papilla. Growth stationary after fifth day. On neutral red agar, colonies are tinted rose, agar yellowing. On potato, there is a white, dry growth with elevations like the contours of a map, also erratic round colonies of pinhead size. On carrot, growth is a white folded membrane with festooned edge. Water of condensation is milky. On beet, growth is white with a rosy tint, bristling with little papillae, border festooned and projecting. On arti- choke, colonies are verrucose, cafe-au-lait in color on ageing. Substrate turns green on sixth day, black on the eighteenth day. On gelatin, surface growth is slow and in the form of little white nodules. In the depths, growth appears arborescent. On gelatin with glycerol, glucose and liver infusion growth is meager. On coagulated serum, colonies punctiform. In hay infusion, a slight pellicle forms Avith whitish flakes settling out and leaving the medium clear. In broth, there is a white deposit at the bottom of the tube with slight tem- porary cloudiness. In bichromate broth, as above, with cloudiness persistent. Acid formation and fermentation with glucose, fructose, maltose, galactose, sucrose ; slightly acid with mannite and dextrin. Litmus milk turns slightly alkaline at first, then is decolorized below. No coagulation. No liquefaction or discoloration of coagulated serum. No liquefaction of gelatin. PARASACCHAROIVIYOES Parasaccharomyces Beurmann & Gougerot, Tribune Med. 42: 502, 1909. The type species is Parasaccharomyces Samlergeri Beurmann & Gougerot based upon Pseudosaccharomyces Busse Samberger. Colony creamy, hyphae straight, long; yeast cells ellipsoid, thick-walled; ascospores not seen. No pellicle produced on liquid media, sometimes a ring with aerial hyphae ; gelatin liquefied, sugars fermented. This genus, originally described from ulcers, has since been reported from onychomycosis and from the respiratory and alimentary tracts. Key to Species Only glucose and fructose fermented. Colony bluish gray, from subcutaneous ulcers. P. Sambergeri. Colony yellowish or grayish brown, from onychomycosis. P. oosporoides. Other sugars fermented. Maltose not fermented, from sputum. P. parachalmersi. Maltose fermented. Sucrose not fermented. Milk coagulated, from sputum. P. Colardi. Milk acidified but not coagulated, from feces. P. intestinalis. Sucrose fermented. Galactose fermented, from sputum. Milk not coagulated. P. irritans. Milk coagulated. p. Talicei. Galactose not fermented. p. crater if or mis. 266 MEDICAL MYCOLOGY Parasaccharomyces Sambergeri Beurmaim & Gougerot, Tribune Med. 42: 502, 1909. Pseudosaccharomyces Busse Samberger, ISbornik Klinicky 5: 466-485, PI. 6, 1904. Isolated from a lesion which started as a vesicle, then a pustule on the left nostril accompanied by pruritus and spread by scratching. The central area a scar covered with fine scales, caused by the peeling of the horny layer. Around this scar is an ulcerous area with the margin elevated, light red. In this area are small pustules, and the crust is thick, uniform, confluent, cover- ing the whole. Pathogenic for mice. In pus, cells spherical or ovoid, thick-walled, yellowish, often in pairs, occasionally showing sprouting. In cultures, cells ellipsoid, thick-walled, with long straight or slightly curved hyphae, cells 10-12/i,. On Zopf medium [water 1,000 e.c, ammonium tartrate 10 gm., dihydrogen potassium phosphate 5 gm., magnesium sulphate 2.5 gm., calcium phosphate 0.5 gm., peptone 10 gm., sucrose 140 gm.], colony thick, bluish gray, margin shining, even, opaque. On potato, surface rough, verrucose, gray or tan gray. In gelatin stab, colony like a large nailhead with only slight growth along the stab. On liquid media, a ring with aerial hyphae in 3 weeks. Gelatin liquefied in 3 weeks ; glucose fermented. Parasaccharomyces oosporoides (Zach) Dodge, n. comb. Blastodendrion oosporoides Zach in Wolfram & Zach, Arch. Derm. Syphilis 169: 102, 103, 1933. Isolated from a case of onychomycosis. Perhaps the organism of Mackin- non (1934), Geotrichoides strain 464, belongs here. Cells ovoid, rarely spherical, thick-walled, with a large vacuole and 1-2 oil globules, 6.5/x in diameter, ovoid cells 5-6.5 x 7.5-9.7/1.. Sprouting at both ends or irregular, after 3 months forming long branched hyphae with occa- sional blastospores at the septa. Hyphal cells elongate, ends rounded, blasto- spores shorter ellipsoid, not lacrimiform. On agar, colony whitish, dull, surface and margin smooth, later becoming yellowish and margin less regular and wavy. On gelatin, colony grayish, dull, margin smooth at first, later wavy, sinking into the gelatin. On gelatin, stab growth superficial with some colonies along the stab. On potato, colony thick, grayish yellow, dull. On carrot, colony white, moist. On maltose broth and autolyzed yeast floccose sediment, no cloudiness and no pellicle. Glucose and fructose fermented. Gelatin liquefied on the ninth day. The following two unnamed species of imperfect fungi may possibly be referred to P. oosporoides but present some minor differences. The first agrees rather closely with Zach's description, while the second seems deserving of varietal rank. It is to be hoped that a sufficient number of strains of this organism may be studied to give us more information on variation. This species should also be compared with Mycocandida onychophila (p. 294). [Monilia sp.] Diibendorfer, Derm. Centralbl. 7: 290-302, 3 figs., 1904. EREMASCACEAE IMPERFECTAE 267 Isolated from a case of onychomycosis. Pathogenic to laboratory animals. Cells long, ovoid or ellipsoid. Hyphae present. Colonies round, hemispheric, elevated with shiny porcelain-like surface. On malt agar, colonies large and flat. On potato, growth in small, hemispheric colonies sending out radiations Avhich join them in monilifonn groups. Growth white, becoming gray in 1-2 months. On alkaline gelatin stab, growth in form of lateral small foliiform projections into medium. In broth and malt extract, a flocculent sediment without turbidity, no pellicle. Glucose fermented, mal- tose and sucrose not. Alkaline gelatin not liquefied but malt gelatin finally liquefied. Var. Gug'gfenheimi Dodge, n. var. Oidium sp. Guggenheim, Arch. Derm. Syphilis 142: 305-309, 1923. Produces onychomycosis. Human inoculation positive. Intravenous in- jection in guinea pig fatal in 10 days. Colonies gray brown, smooth, shining, round elevations, after 2 weeks becoming silvery powdery, then gradually resuming gray broAvn color. No growth on malt agar. In beef broth, gray sediment appears but no turbidity. Glucose fermented, gelatin liquefied. Parasaccharomyces parachalmersi (Castellani & Chalmers) Dodge, n. comb. Monilia parachalmersi Castellani & Chalmers, Man. Trop. Med. ed. 3, 1087, 1919. Mycelohlastanon parachalmersi Ota, Jap. Jour. Derm. Urol. 28: [4], 1928. Isolated, from sputum. Acid formation and fermentation with glucose, fructose, galactose, sucrose, and inulin. Milk coagulated. Gelatin slowly liquefied. Parasaccharomyces Colardi Dodge, n. sp. Monilia sp. Jaumain & Colard, C. R. Soc. Biol. 93: 858-860, 1925; Colard & Jaumain, Bruxelles Med. 5: 1503-1506, 1925. Isolated from the sputum in a case of bronchomycosis (fatal) in the Belgian Congo which was under observation in 1924-1925. Shortly before death, the same organism was isolated from ulcers in mouth and pharynx. Pathogenic for rabbits. In sputum, cells 4-5 x 2-3/a, granular, often united in pairs. Yeast cells and branched hyphae bearing terminal chlamydospores. On solid sugar media, mostly yeast cells or occasional short chains of long cells forming pseudo- mycelium. Optimum temperature 37° C. On solid sugar media, colony thick, creamy white, not changing color with age. On can'ot sediment, some floating flakes in the liquid, no pellicle or ring. Abundant sediment in malt extract. Acid and fermentation with glucose, fructose, maltose, and dextrin. Acid only with galactose and sucrose. No action with lactose, mannite, inulin, or starch. Litmus milk acidified, coagu- lated after 8 days in the incubator when it again becomes alkaline. No action on coagulated serum. Glucose gelatin liquefied in 10 days. 268 MEDICAL MYCOLOGY Parasaccharomyces intestinalis (Mattlet) Dodge, n. comb. Blast ode ndrion mtestinale Mattlet, Ann. Soc. Beige Med. Trop. 6: 16-17, 1926. One of the organisms isolated from stools of patients suffering from dysentery in Belgian Congo. In potato decoction at 37° after 3 days, yeast cells ovoid, 2.5-3.5 x 5-8/i, vacuolate, sprouting to give spherical cells, 2-3/t in diameter which soon elongate ; hyphae, simple or slightly branched, composed of 12-20 cells with small area of contact. After 30 days the yeast cells contain fat droplets and some are thick-walled, spherical, 7-8/a in diameter with a crown of granula- tions as in Atelosaccharomyces pyogenes. The hj^phae become firmer, not easily dissociated, with larger areas of contact between cells. On Sabouraud agar at 37° C, colony round, white, dull. Later the center yellows and crumples, with radial folds at the lobate margin and verrucose projections into the medium. On gelatin stab, colony develops similarly with abundant granulations, as large as 1 mm. in diameter, along stab. In potato decoction, white lumpy sediment. Optimum for culture, 37° C. Milk slightly acidified. Fermentation of glucose, fructose, maltose, galactose ; none of sucrose, lactose, mannite, dextrin, inulin. Gelatin liquefied with evolution of gas. Parasaccharomyces irritans (Mattlet) Dodge, n. comb. Blast odendrion irritans A Mattlet, Ann. Soc. Beige Med. Trop. 6: 18-19, 1926. Isolated from the sputum in several cases of mycosis of the respiratory tract with dry cough in long drawn-out fits of varying degrees of severity. Treatment with potassium iodide, emetine, and arsenicals gave improvement, although the parasite generally remained in the sputum. In sputum, yeast cells, spherical or ovoid, 2-5/a in diameter, sprouting or in groups. After 3 da3\s at 37° in potato decoction, single cells, ovoid or spherical, with numerous granulations and rather rare types of cellular group- ings. Spherical cells up to 6/* in diameter, ovoid cells 3.5 x 7/a. Groupings consist of an axis of elongate cells, 10 x Sfi, from whose septa grow either lateral filaments, or spherical cells, or chains of cells. Between the elongate cells sometimes grow intercalary spherical cells. Free ends of cells always rounded. Same after 30 days. Optimum temperature for growth 37° C. On Sabouraud agar, colony round, cream colored at the smooth margin with yellow center, surface smooth and slightly shining, the center wrinkling slightly in age, no submerged mycelium. On gelatin stab, small cup of lique- faction after 20 days. In depths numerous granulations along the stab, medium displaced. In potato decoction, sediment of grayish white clots which break up on agitation. Fermentation of glucose, fructose, maltose, galactose, sucrose, none of lactose, mannite, dextrin, inulin. Slight acid formation with milk but no coagulation. Slight gas with gelatin, also liquefaction. The following strain B described by Mattlet differs in minor particulars but scarcely seems deserving of varietal rank. EREMASCACEAE IMPERFECTAE 269 Blast odendrioii irritans B Mattlet, Ann. Soc. Beige Med. Trop. 6: 18-20, 1926. Monilia sp. (case V) Mattlet, Ann. Soc. Beige Med. Trop. 4: 173, 1925. Isolated from cases clinically similar to those of the species. In the case V Soter, Aspergillus giganteus was isolated by Mattlet at the same time as Blast odendrioii irritaris. After 3 days at 37° C. in potato water, many spherical cells, 6/x in diameter, and ovoid cells 7-8 x 4/t, with a large central vacuole. Rare elongate cells, 8 X 3/A, forming hyphae when attached end to end. At the septa, sprout single elongate or spherical cells. After 30 days all cells show fatty granulations. On Sabouraud agar, appearance as in P. irritans A, except margin lobate and tufts of hyphae penetrating deeply into substrate. Action on sugars differs from P. irritans A only in intensity. Litmus milk at first slightly acid, then alkaline. Gelatin liquefied, gas evolved. Parasaccharomyces Talicei Dodge, n. sp. Monilia sp. Talice & Mackinnon, Bol. Inst. Clin. Quiriirg. Univ. Buenos Aires 4: 502-519, 13 figs., 1928. Isolated from the sputum of a case simulating pulmonary tuberculosis without showing presence of Mycobacterium. Pathogenic to rabbits on in- travenous injection, to white rat on intraperitoneal injection, but not found pathogenic to guinea pig. Rare hyphae in sputum. Blastospores present. In culture, blastospores of variable size, spherical, 2-11/^ in diameter, averaging 5-6/t, ovoid 5-7/a in diameter, sometimes forming chains in cultures more than 3 days old. Blasto- spores germinate by septate hyphae, 10-30 x 2/t, which in turn give rise terminally and laterally to hyphae and blastospores. Hyphae increase in number as the culture ages, but yeast forms always predominate. In very old cultures the hyphae form arthrospores. Protoplasm generally granular and shows oil droplets. Gram-positive, stained with May-Giemsa stain. Optimum temperature for growth 36°-38° C, grows anywhere between 18° and 40° C., does not grow above 45° C. Giant colony, after 16 days, is 4-5 cm. in diameter, approximately circular, granular at the center, with radiating furrows which end in indentations at the margin, generally white but assumes color of the medium. On plain or Gorodkova agar, a moist and white covering. On potato, colonies of irregular outline, verrucose. On caiTot, surface colonies which grow rapidly and be- come folded. On Sabouraud glucose agar, growth very good, forming a moist thick covering which acquires the color of the medium. On ordinary Sabour- aud agar or Sabouraud with maltose, groAvth similar but less abundant. On Sabouraud conservation agar, growth poor, verrucose. In glucose broth, a very light pellicle which clings to sides of tube. Similar on Raulin's liquid. Acidification and fermentation with glucose, maltose, galactose, sucrose, inulin. 270 MEDICAL MYCOLOGY and levulose, none with lactose, mamiite and dulcite. Litmus milk acidified and coagulated, then alkaline. Slight growth on coagulated serum. Gelatin rapidly liquefied. Parasaccharomyces crateriformis Dodge, n. sp. Oidium sp. Spiethoff, Jahrb. Hamburg. Staatskrankenanst. 9: 167-208, Ph. 14, 15, 1905. Isolated from lesions about the genitalia and adjacent thigh, also from the urine of a diabetic woman. Pathogenic for laboratory animals. Cells spherical to ovoid with some filaments. Colonies on agar, glucose agar, malt agar, and blood agar, white, moist, without aerial hyphae ; on blood serum, growth poorer ; on gelatin, colony thick, white, crateriform, margin with short radial furrows. On potato, colony at first white, moist, becoming dry and chalky. In liquid media, a ring but no pellicle developed, with a fine flocculent sediment. Coagulation of milk usually negative but observed after several days in a few tubes. Glucose, sucrose, and maltose fermented. Gelatin liquefied. MONILIA Monilia Bonorden. Handb. AUg. Mykol. 76, 1851. Candida Langeron & Talice, Ann. Parasitol. Hum. Comp. 10: 54-56, 1932. The type species is Monilia Candida Bonorden {Monilia Bonordeni Vuil- lemin) . Colonies creamy, thick, convex, beginning by bipolar sprouting of the blastospore and followed by extensive branching, blastospores appearing late. Blastospores ovoid, produced only terminally. Pseudomycelium of ellipsoid cells, terminal cells prolonged into simple or compound chains, no verticils (Fig. 49). Pellicle developing in liquid media, gelatin liquefied, sugars fer- mented. This group is predominantly saprophytic. It is quite possible that Myco- torida Will may belong here, but its morphology has been so poorly described that one cannot be certain. The description of M. Bonordeni is included here because pathogenic cultures have so often been incorrectly referred here. M. Kochi, inadequately described, should be recognized by its red color, Monilia Bonordeni Vuillemin, Bull. Soc. Myc. France 27: 140, 1911. Monilia Candida Bonorden, Handb. Allg. Mykol. 76, 1851, not Persoon, Syn. Meth. Fung., 693, 1801. Endomyces candidus Castellani, Lancet 1: 13-15, 1912. Mycelohlastanon candidum Ota, Jap. Jour. Derm. Urol. 28: [4], 1928. Candida Bonordeni Basgal, Contr. Estudo Blastomycoses Pulmonares, 49, 1931. Forms a woolly, granular, snow-white coA^ering on rotten wood, about 2 mm. thick. Not pathogenic. EREMASCACEAE IMPERFECTAE 271 Cells spherical to short ovoid. In older cultures and in pellicles also long sprouting- cells and very long, septate branched hypliae, bearing terminal chains of ovoid or spherical white blastospores. Blastospores sprouting. Giant colonies usually pure white, moist, shining, elevated. With age the surface becomes somewhat drier, with a depression in the center which ap- pears hairy. The periphery is correspondingly elevated. The gelatin adjacent to the surface colonies is somewhat depressed. Th^ colonies in the depths of the medium appear spherical or star-shaped. Culture lias a pleasant, sour Fig. 49. — Monllia (Candida Langeron & Talice). 1, chains of ellipsoid blastospores; 2, chains at tips of filaments ; 3, terminal chains and veiticlllate branching. (After Langeron & Talice 1932.) odor. On malt gelatin streak, growth is broad, thick, white, with deep folds starting parallel to the streak and ending perpendicular to the periphery. With age, colony becomes dry and powdery. Growth similar on stab though not much in depths of medium. In malt extract, beer, or whey, a pellicle is formed. In the latter at 25° C, abundant, flocculent growth. Ferments glucose, fructose, maltose, and sucrose. Malt gelatin slightly liquefied in 2 weeks, completely in 6-8 weeks. 272 MEDICAL MYCOLOGY Monilia Kochii (Wettstein) Saccardo, Syll. Fung. 10: 518, 1892. Rhodomyces Kochii Wettstein, Sitzungsber. K. Akad. Wiss. Wien, 91 : 33- 58, 1885. Rhodomyces eruhescens Apel apud Ascher, Zeitsclir. Hyg. Infektions- krankh. 34: 475-481, PI. 5, 1900. Candida Kochii Basgal, Contr. Estudo Blastomycoses Pulmonares, 49, 1931. Isolated from human sputum repeatedly over a period of 2 years. In the stomach of a cat fed with milk, it sprouted and formed conidia. Seems to be connected with pyrosis. R. ruhescens was isolated from the placenta and fetal skin of a guinea pig, but is not pathogenic for experimental animals. Mycelium in substrate colorless, thin, one- to several-celled. Hyphal cells 20-60 X 6-16/t with thin membrane. Conidiophores rise above substrate, rose red to yellowish red, formed of spherical or short cylindric cells, much branched. Conidia in chains which finally dissociate, ovoid to polj^hedral, 5-16ytt in di- ameter or 15-20 X 6-15/i, with relatively thin wall and hyaline content, finallj^ breaking up into a powdery mass. Intercalary chlamydospores also present. Hyphal anastomoses observed. Colonies rounded, rose-red or red-yellow, covered by a pulverulent conidial layer 1-2 mm. thick, aerobic. Sugars not fermented. Milk slowly coagulated and digested. No ring or pellicle on liquid media. The species of Oidium referred to as Oidium rose by Sartory & Orticoni, Rev. Path, Comp. 14: 176, 1914, but not fully described, perhaps belongs here. Hyphae repent, 3-4/a in diameter, rose color. Fertile hyphae terminated by a chain of ovoid, pale rose spores, 2.5-3 x 2fi. Not pathogenic but found along with pathogenic Cryptococcus sp. described in same paper. SYRINGOSPORA Syringospora Quinquaud, Arch. Physiol. Norm. Path. 1: 290-305, PI. 8, 1868. Wnantiothamnus Pinoy apud Brault & Masselot, Ann. Derm. Syphiligr. V, 2: 602, 1911. Mycotorula Langeron & Talice, Ann. Parasitol. Hum. Comp. 10: 43-48, 1932. The type species is Syringospora Robini Quinquaud (Oidium albicans Robin). Colonies creamy, thick and convex, beginning by polar sprouting of a blastospore, followed by the progressive branching of the pseudomycelium ; blastospores spherical or ovoid, rarely elongate, arranged in simple verticils at the septa along a hypha; pseudomycelium formed of short cells, each typically terminated by a verticil of blastospores; terminal cell similar, very rarely terminated by a short chain ; verticils simple, regularly spaced, some- times limited to 4 or 6, sometimes in dense clusters (Fig. 50). Gelatin lique- fied, sugars fermented. EREMASCACEAE IMPERFECTAE 273 With some hesitation has Enantiothamnus been referred here, on account of its morphology, although several characters suggest a relationship with Proteomyces. Biochemical characters which often provide a clue to the rela- tionships are not given in the original description. Langeron & Talice (1932) doubtfully referred it to their Geotrichoides (Candida Berkhout). The members of this genus grow predominantly on mucous membranes of the respiratory and alimentary tracts and about the external genitalia, Fig. 50. — Syringospora (Mycotoi-ula Langeron & TalicG) showing regular, dense verticils with- out terminal chains. (After Langeron & Talice 1932.) extending into the urethra in man. They seem to be saprophytes or mild parasites which grow much better on artificial media and produce injury largely by mechanical and perhaps chemical irritation rather than by invading the tissues. I have also referred here several partially described organisms found growing on the moist folds of skin, causing local irritation often clini- cally resembling those caused by Epidennophyion. 8. Braulii (Enantiotham- 7ius) alone in the group produces subcutaneous lesions. 274 MEDICAL MYCOLOGY Key to Species Pellicle formed on liquid media. Verticils dense, cells nearly spherical. S. albicans. Verticils scanty (4-6 cells), cells slightly elongate pointed at one end. S. Braulti. No pellicle on liquid media. Gelatin not liquefied. From moist folds of skin. Colony margin even. S. interdigitalis. Colony margin dendroid, sugars fermented. S. cutanea. From digestive tract. Maltose not fermented. S. Tonge. Maltose fermented. s. psilosis. Gelatin liquefied. Sugars not fermented, from cornea. S. Cavarae. Galactose and sucrose fermented. ^S^. Issavi. Galactose and sucrose not fermented. S. Negroni. S3nnngospora albicans (Robin) Dodge, n. comb ^porotrichum sp. Griiby, C. R. Acad. Sci. Paris 14: 634-G35, 1842. Oidium albicans Robin, Hist. Nat. Veg-. Paras. 488-513, 1853. Stemphylium polymorphum Hallier, Die Pflanzl. Paras, des Menschl. Korpers, 1866. Syringospora Rohini Quinquand, Arch. Pliysiol. Norm. Path. 1: 290-305, PI. 8, 1868. Saccliaromyces albicans Reess, Sitzungsber. Phys. Med. Soc. Erlangen 9: 190-195, 1877. Mycoderma vini Grawitz, Deiitsch. Zeitschr. Prakt. Med. 1877: 209-211, 220-223, 1877. non alioram. MonHia Candida Plant, Beitr. z. Syst. Stellnng d. Soorpilzes, 16 pp., 1885, non Bonorden. Dematium albicans Lanrent, Bnll. Soc. Beige Micr. 16: 14, 1890. Monilia albicans Zopf, Die Pilze 478-480, 1890. Endomyces albicans ,Johan-01sen, Centralbl. Bakt. II, 3: 276, 1897, non Vuillemin. Parasaccharomyces albicans Froilano de Mello & Gonzaga Fernandes, Arq. Hig. Pat. Exot. 6: 271-272, 276-277, 1918. Myceloblastanon albicans Ota, Jap. Jour. Derm. Urol. 28: [4], 1928. Candida albicans Basgal, Contr. Estudo Blastomycoses Pulmonares, 49, 1931. Mycotorula albicans Langeron & Talice, Ann. Parasitol. Hum. Comp. 10: 44, 1932. The disease caused by the group of organisms centering about Syringo- spora albicans has been known clinically since the time of Hippocrates as "stomata aphthodea" and Galen "aphtae alba" and ''aphtae infantum." Later it became known as "aphta lactamen" and "aphta lactantium," and EREMASCACEAE IMPERFECTAE 275 in modern literature thrush, muguet, sapinho and Soor. Langenbeck (1839) recognized the presence of the fungus in the disease, although he did not differentiate it from typhus, which it often follows, but Berg (1842) discovered the constant association of the fungus and gave a few morphologic details. Various workers during this decade place the organism in various genera without adding greatly to existing knowledge. Robin (1847) described and figured the organism, placing it in Oidinm without naming it until the revised and greatly enlarged edition of his work in 1853. Haussmann (1870) followed Robin's opinion in his Die Parasiten der weihlichen Geschlechtsorgane. Quin- quaud again studied the organism and placed it in his new genus Syringo- spora, describing the characteristic clusters of blastospores. Many minor works and case histories appeared without affecting nomenclature until Grawitz (1877) called it the same as Mycoderma vim, which opinion gave rise to polemics. While this was unfortunate, Grawitz did call attention to the differences of yeast form and mycelial form, and described chlamydospores. He discusses the action of media on morphologj^ but his observations are to be distrusted on account of the crude state of cultural technic, so that he may have been working with mixed cultures. Reess (1877) showed that the organ- ism was distinct from Mycoderma vini and called it Saccharomyces albicans. This gave rise to further polemics (summarized by Fisclil). Kehrer (1883) studied the physiology of the organism from the standpoint of mode of infec- tion and treatment. Plant (1885) was the first to apply modern cultural technic, and he identified the mycelial form with MoniJia Candida Bonorden on decaying wood. Stumpf (1885) concluded he had two organisms, one filamentous and one yeast, both liquefying gelatin. Baginsky (1885) studied the organism on various media, and Klemperer (1885) produced experimental mycoses from intravenous injections. Grawitz now abandoned his view of relationship to Mycoderma vini. Plant (1887), after a long and detailed studj' with much new data, reaffirmed the identity of his organism with Monilia Candida Bonorden. The first connection between yeast and mycelial forms was proved by Audry (1887), showing that the former Avere common on solid media, the latter in liquids. He described his colonies as follows : Colonies lobulate, pure white, mammillate on gelatin, no liquefaction. On agar and glycerol agar colonies smooth, whitish. On potato, dirty white ; on liquid media, broth turbid, no pellicle, cells elongate in chains. This description is still too gen- eralized to place the organism definitely, but it represents an advance over those of previous workers. In 1890 Linossier & Roux studied the physiology in great detail, giving extensive notes on carbon and nitrogen metabolism without definitely describing biochemical reactions. They describe their or- ganism as producing white, elevated, creamy colonies, with surface slightly furrowed on cooked carrot. At first the yeast cells predominate, then a short period of some filaments and then yeast cells again. On liquid media, the filamentous forms predominate except in malt extract. On most fruits (ex- 276 MEDICAL MYCOLOGY cept melon) and on peptone gelatin, the yeast form is abundant, while on sucrose gelatin, both fonns are found. No ascospores observed, chlamydo- spores not uncommon (Fig. 50). The description of Fischer and Brebeck (1894) is the first which enables us to recognize the organism from its cultural description, consequently it may be taken as a standard for the description of cultural characters and associ- ated with the morphology so well figured by Quinquaud in 1868. This organism or rather the various strains (mostly very imperfectly de- scribed morphologically and culturally) referred here by their authors, seem to be largely facultative parasites, which may invade many tissues when bodily resistance is lowered by disease, senility, etc. The organism is most frequently reported on infants and aged persons, usually in the mouth. This or similar organisms have been reported from the throat in angina, and from the tonsils, the nipples of nursing mothers, occasionally on skin of badly in- fected infants, bronchial tubes (old cases probably to be referred to other species of this group), in the bladder of diabetics, in female genitalia, and in the alimentary canal. It will be noted that many of the cases cited by Fischl (1919) date from the time when it was customary to call any mycelium-pro- ducing yeast from the human cases Monilia albicans or one of its synonyms without comparison with descriptions of previous strains. The only way to prevent further confusion seems to be to take the description of Fischer & Brebeck (1894) as a standard and name strains which are not conspecific with it as something else. Occurring on the mucous membrane of the vagina and in the mouths of infants. Pathogenic to rabbits when inoculated in eye. In young malt extract cultures, cells spherical or ovoid, very variable in size, sprouting. Also occasional elongate cells connected in short chains or longer pseudomycelium. In cultures from room temperatures to 27° C, cells show large vacuoles and one or more metachromatic granules. In malt ex- tract cultures several weeks old, spherical cells 8/a or more in diameter, also short-ovoid, occasional elongate-ellipsoid and filiform cells. Also longer, sep- tate, branched mycelium like that of molds. Similar morphology on old whey cultures with the spherical sprout cells attaining a diameter of 18-20/i,. Each cell contains 1-4 fat droplets 1.5-4/x in diameter (Fig. 51). Giant colonies white to yellow-white, moist, shining, projecting to 2 mm. above the gelatin, guttiform to conic, pulpy in consistency. On gelatin streak, a thick, yellowish white, moist colony which (after liquefaction of the sub- strate) sinks to the bottom in the form of a thick, grayish powdery white or pulpy sediment, leaving the gelatin clear. On stab growth in all directions like a root system. On malt extract between 25° and 27° C, a smooth, dull grayish white pellicle. Glucose, levulose, and maltose fermented. Sucrose only fermented after it has been inverted by the acids formed. Gelatin rapidly liquefied (liquefaction complete in 2 weeks). EREMASCACEAE IMPERFECTAE 277 Syring-ospora Braulti (Pinoy) Dodge, n. comb. Ena7itiothamnus Braulti Pinoy in litt, apucl Brault & Masselot, Ann. Derm. Syphiligr. V, 2: 592-602, 7 figs., 1911; Brault, Bull. Soe. Path. Exot. 7: 90-91, 1914; Bull. Mem. Soc. Chirurg. Paris 37: 405-407, 1911 [original case history with cultural characters but organism not named]. Monilia Braultii Vuillemin, Champ. Parasit. Homme Anim. 87, 1931. Blast odendrion Braulti Langeron & Talice, Ann. Parasitol, Hum. Comp. 10:62,1932. Isolated from slight crateriform tumors on buttock of an Arab in Algeria. Craters opened in a few days, exuding pus, then healed(?). Epidermis not altered. Tissues infiltrated. Subcutaneous injection gave slight lesion in rat, doubtful in guinea pig ; nonpathogenic to rabbit and hen. On carrot broth^ hyphae 2.1yu, in diameter, cells 6.5-10/a long, end cells swollen to 2.8 x 3-7ja. A few branches several cells long. Spores verticillate Fig. 51. — Syringospora chlamydospores. (After Langeron & Talice 1932.) at septum, 2-2.5 x 1-1.5/a. Thallus easily dissociating. Stained by crystal violet, gentian violet, Ziehl, Giemsa, less by thionin, Borrel blue. Loeffler blue, and Unna blue. Cultures groAv at room temperature and 37° C. Colony on Sabouraud agar, colony dull white slightly yellowing, with slight depressions. Margin thicker, finely folded, more brilliant. On glucose agar, after 48 hours, colony thick, center dense, becoming yellowish, margin thinner. Glycerol-glucose agar seems more favorable, center yellow, granular to mammillate, very thick, margin thin, grayish, folded. On potato, colony small, meager. Neither carrot, carrot- glycerol, nor carrot broth very favorable. On ordinary agar, whitish, thick velvet. Velvet on stab at surface with grayish streak in deeper layers. On broth, pellicle forms and medium becomes cloudy. Syringospora interdigitalis (Pollacci & Nannizzi) Dodge, n. comb. Crypt ococcus interdigitalis Pollacci & Nannizzi, I Miceti Patog. 5: No. 44, 1926; Marengo, Arch. Ital. Derm. Sifiligr. Venereol. 1: 464-477, 2 figs., 1926. 278 MEDICAL MYCOLOGY Mycotorula interdigitalis Radaelli apud Flarer, Arch. Ital. Derm. Sifiligr. Venereol. 7: 415-478, 11 figs., 1931. Torulopsis pidcherrima var. variahilis Lodder, Anaskosporogenen Hefen 1: 144-146, 1934. Isolated from interdigital lesions. Cells mostly spherical 3.5-4/a in diameter, thin-walled, uniguttulate. Older cells spherical or ellipsoid 6-8.5 x 5.5-7/a, single or in pairs, thickrwalled. Pro- toplasm at first homogeneous and granular, later vacuolate, blastospores some- times solitary, sometimes in verticils about cylindric septate hyphae, 2-3.5/a in diameter, sparingly branched. On Pollacci agar, colonies punctiform, moist, creamy, yellowish. Flarer reports colony dense, creamy, whitish, shining; margin smooth, regular, and confluent; slightly grayish when old. On malt agar, colony yellowish, shining, slightly radially striate, margin smooth. On malt gelatin, colonies irregularly reddish, flat, smooth, radially striate, margin sinuous. On malt extract, a thin ring and thin, slimy pellicle which soon sinks, slight odor of esters. On broth, an incomplete ring with a cottony, floccose sediment. On alcohol, a thin pellicle. Glucose, fructose, and mannose fermented. Gelatin liquefied in 75 days. Syringospora cutanea Dodge, n. sp. Mycelohlastanon cutanenm var. Takahashi, Jap. Jour. Derm. Urol. 29: 224- 251, 318-331 [26-31], 1929. Isolated from mycosis of interdigital spaces as well as of gluteal, perianal, inguinocrural, and axillary folds. In one case, isolated from neck of infant. Yeast cells ovoid or ellipsoid, 2-4 x 3-5/*, granular at first, becoming vacuo- late and thick-walled. Mycelium 2-5/* up to 50/*, septate. Mycelium forma- tion is stimulated by the following factors : lowered temperature, high oxygen tension, slight acidity of substrate, depletion of nutrients, addition of carbo- hydrates, especially glucose, addition of "koji, " growth on carrot, on man, or other mammals. Blastospores appear at the septa, often in dense clusters suggestive of a mulberry. No asci formed. Colony on malt agar or Sabouraud glucose is brownish white, smooth, shining and creamy, with the margin showing dendroid projections of my- celium into the medium. On broth, producing a slight turbidity, ring, sedi- ment, but no pellicle. Growth better on glucose broth. Glucose, maltose, fructose, mannose fermented, but not sucrose, galactose, raffinose, dextrin, lac- tose, or mannite. Gelatin not liquefied, though dendroid mycelium appears along the stab. Malt gelatin slowly liquefied. Milk acidified and coagulated, the optimum temperature for this being 37° C. Syring"Ospora Tonge (Mattlet) Dodge, n. comb. Monilia Tonge Mattlet, Ann. Soc. Beige Med. Trop. 6: 22, 23, 1926. Isolated from cases showing sjonptoms of true dysentery and of enteritis. No amoebae or cysts found. Patients natives of Belgian Congo. Ameliora- tion of symptoms and even cure on treatment with light purgatives in con- junction with emetine injections. EREMASCACEAE IMPERFECTAE 279 After 3 days at 37° C. in potato decoction, blastospores, spherical up to 7/i in diameter, each containing a big vacuole, sprouting at several points at once with these daughter cells sprouting in turn without separating from the mother cell. Hyphae formed of cells that lengthen with age, beginning about 20 X 2.5/A and ultimately attaining 30 x 1.5 fx. The protoplasm of the cells is reduced to a thin layer along the wall. Verticils of spherical cells at the septa; lateral branches rare. After 30 days intercalary and terminal chlamy- dospores, which have a double membrane and a large vacuole, separate and float free. Optimum temperature for growth 37° C. On Sabouraud agar at 37° C, colony round, dull white, with a smooth margin at first, developing outgrowths composed of large, contorted hyphae, which also grow down into the medium. On gelatin stab, culture white, with denticulate margin, horizontal hyphae growing in depth of culture give a growth of inverted cone shape. On potato decoction, white flaky sediment. Ferments glucose and fructose, not maltose, galactose, sucrose, lactose, man- nite, dextrin, or inulin. Neither acid formation nor clotting in milk. No liquefaction of gelatin. Syring-ospora psilosis (Ashford) Dodge, n. comb. Monilia sp. Ashford, Jour. Amer. Med. Assn. 64: 810, 811, 1915; Amer. Jour. Med. Sci. 150: 680-692, 1915. Monilia psilosis Ashford, Amer. Jour. Med. Sci. 154: 157-176, 1917 [August] . Parasaccharomyces Ashfordi Anderson, Jour. Infect. Dis. 21: 341-387, Pis. 3-8, 1917 [October]. MyceloMastcmon psilose Ota, Jap. Jour. Derm. Urol. 28: [4], 1928. Mycotorula psilosis Langeron & Talice, Ann. Parasitol. Hum. Comp. 10: 47, 1932. Candida psilosis Almeida, Amiaes Fac. Med. Sao Paulo 9: 11, 1933. Isolated from a patient suffering from sprue in Puerto Rico. Pathogenic to guinea pigs and rabbits. In young cultures cells are spherical or slightly ovoid ; in old cultures cells more variable, ovoid, elongate, ellipsoid, spherical, or irregular. Giant cells common, spherical, 3.5-5/a in diameter, Avitli refractile oil droplet. Septate hyphae appear in hanging drop of gelatin and in old cultures. Hyphal cells 2-4/x in diameter, branching not uncommon but not t^'pical. Sprouting may occur anywhere in young cultures but near ends of cells in old cultures. It is the normal method of multiplication. On glucose agar streak, colony filiform, elevated, glistening, chalk-white, and smooth. Later center may become rugose or pitted, margin even or fila- mentous. On gelatin stab, growth at first filiform, later developing scattered bushy clusters of filaments. In liquid sugar media and malt extract there is a ring, but no pellicle. Liquid sugar media become at first acid, then more alkaline. Glucose, maltose, and fructose fermented. "Occasionally sucrose 280 MEDICAL MYCOLOGY and galactose" — Anderson [probably due to sugar impurities or perhaps strains of Castellania faecalis]. Litmus milk alkaline but not clotted in 2 weeks. "Gelatin rarely liquefied" — Anderson. Syringospora Cavarae (Pollacci & Turconi) Dodge, n. comb. Cryptococcus Cavarae Pollacci & Turconi in Bencini & Federici, Atti R. Accad. Fisiocrit. Siena X, 3: 746-766, 8 figs., 1928. Isolated from primary corneal lesions in Siena, Italy. Lesions reproduced in the eye of a rabbit. Cells spherical, 4-7/*, or ovoid, 6-9 x 4-6/i, sprouting, wall thick, homo- geneous or uniguttulate, rarely biguttulate, hyaline or yellowish ; later the cells elongate to subcylindric 14-24 x 3-5/*, sprouting only from the apex, forming a mycelium, septate, simple, or slightly branched. Blastospores in small verticils of only a few cells each. On Pollacci agar, colony disciform, creamy, moist, milky white, then slightly yellowish, slightly elevated in the center, plane, or more or less rugose. Gelatin rapidly liquefied, milk not coagulated, no fermentation of sugars, slight assimilation of glucose, fructose, maltose, and galactose. Syring-ospora Issavi (Mattlet) Dodge, n. comb. Monilia Issavi Mattlet, Ann. Soc. Beige Med. Trop. 6: 21-22, 1926. Case history not given. After 3 days at 37° C. in potato decoction, three types of growth: (1) Blastospores, spherical, 5-6 fi, or ovoid, 7 x 4/*, containing a vacuole. (2) Groups, sometimes in chains, of several elongate cells (6-8 x 2.5-3.5/*), bearing at ends and at septa, spherical or ovoid blastospores. (3) Hyphae long, made up of elongate and narrow cells (3.0 x 26.5/t), not much narrowed at the septa, free ends rounded, with either spherical blastospores or other hyphae at the septa. After 10 days the hyphae form a felt in whose interstices appear free cells, some of which enlarge considerably, becoming, when ovoid, 8 x 4/*, and, when elongate, 15 x 2.5/*. In some young hyphae the end of a cell swells. After 30 days there is no further change. On Sabouraud agar at 37° C, colony white, somewhat shining, yellowing, and thickening with age, becoming irregular with radiating striations at mar- gin, giving a fringed appearance. Abundant growth of hyphae into medium, giving it a granular appearance under a magnifying glass. On gelatin stab, same appearance at surface, medium becomes cloudy beneath, with gas bubbles. In potato decoction, flaky white sediment. Optimum temperature 37° C. Litmus milk at first acid, then decolorized. Fermentation of glucose, fruc- tose, maltose, galactose, sucrose, dextrin; none of lactose, mannite, inulin. Milk coagulated. Slight evolution of gas with gelatin. Syringospora Negroni Dodge, n. sp. Cryptococcus sp. Negroni, Rev. Soc. Argentina Biol. 6: 648-652, 1930; Rev. Univ. Buenos Aires II, 29: 360-364, Figs. 60-62, 1931. EREMASCACEAE IMPERFECTAE 281 Isolated from moist intertriginous lesions, once from interdigital mycosis of the toes, once from vulvitis, and once from the epidermis of the male geni- talia. [Cases very briefly described and not figured.] Yeast cells isolated, spherical or ovoid, 4-5ju, in diameter, rarely 3-10/i. Rudimentary mycelium present, hyphae short, septate moniliform, blasto- spores borne in verticils at the nodes and terminally. Yeast cells with a large vacuole and a polar oil globule. Chlamydospores present. No ascospores on Gorodkova agar or on gypsum block. Growth good on Sabouraud agar and malt agar, colonies confluent, form- ing a thick, white, moist, creamy layer. Giant colony on malt, 4 cm. in 25-30 days, circular, margins festooned, center slightly elevated, occasionally conic and granular, shining, humid, grayish white, with 4-5 radial furrows. On carrot, growth good, creamy, white, no spores. Malt extract fermented, with a ring and sediment. Litmus milk coagulated and acidified. Glucose, fruc- tose, and maltose fermented; not lactose, sucrose, or galactose. Malt gelatin fermented and liquefied. Optimum temperature 30° C, maximum about 45°, and minimum 12°-15° C. The following species are placed here from their morphology, but their descriptions are too incomplete to refer here with certainty. It is possible that some of them may belong in synonymy with better described species. The organisms of Hasegawa (p. 282) and of Braafladt (p. 282) may very pos- sibly belong elsewhere. Syringospora uvae (Pollacci & Nannizzi) Dodge, n. comb. Cryptococcus uvae Pollacci & Nannizzi in Motta, Atti E,. Accad. Fisiocrit. Siena IX, 17: 636, 1 pi., 1925. Monilia uvae Vuillemin, Champ. Parasit. Homme Anim. 48, 1931. Isolated from lesions on uvula, sometimes on pillars of fauces. Cells spherical, 4-8/a in diameter or ovoid, 7.5-9 x 9-10/*, some as high as 15/1. Sprouting polar. Hyphae septate, 2-2.5/* in diameter, alternately branched with glomerules of blastospores at tips. On Pollacci agar, colonies round, 1-2 mm. in diameter, moist, projecting 0.3-0.4 mm. above substrate, and 0.5-1.0 mm. into it. Radiating hyphae in the upper part of the agar extend 2-4 mm. On malt extract agar, colony gray, dull, smooth, margin smooth. On malt gelatin, colony gray, shining, flat, center slightly elevated, margin somewhat sinuous. In liquid media, floccose colonies form on the surface and settle to the bottom, the liquid remaining clear. No growth on alcohol. On malt extract, a few islets, a thin ring, and sediments. No fermentation; gelatin not liquefied in 57 days at 15° C. Syringospora Otae (Nannizzi) Dodge, n. comb. Cryptocococcus Otae Nannizzi, Tratt. Micopat. Umana [Pallacci] 4: 329, 330, 1934. Cryptococcus de Burnier (cas C) Ota, Ann. Parasitol. Hum. Comp. 2: 49-51, Fig. 8, 1924. Isolated from a case of epidermomycosis. 282 MEDICAL MYCOLOGY On malt agar at 25° C. after one day, cells ovoid or slightly elongate, but often spherical 8 x 5/t, thin-walled, solitary or in pairs, sometimes in short chains or small groups. Cells mostly solitary after a week, walls thicker. On carrot, elongate cells also occur, 15 x 4/x, ends rounded, forming longer chains. Blastospores in simple verticils at the septa, often separating soon leaving one or two which elongate to form branches. On malt agar, colonies white or slightly .yellowish, smooth, margins defi- nite. On malt extract, no ring or pellicle, but sediment formed. Syringospora Hasegawae, Dodge, n. sp. Oidium alhicans var. Hasegawa. Jap. Jour. Derm. Urol. 28: 1104-1124, 7 figs. [86-87], 1928. Isolated from three ulcers size of hen's egg on lower leg; clinically re- sembling Zymonema dermatitidis. Pathogenic to mice and guinea pig. Cells S-6fi in diameter, thick-walled, spherical, some also ovoid or elongate. Chlamydospores 7.5^. No asci. Conidia at the septa. Colony, on malt agar, shows characters of a wild bottom yeast. On sugar agar, produces milky white disc with brownish tone. Fermentation negative. Gelatin liquefied. Milk coagulated, with production of acid. Doubtful Position Cryptococcus sp. Braafladt, China Med. Jour. 35: 30-35, 1921. Isolated from chronic sinuses on breasts of Chinese woman. Breasts removed surgically. In a second case, that of a foreign woman, in which the organism seemed to be the same, there was slight pain in the chest, cough with bloody sputum, dyspnea, and weakness. Cough developed in Chicago before going to the Orient. There is nothing in the article to show which case belongs to the following organism or whether characters of the two strains may not have been confused. The early colonies form yeast cells only; later clusters of spherical, gram- positive spores attached to long, slender, gram-negative hypliae containing some gram-positive granules. Still later thicker gram-negative hyphae con- taining spherical or ovoid spores. The transition from yeast cells to hyphae occurs only on solid media and apparently is not reversible. Colonies on agar, circular, flat, opalescent, 2-3 mm., after 3 weeks begin- ning to grow aerial hyphae. On nutrient broth at 37° C, grayish pellicle, heavy sediment at the bot- tom of the test tube. No pellicle from subcultures. The first case suggests Zymonema dermatitidis or AtelosaccJiaromyees hom,inis. BLASTODENDRION Blasiodendrion Ota, Derm. Woch. 78: 216-264, 1924; Ciferri & Redaelli, Atti 1st. Bot. R. Univ. Pavia III, 2: 129-146, 1925. fOidiodendron Robak, Nyt. Mag. Vidensk. 71: 243-255, 1932. EREMASCACEAE IMPERFECTAE 283 The type species is Blastodendrion Krausi Ota. Colony creamy, thin, beginning from the germination of thick-walled blastospore, forming a dendroid mass by sprouting, either bipolar or multipolar, rarely cruciate; blastospores polymorphous, the lacrimiform or pyriform type predominating; pseudomycelium more or less developed, little branched, less easily dissociable than in related genera, forming dendroid masses with ascend- ing branches parallel or suggesting the branching of sporophores in Penicillium, cells mostly elongate, pyriform, hyhpae terminated by a chain of lacrimiform blastospores or by a long slender filament; verticils occasionally present, formed of lacrimiform blastospores. Found in a variety of conditions, the most conspicuous being a group on moist skin and about nails. This whole group needs further study to differen- tiate it both morphologically and culturally from related groups. Langeron & Talice made a good beginning but much more needs to be done. Key to Species Pellicle produced on liquid media. Cells 5 X 10m. ^- Krausi. Cells 2-4^1. B. intermedium. PTo pellicle on liquid media. Glucose, fructose, and maltose fermented. On mucous membranes. B. Finoyi. On moist folds of skin. B. Favrei. Only glucose fermented. B. elongatum. Sucrose fermented. B. Kayongosi. Galactose fermented. Cells 50-60/x long. B. cutaneum. Cells up to lOfi long. B. gifuense. Blastodendrion Krausi Ota, Derm. Woch. 78: 229-230, 1924. "Pferdehefe II." In 24-hour-old cultures may be found rudimentary branched mycelium of round or long, ovoid cells not easily breaking apart, cells 5 x lO/x, many small fat granules. In 3-day cultures, cells 15/x long. Mycelial habit retained up to a month, fat drops peripheral to a central vacuole. On carrots, morphology is similar, but with much longer cells. On malt extract, a pellicle appears and on inner wall of tube, cells 10-20 X 3-4/x, with abundant branching. Also branching forms found in sediment. Blastodendrion intermedium Ciferri & Ashford, Porto Rico Jour. Publ. Health & Trop. Med. 5: 91-105, 4 pis., 1929. Isolated from the human intestine in a case of sprue, but not pathogenic for rabbits, and since Syringospora psUosis (Monilia psilosis) was also present in abundance, probably not connected with sprue. ■ 284 MEDICAL MYCOLOGY Cells spherical to ellipsoid or ovoid, rarely of other shapes, normally 2-4/4, with small cells 1-1.5/x and giant cells up to 10/x (Fig. 52). Giant colonies white, with one or more central craters, margin lobular, and denticulate (type II of Will). On liquid media, pellicle, ring, and sedi- ment present. Ferments glucose, fructose, sucrose, maltose, trehalose, meli- tose and galactose; fails to assimilate lactose and all alcohols except glycerol ; assimilates peptone and asparagin, ammonium sulphate slightly, but not nitrites or nitrates. Gelatin not liquefied. 4 ^ Fig. 52. — Blastodendrion intermedium. A, mycelium and blastospores ; B, actively sprouting blastospores ; C, germination o° blastospores. Blastodendrion elongatum Mattlet, Ann. Soc. Beige Med. Trop. 6: 17-18, 1926. One of the organisms isolated from stools of patients suffering from true dysentery in Belgian Congo. Also isolated in one case of European resident suffering from enteritis. After 3 days in potato decoction, spherical cells, about 6/a in diameter, or ovoid cells, 8 x 4/a, containing large vacuoles, also long narrow cells, 3 x 12-23/*. Cell groupings do not give rise to simple filaments. Blastospores spherical, ovoid, or elongate at the septa. EREMASCACEAE IMPERFECTAE 285 On Sabouraud agar at 37°, colony dull white, finally yellowing at the center, with radial i'olds; margin lobed, and numerous verruciform projec- tions into medium. On gelatin stab, appearance at surface the same, but in depth, numerous small granules. In potato decoction, white lumpy sediment which breaks up somewhat on shaking. Optimum temperature for growth 37° C. Litmus milk at first acid, then alkaline. Glucose fermented; fructose, sucrose, and dextrin slightly fermented ; no fermentation of maltose, galactose, lactose, mannite, or inulin. Gas evolution with gelatin. Blastodendrion Kayongosi Mattlet, Ann. Soc. Beige Med. Trop. 6: 20-21, 1926. Isolated from a grayish white covering on tongue and tonsils of five-year- old boy in Belgian Congo, which prevented him from eating. Elevation of body temperature also observed. Scraping and application of tincture of iodine caused amelioration of symptoms. Patient not seen again. In scrapings, blastospores spherical, 2-3/^ in diameter, others ovoid or elongate of analogous dimensions. After 3 days' growth in potato decoction at 37°, cells mostly rounded with an oil droplet, maximum diameter 6/*. After 30 days, hyphal forms appear, much branched, elongate cells averaging 7 x 2.5/*. Branches sometimes reduced to a single spherical or elongate cell. Optimum temperature for growth 37° C. On Sabouraud agar, colony round, dull, white with center later yellowing and wrinkling margin with radial striations which cause fine indentations. On gelatin stab, surface of same aspect as on Sabouraud agar. Fine points of growth along the stab, with displacement of medium. In potato decoction, flaky sediment which separates on shaking, no pellicle. Glucose, fructose, maltose, galactose, and sucrose fermented, not lactose, mannite, dextrin, or inulin. Slight acid production in milk, no clotting. Gelatin not liquefied, but gas produced. Blastodendrion Pinoyl (Castellani) Langeron & Talice, Ann. Parasitol. Hum. Comp. 62, 1932. Endomyces Pinoyl Castellani, Lancet 1: 15, 1912; Brit. Med. Jour. 2: 1208-1212, 1912. Monilia Pinoyi Castellani & Chalmers, Man. Trop. Med. ed. 2, 826-827, 1913. Mycelohlastanon Pinoyi Ota, Jap. Jour. Derm. Urol. 28: [4], 1928. Odlund & Hoffstadt, Arch. Derm. Syphilol. 20: 335-338, 1929, give good description and drawings of their ease of Monilia Pinoyi infection of the penis and vagina (Fig. 53). Originally isolated from sputum in case of bronchomycosis. Acidification and fermentation with glucose, fructose, and maltose. Milk not clotted, coagulated serum, and gelatin not liquefied. Permanand (1922) gives the following cultural characters. Colonies dull white, smooth with an ivory-like surface, sticky, dense, not adherent to the medium. 286 MEDICAL MYCOLX)GY Lasseur & Servet (1922) describe cultures as follows. Opaque, white ele- vated and round on Sabouraud maltose. On gelatin, velvety, thick, elevated, pure white. Growth poor on potato and turnip. Acidification and fermenta- tion with glucose, fructose, and maltose, not with other sugars. Growth on coagulated serum poor, no liquefaction. Gelatin not liquefied. Organism gram-positive. Blastodendrion Favrei (Ota) Dodge, n. comb. Levure de Favre Ota, Ann. Parasitol. Hum. Comp. 2: 51, 1924. Fig. 53. — Blastodendrion Pinoyi. (After Odlund & HofCstadt 1929.) Mycelohlastanon Favrei Ota, Ann. Parasitol. Hum. Comp. 3: 181-184, 1 fig., 1925. Cryptococcus (Mycocryptococcus) Favrei Pollacci & Nannizzi, I Miceti Patog. 9: 83, 1929. Candida Favrei Almeida, Annaes Fac. Med. Sao Paulo 9: 11, 1933. First case cited, Favre & Ota, C. R. Soc. Biol. 88: 222-224, 1924. Isolated from pruriginous dermatitis of inguinal and axillary folds. Sub- cutaneous or intraperitoneal injections pathogenic to rabbit and guinea pig. Morphology on carrot : cells spherical, ovoid, ellipsoid, often elongate, isolated or in twos, occasionally agglomerated in larger number. In deeper EREMASCACEAE IMPERFECTAE 287 layers cells are in chains, 2.5-3.5 x 10-20//., developing by budding, chains branched, cells larger at one end. Same, but less pronounced, on glucose agar, malt agar, or bouillon. True mycelium in 15 days on carrot or in potato decoction. Hyphae slender, 1-2/x, septate as in Mycoderma lactis. Maximum temperature for growth 50°-52° C, optimum temperature 35°- 40° C, no growth at 12° C. In 12 days at 25° C, giant colony attains size of one franc piece. On Pollacci agar, colony irregular, white to yellowish, smooth, moist. Colony on malt agar at 25° C, for 24 hours, round, sometimes elliptic or oval, 3-7 cm. Refractive granules fuse on fifth day. Giant cells 10/a with numerous fatty granules. Growth on carrot similar. In malt extract, a grayish white sediment appears in 24 hours, but no pellicle or ring in 25 days. After a month, some cells tend to creep up the sides of the tube. No spores formed on Gorodkova agar, carrot, or plaster block. The giant colony appears gray- ish white, humid, brilliant, margin rounded and distinct, rays from center. Organism does not invert sucrose; moderate fermentation of glucose, fructose, and maltose, none of other sugars. Gelatin not liquefied. Blastodendrion cutaneum (Ota) Dodge, n. comb. Mycelohlastanon (Mycelorrhizoides) cutaneum Ota, Derm. "Woch. 78: 232- 234, 1924. [Oidiomyces unguium] Frei, Arch. Derm. Syphilis 129: 404-433, 1921, or- ganisms of Fabry, Miinchen. Med. Woch. 65: 1557-1558, 1917; Ibid. 66: 321, 1918; Berendsen, Arch. Derm. Syphilis 126: 751-763, 1919; Samson, Derm. Woch. 76: 473-481, 1922. In 24-hour-old cultures on malt agar at 25° C., cells are mostly ovoid, thin- walled with large vacuole and very small oil globule. Cells, sometimes in chains, usually 2-5/1, (sometimes 9/i) in diameter, 50-60/1 long, Ifi in diameter at rounded end and 3/^ at point. In 10 days, cell chains have broken up, cell walls are thickened, and large fat granules appear. In 30 days, they have broken down into ovoid forms, interior with large vacuole and fat drops. On carrots, mycelium penetrates substrate as slender cells, 2-3/t in diameter; hyphae dichotomous. No pellicle on malt extract. Fermentation with glucose, fructose, mannose, galactose and maltose only. Var. Fujii Dodge, n. var. Mycelohlastanon cutaneum var. 7. Fujii, Jap. Jour. Derm. Urol. 31: 959- 983, J5/i^5. [71-72], 1931. Organism isolated from an infection on the hand of a Japanese farmer. Three years before the middle finger became grayish in color and translucent, slightly swollen. Surface dull, edematous, erj^thematous, then cracked and became vesiculose with little pruritus. Organism injected intraperitoneally into mice, was fatal to most within 72 hours. Cells 15-13-10 X 7.5-5.8/1 in diameter, spherical, ovoid, or polyhedral. Pyri- form cells show diameter of 3/i, at smaller end. Cell wall thin. On Gorodkova 288 MEDICAL MYCOLOGY agar, hypliae 3-4.5/* in diameter, transparent, flexuous, up to 45/a long. Blasto- spores showing nucleus and vacuoles 4/i in diameter. On Sabouraud glucose agar after 10 days, colony circular, 1.5 cm. in di- ameter, surface flat, margin thinning, center slightly elevated, moist, grayish yellow. On malt agar, in 10 days colony circular, 1.5 cm. in. diameter, yellow- ish white, umbilicate, shining. By the twentieth day 2 cm. in diameter with radiating rays which become closer together with age, surface moist, center gradually depressed, medium not discolored. On glucose gelatin, at tenth day colony round, 1 cm. in diameter, smooth, moist, sticky, grayish white. On twentieth day colony same size with margins denticulate. On malt extract in 10 days, liquid cloudy and sediment present ; no pellicle or ring mentioned. Acid production with fructose and mannose. Fermentation of galactose, arabinose, xylose, rhamnose, and maltose. None with glucose (!), lactose, raffinose, inulin, mannite, peptone, or starch. Gelatin not liquefied. Blastodendrion gifuense (Taniguchi) Dodge, n. comb. Mycelohlastanon gifuense Taniguchi, Jap. Jour. Med. Sci. xiii, Derm. Urol. 1: 74-94, Pis., 1-2, 1926. Producing interdigital blastomycosis among paper workers, locally known as sadare, in the province of Gifu. In the manufacture of mino paper, the workers keep their hands in water practically the whole day. Paper made from bark of Broussonetia papyiHfera Vent, or Quercus glandulifera Blume, bark bleached with CaOCl, washed thoroughly with water, then a maceration of the root of Hibiscus japonicus Miq. or H. Manihot L. is added for the slime which holds the paper fibers together and makes it suitable for umbrellas. In a village of 50 workers examined, of 30 infected, most had used local remedies, juices of plants. Also some attempt had been made at the factory by dipping the hands in CUSO4 solution or HgClj, the former ineffective, the latter effective in 1 :10,000 dilu- tion. Organism was isolated from raw root of Hibiscus, from the preparation used in paper making, as well as from the paper makers. Pathogenic for mice. Cells ovoid, thick-walled, 2 x 4-3 x 5/a, up to lO/i long. Optimum tempera- ture 25°, growth very slow at 37° C. No asci seen. On glucose and malt agar, growth good, colonies round, yellowish white, smooth, margins sharp, moist, shining, center slightly elevated, deeper col- ored, whole colony browns in drying out. On carrot, growth good, yellowish or clay color. On malt extract, turbid in 2-3 days, with sediment, but no pel- licle or ring. Ferments maltose, mannose, fructose, glucose (one strain fer- mented galactose and one strain raffinose) as does Mycelohlastanon cutaneum (Fabry, etc.). Blastodendrion Arzti Ota, Derm. Woch. 78: 232-234, 1924. On malt agar, cells ovoid or long ellipsoid, 2-6 x 3-10/i,, others 2-3 x 30-60/1 with rounded ends, one end usually rather thicker than the other, arranged in loose chains ; other cells 10-20 x 3-5/i with pointed ends ; round cells have one or two fat drops, long cells, many small ones. In hanging drop, branch- ing chains are formed which cling together for several days. Gas with levulose, sucrose, and raffinose only. EREMASCACEAE IMPERFECTAE 289 Myceloblastanon (Mycelorrhizoides) Gruetzii Ota, Derm. Woch. 78: 226, 264, 1924. This is proposed as a nomen nudum. Blastodendrion erectum Camargo, Agentes Etiologicos do Sapinho 60, 1934 (nom. nud.). REDAELLIA Redaellia Ciferri, Arch. Protistenk. 71: 424-428, Fig. 3, 1930. The type species is Redaellia elegans Ciferri. Fig-. 54. — Redaellia elegans Ciferri. (After Ciferri 1930.) Colony growth slow, elevated, cerebriform, irregularly convoluted, hyphae hyaline, septate, branched with many fusiform blastospores at the tips ; blasto- spores germinating either by sprouting or by hyphae. Only a single species of doubtful pathogenicity has been referred here. Redaellia elegans Ciferri, Arch. Protistenk. 71: 424-428, Fig. 3, 1930. Isolated along with Epidermophyton ruhrum from a case of tinea axillaris in Santo Domingo. Blastospores apical, 1-20/x, usually S-lOfx, on the top of a slightly inflated cell, more or less fusiform. Ciferri states pyriform, but his drawings show uniformly fusiform blastospores, 6-8 x 2.5-3^, which he was unable to germi- nate in hanging drop; gelatin not liquefied, sugars not fermented (Fig. 54). 290 MEDICAL MYCOLOGY I am doubtful whether this genus really belongs to the Eremascaceae. Evidently a contamination, suggesting a primitive Basidiomycete or perhaps an unreported member of the Tulasnellaceae which grew in the cultures. The drawings do not suggest any other filamentous yeast. The mycelium is largely of the raquet type, if the drawings are correct. MYCOTORULOIDES Mycotoruloides Langeron & Talice, Ann. Parasitol. Hum. Comp. 10: 48-54, 1932. The type species is Mycotoruloides triadis Langeron & Talice. Colonies creamy, thick, convex, beginning by bipolar sprouting of a blas- tospore followed by progressive branching of the pseudomycelium. Blasto- spores spherical or ovoid, arranged in verticils, arising at the apical portion of I I Fig. 55. — Mycotoruloides. 1, young mycelia ; 2, mature mycelia. 1932.) (After Langeron & Talice the pseudomycelial cells, no terminal chains of cells. Pseudomycelium formed of short cells, each cell producing a verticil of blastospores at its apex. The verticils are less regularly spaced than in Syringospora and are usually com- pound, producing an ovoid mass of blastospores whose long axis is more or less perpendicular to the main axis of the pseudomycelium (Fig. 55). Some branches develop more than others, giving an irregular appearance. Occasion- ally a branch grows out and is terminated by short chains from its terminal verticil. Gelatin not liquefied. I Key to Species M. unguis. No pellicle on liquid media, from nails. Pellicle or ring on liquid media. Glucose, maltose, and sucrose fermented, galactose acidified, gelatin not liquefied, from chronic ulcers of tongue. M. argentina. EREMASCACEAE IiMPERFECTAE 291 Glucose and maltose fermented. Gelatin liquefied, no action on gitla(;tot:o, i'loni iungs. ,1/. triadi.i. Gelatin not liquefied, acid produced on galactose, from tongue. M. aldoi. Mycotoruloides arg-entina (Vivoli, Avellaneda & Bardessi) Dodge, n. comb. Monilia argentiiia Vivoli, Avellaneda & Bardessi, Reunion Soc. Argentina Patol. Keg. del Norte en Tucuman 7: 239-277, 37 figs., 1932. Geotrichoides tumefaciens Talice & Mackinnon p.p., Reunion Soc. Argentina Patol. Reg. del Norte en Santiago del Estero 8: 161, 162, 1933. Ulcerous lesions on tongue, complicated by syphilis. Biopsy showed fila- ments 20-60 X 2/A, unbranched, septate. Autopsy showed nodular lesions in lungs. Pathogenic for rats, rabbits, and guinea pigs. On Sabouraud agar, yeast cells spherical or ovoid, 4-7yu, in diameter and liyphae unbranched, terminal cell somewhat clavate, septate, up to 60/* blasto- spores polymorphic ; coremia frequent. Colonies shining, pasty, not adherent, confluent, margins irregular with fine radiating hyphae. On malt agar, colony slightly granular, yellow. On Pollaeci agar, colony spherical, at first yellowish white, becoming thick, rugose, dry. On potato glycerol, colony thick, grayish white ; on carrot glycerol, colony about 2 mm. thick, white rugose, dull, partly covered by a fine dark brown layer; on beets, white pasty colony which assumes a rose tint. On gelatin, growth good ; on coagulated serum, growth poor ; on both, colony white with some hyphae. In malt extract, or Raulin's liquid, white powdery sediment, no ring or pellicle. On sugar broths, thin white pellicle ; glucose, maltose, and sucrose fermented, slight acidity in fructose, galactose, and lac- tose but at the end of a month all sugar broths have become alkaline. Milk coagulated, gelatin and serum not liquefied. Mycotoruloides unguis (Weill & Gaudin) Langeron & Talice, Ann. Para- sitol. Hum. Comp. 10 : 50, 1932. Spicaria unguis Weill & Gaudin, Arch. Med. Exp. 28: 461-463, PI. 12, 1919. Monilia unguis Vuillemin, Champ. Paras. Ilomme Anim. 85, 1931. Found on nails in two cases of onychogi-yposes with splitting of nail into white and yellow layers. Mycelium slender, 1.5fi in diameter, septate, sometimes closely entangled and bearing spherical or ovoid blastospores. Mycelium branched, septate with septa 3-6/* apart. Verticillate pyriform buds at level of the septa give rise to verticillate chains of blastospores which are hyaline, rounded, or pyriform, 3 X 4-5/x. Organism grows on all the usual substrates, best at 37° C. On Sabouraud maltose agar, growth fleshy, white [or yellowish if the maltose is colored], convoluted. On potato, colony humid, grayish, 2-3 mm. thick, chalky on desic- cation. Growth vigorous on carrot. In giant colonies, growth is 10-12 mm. in 10 days, dirty white, brilliant witli densely twisted masses of hyphae about 1.5 cm. thick. In Raulin's liquid a scanty white granular sediment appears at the bottom of the tube. 292 MEDICAL MYCOLOGY Mycotoruloides triadis Langeron & Talice, Ann. Parasitol. Hum. Comp. 10: 48,1932. Monilia sp. Brocq-Rousseii, Guillierniond & Cilleuls, Ann. Parasitol. Hum. Comp. 5: 48-62, 1927. Isolated from an infection of the lung, clinically suggestive of tuberculo- sis. After the diagnosis was established, patient cured in a short time by potassium iodide. Pathogenic to guinea pig and rabbit. On malt extract developing as a yeast, cells spherical 2.8-8^i in diameter, or ovoid, 4-8 x 6-10/*, united in small groups, then developing a septate, branched mycelium of normal slender cells and swollen cells. Giant cells 10.5-18/1 sometimes up to 40/i in diameter. Mycelium early appearing on malt extract agar, yeast cells predominant in the pellicle. On agar, colonies white, almost transparent, thick. On Sabouraud agar similar, but colony whiter and more abundant, confluent. On malt agar, yeast decoction agar, and Gorodkova colony humid and white. On malt agar after 15 days, colony circular, yellowish white, moist, center granular reticulate, margin smooth with a few radial fun-ows, composed of large lobes. After 2 months, center is thicker, mesenteroid in appearance, surrounded by a broad smooth zone with radial furrows, margin thinner and finely festooned. No change of color in neutral red agar, no blackening in lead subacetate agar; on litmus sucrose agar, slight reddening of the medium, soon turning back to blue. On litmus lactose agar, slight growth, no change of color. On carrot and potato, colonies white and confluent. On gelatin, colony creamy white, growth slow. In gelatin stab, growth conic above, suggesting a pipe cleaner. On coagulated serum, growth slight. On malt extract, whitish sediment with fermentation, then a ring forms and about the fifteenth day spreads to cover the surface, brownish yellow but no true pellicle. In yeast decoction, grayish islets which coalesce to form a complete delicate pellicle. No indol on pep- tone. Milk coagulated in 12-15 days, curd digested, no acidity developed. Albumin digested. Starch not hydrolyzed. Sucrose inverted; glucose, fruc- tose, maltose, and dextrin fermented. No action on galactose, lactose, and raffinose. Liquefaction starts on the tenth day and is completed in 2 months. Mycotoruloides aldoi (Pereira Filho) Langeron & Talice, Ann. Parasitol. Hum. Comp. 10: 49, 1932. Monilia aldoi Pereira Filho, Jour. Trop. Med. Hyg. 27: 8-10, 1927. Candida aldoi Castellani & Jacono, Jonr. Trop. Med. Hyg. 36: 317, Fig. 46, 1933. Isolated from a case of hypertrophy of the tongue with white spots ap- pearing. Intraperitoneal injection caused death of rabbit in 3 days, intra- venous injection caused death of guinea pig in 2 days. Organism gram-positive. Mycelium reported but no ascospores. In culture gave small, smooth white colonies composed mostly of yeast cells with some mycelium. On "yeast medium" (equal quantities of yeast and 3% glucose; solution autoclaved at 115° C. for 15 minutes) gave rough EREMASCACEAE IMPERFECTAE 293 colonies and niycelinm. In broth, a slight turbidity appears with a white powdery deposit. In peptone water, no turbidity, but white flakes on walls and at bottom. No acid formation here or in milk. In yeast extract, turbidity and pellicle at sides of tube. Glucose, fructose, and maltose fermented. Acid for- mation with sucrose and galactose. No action with lactose, mannite, dextrin, spores (Fig. 56). Gelatin not liquefied. MYCOCANDIDA Mycocandida Langeron & Talice, Ann. Parasitol. Hum. Comp. 10: 56-58, 1932. The type species is Candida mortifera Redaelli. Colonies creamj^, sometimes thick, more often thin, flat, iridescent, trans- parent at first, gently sloping from the center, sometimes coremia pres- ent; colonies beginning by bipolar sprouting; blastospores appearing late, dimorphous, ovoid or elongate, the latter predominant, much less numerous than in the preceding genera. Pseudomycelium well developed, highly branched (suggesting a fir tree), hyphae ending in a group of blastospores, a short chain, or a single elongate cell. Verticils not developed, the apical por- tion of a pseudomycelial cell not producing more than two opposite blasto- spores (Fig. 56). Gelatin not liquefied. Key to Species Colony yellowish, becoming rose color. M. rosea. Colony not becoming rose color. Maltose fermented, cutaneous lesions. M. Shutetzlcyi. Maltose not fermented, intestinal tract. M. parapsilosis. Action on maltose unknown, nails. M. onychophila. Mycocandida rosea (Zenoni) Dodge, n. comb. Oidium roseum [non liquefaciens] Zenoni, Lo Sperimentale 66: 33-66, 1912. Monilia rosea Castellani & Chalmers, Man. Trop. Med. ed. 2, 829, 1913. MyceloUastanon roseum Ota, Jap. Jour. Derm. Urol. 28: [4], 1928. Candida rosea Almeida, Annaes Fac. Med. Sao Paulo 9: 11, 1933. Isolated from a fatal case of hepatitis. Pathogenic for rabbits. Colony yellowish, rounded, granulose, waxy, shining. On agar after 12-15 days, there is a definite odor and distinct rose coral color. In plain broth, floccose sediment. In brain broth, liquid becomes turbid with white powdery sediment and yellowish white flocci. Gelatin not liquefied. Mycocandida Skutetzkyi (Ota) Dodge, n. comb. Cryptococcus Skutetzkyi Ota apud Nannizzi, Tratt. Micopat. Umana [Pol- lacci] 4: 331, 1934. Cryptococcus de Skidetsky Sa.sakawa, Centralbl. Bakt. I, 88: 273, 1922; Ota, Ann. Parasitol. Hum. Comp. 2: 53-55, Fig. 10, 1924. Isolated from cutaneous lesions, case never published. 294 MEDICAL MYCOLOGY Yeast cells spherical, 5/a in diameter, or ovoid with some raquet mycelium, producing single blastospores at the septa. Mycelium not highly developed, hyphal cells 5/* in diameter, walls somewhat thickened. On malt agar, colony grayish or yellowish, smooth, center granular, with slight rays or fine folds. On malt extract, producing a thick grayish ring with sediment. By Lindner method, reported to ferment glucose, fructose, mannose, galactose, and maltose, but no action on sucrose, lactose, or raffinose. Myco Candida parapsilosis (Ashford) Dodge, n. comb. Monilia parapsilosis Ashford, Am. Jour. Trop. Med. 8: 518. 1928. Candida parapsilosis Camargo, Agentes Etiologicos do Sapinho 58, 1934. Nonpathogenic to guinea pigs and rabbits. Produces growth of fir tree shape along gelatin stab. Maltose not fer- mented. Gelatin not liquefied. Fig. 56. — Mycocandida, showing branching and reduced verticils. (After Langeron & Talice 1932.) Mycocandida onychophila (Pollacci & Nannizzi) Langeron & Talice, Ann. Parasitol. Hum. Comp. 10: 58, 1932. Monilia onychophila Pollacci & Nannizzi in Marengo, Archivi di Biol. 3: 4:25-36, 2 fi.gs., 1926. Mycotorula onychophila Ciarrocchi, Giorn. Ital. Derm. Sifilol. 74: 415-429, 3 pis., 1933. Isolated from paronychomycosis. Probably organism of Connor (1933) should be referred here. In pus cells spherical hyaline 3-5/^, same cells seen in nails. Cells spherical or slightly ovoid uniguttulate, solitary or in chains, mostly 3.5-5iU in diameter; in older cultures mycelium present. Somewhat sparingly branched, septate, hyaline, 2-S/x in diameter, branches cylindric. terminal cell I ( I EREMASCACEAE IMPERFECTAE 295 clavate, bearing a chain of blastospores, 3-8 cells long ; blastospores spherical, ellipsoid, or citriform, uniguttulate, hyaline, 3.5-6^ in diameter, or 3.5-6.5 x 3-5/x. On Sabouraud agar, colony creamy, surface smooth and shining, whitish tending to yellow in age. On Pollacci agar, creamy, but the surface is slightly granular, yellowish becoming brownish. On potato and carrot, creamy whitish, less shining than on agar media. On Raulin's liquid medium at 20° C, slight pellicle and ring with slight sediment. On broth, ring more highly developed, pellicle not reported, sediment without turbidity, glucose and fructose fer- mented (other sugars not reported) ; gelatin liquefied. Mycocandida mortifera (Redaelli) Langeron & Talice, Ann. Parasitol. Hum. Comp. 10: 58, 1932. Candida mortifera Redaelli. Isolated from the lungs. Mycelium, septate, 10-20 x 2-2. 5/*; blastospores spherical or ovoid, in chains of 3-6 cells, 4-5/i. in diameter ; clilamydospores abundant in old cultures. On carrot agar, colony creamy, light yellowish, convex, margins sinuous. On carrot, colony elevated, center convex, margins lobed, white or slightly cream colored, opaque. Giant colony on malt gelatin (30 days) plane, white or slightly yellowish with three concentric zones and denticulate margins. On malt extract, an incomplete, irregular, semitransparent ring, no pellicle and slight, floccose, powdery sediment. On glucose broth, no ring or pellicle, slight sediment. Glucose, fructose, sucrose, mannite, galactose, and lactose fer- mented ; maltose not fermented ; no reduction of methylene blue ; milk not clotted; gelatin liquefied. PSEUDOMONILIA Pseudomonilia Geiger, Centralbl. Bakt. II, 27: 97-149, 2 pis., 4 figs., 1910. The type species is Pseudomonilia alhomargi^iata Geiger. Cell shape variable in young cultures, sprout cells in old cultures ; more or less branched mycelium formed of elongate sprout cells but no true septation ; giant cells common in old cultures. Pellicle well developed, very little sedi- ment; no ascospores, no alcoholic fermentation, sugars variously assimilated; gelatin not liquefied. Key to Species Milk not coagulated. P. matalensis. Milk coagulated. Acid formed in lactose. P. inexpectata. Acid formed in sucrose. P. alessandrina. Pseudomonilia matalensis (Castellani) Dodge, n. comb. Oidium mafalense Castellani, Lect. on the Higher Fungi in Relation to Human Pathology. R. Coll. Phys. 1915 ; Castellani & Chalmers, Man. Trop. Med. ed. 3, 1096, 1919. 296 MEDICAL MYCOLOGY Mycoderma mataleiise Brumpt, Precis Parasitol. ed. 3, 1084, 1922. Pseudomycoderma matalense Ciferri, Arch. Protistenk. 71: 436-443, 1930. Geotrichum Matalense Castellani, Jour. Trop. Med. Hyg. 35: 278-279, 3 figs., 1932. Isolated from ulcers and from sputum of cases of bronchitis in the tropics (Ciferri 1930). Mycelium abundant on agar, little branched, septate, 2-5/* in diameter, generally 3-4/*, blastospores sprouting, 5-15 x 2-4/i, generally 10 x 3/t, isolated or sprouting to form chains. Colony effuse, white on agar and gelatin. On liquid media, producing islets promptly white, cottony above, mucus, yellowish below, finally becom- ing rose violaceous pellicle, practically no sediment. Sucrose not inverted, milk slightly acidified, no fermentation of sugars, no liquefaction of gelatin. There seems little to distinguish var. Chapmani Castellani, Jour. Trop. Med. Hyg. 35: 278-279, 1932. Pseudomonilia inexpectata (Mazza, Niiio & Egiies) Dodge, n. comb. Monilia inexpectata Mazza, Nino & Egiies, Bol. Inst. Clin. Quirurg. Univ. Buenos Aires 5: 284-288, 1930. Mycocandida inexpectata Talice & Mackinnon, Reunion Soc. Argentina Patol. Reg. del Norte in Santiago del Estero 8: 164, 1934. Isolated from the drops of pus expressed from inflammatory processes in- volving the fingers, particularly around the nails, in a woman, a native of Buenos Aires, Argentina. Ordinary medicaments of no avail. Intraperitoneal injection to guinea pig and mouse produced abscesses. On Sabouraud glucose agar at 37° C, or room temperature, in 24-48 hours, abundant growth, colonies cream white, dry in appearance, opaque surface, not adhering to substrate ; after 5 days several colonies about 1 cm. in diam- eter, center smooth, elevated, margins deeply folded. Growth similarly abun- dant on simple agar and broth media. On agar, slight growth in 24 hours. On potato with 8% glycerol, good growth in 24 hours, colony cream-colored, dry in appearance; no pigmentation of medium. On Drigalski-Conradi me- dium, colony grows without modification of medium at first, then slight evolu- tion of gas. On Gougerot gelatin medium, growth good. On coagulated human serum, organism grows Avith digestion of the part of medium disturbed by puncture. In Sabouraud glucose broth, turbidity and sediment, with for- mation of creamy white pellicle which ascends walls of tube. In plain broth, uniform turbidity and sediment after 24 hours. In Raulin's liquid, turbidity, sediment, and pellicle ascending sides of tube. No fermentation of sugars. Acid produced in maltose, lactose, dextrin, and glucose. No indol. Milk coagulated in 24-48 hours. Gelatin not liquefied. Indistinctly positive or negative to Gram's stain; with Gueguen's mixture preparation colored intense blue with tiny points of brick red, probably cor- EREMASCACEAE IMPERFECTAE 297 responding to fat droplets stained with the Sudan II. With "panoptic" stain, yeast elements blue, hyphae part blue and part purplish red, giving a laminated appearance. Pseudomonilia alessandrina (Panayotatou) Dodge, n. comb. Monilia alessandrina Panayotaton, Jour. Trop. Med. Hyg. 33: 17-18, 1930. Isolated from the mucopurulent sputum in a case of bronchitis. Medicated with KI. Intravenous injection into rabbit caused generalized infection and finally death. Organism reisolated. Cells yeastlike, 3-4/a in diameter. Gram-positive. On Sabouraud agar, abundant white growth. Wrinkled colonies on beef agar. Pellicles formed in maltose and sucrose. Acid formation with glu- cose? ["gelose'^'], maltose, sucrose, dextrin, raffinose, galactose, and dulcite. No fermentation of any medium. No indol or HoS formed. Milk acidified and coagulated on fourth day. Coagulated serum and gelatin not liquefied. Doubtful Position The following species have been so vaguely described that I have not been able to refer them even doubtfully to genera of the Eremascaceae Imper- fectae, or I have been unable to locate a copy of the original descriptions. Monilia Botrytis Vuillemin, Champ. Parasit. Homme Anim. 87, 1931, Botrytis sp. Auclie & LeDantec, Arch. Med. Exp. I, 6: 853-861, 1894. Botrytis pyogenes Fayod, 1894. Imperfectly described and not easy to place. Cultures lost. Beurmann & Gougerot, Arch, de Parasitol. 15: 88, 1910, think it is a Sporotrichum. Candida pinoysimilis Castellani, Jour. Trop. Med. Hyg. 36: 312, Fig. 37, 1933. Cryptococcus pinoysimilis Castellani, Med. Press Circular 136: 441, 1933. Isolated from the surface of a blastomycetic, verrucoid lesion, patho- genicity doubtful. Cells spherical or ovoid, 3.5-7/^. Some hyphae noted. On glucose agar, smooth, white. Litmus milk made slightly alkaline, gelatin and serum not liquefied. Sugar fermentation doubtful [reports fer- mentation of glucose, fructose, and maltose in text, denies it in accompanying table] . Acid produced in glucose, fructose, sucrose, lactose, glycerol, inulin, and perhaps xylose. Saccharomyces intertrigos Touraine-Desvaux, Contr. etude des epider- momj^coses a levures . . . These Fac. Med. Paris 203: 1-44, 1923. [Unnamed organism] Gougerot & Gancea, Bull. Soc. Derm. Syphiligr. 25: 335-339, 1914; Hudelo & Montlaur (1914). Cryptococciis de Gougerot et Gancea Ota, Ann. Parasitol. Hum. Comp. 2: 43-44, 1924 Cryptococcus Gougeroti-Ganceae Ota apud Nannizzi, Tratt. Micopat. Umana [Pollacci] 4: 309, 310, 1934. 298 MEDICAL MYCOLOGY Isolated from an interdigital mycosis of the feet, persistent over 2 years under a variety of treatments. Intravenous injection into rabbit produced rapidly fatal septicemia or pyemia with metastatic abscesses and generalized peritoneal infection. Hyphae, pseudomycelium, and sprout cells present. Colonies white, spreading, creamy, shining, smooth or with center slightly folded and surrounded by a smooth zone. Monilia Montoyai Castellani & Chalmers, Man. Trop. Med. ed. 2, 1514, 1913 (nomen nudum). Isolated from red pinta. Monilia pulmonalis (Plant) Sartory, Champ. Parasit. Homme Anim. 709, 1922, non Castellaiii. Parendomyces pulmonalis Plant apud Mautner, Centralbl, Bakt. I, 74: 207- 208, 1914. [Case history in Wiener Med. Woch. 64: Col. 1055-1067, 1914.] Candida pulmonalis Almeida, Annaes Fac. Med. Sao Paulo 9: 11, 1933. Mycelohlastanon pulmonare Ota, Ja]3. Jour. Derm. Urol. 28: [4], 1928 (per- haps Monilia pidmonalis of Castellani). Monilia Mautneri Nannizzi, Tratt. Micopat. Umana [Pollacci] 4: 366, 367, 1934. Isolated from sputum. Not pathogenic to rabbit. Spores smaller than in Monilia Candida Bonorden. No mycelium. Monilia Samboni Castellani, Proc. R. Soc. Med. (Derm.) 6: 36, 1912. Saccharom.yces Samboni Castellani & Chalmers, Man. Trop. Med. ed. 2, 768, 1913. Isolated in intertriginous dermatitis of scrotocrural and axillary regions. Skin reddens with possibly slight exudation, borders marked but not elevated. Little pruritus. Treatment : KMn04 solution, 1 :4,000, or resorcin, 1 :100, followed by dusting with a powder composed of boric acid and talc. Cells 6-8^ in diameter. Colonies white, rapidly confluent. Monilia vaginalis Escomel, Bull. Soc. Path. Exot. 17: 922, 1924. Endomyces sp. Carpano, Ann. Ig. Sperim. 22: 435-450, PI. 10, 1912. Isolated from pleuropulmonitis in a horse. Disease reproduced in rabbit and guinea pig. In liquid from lungs, cells very variable in size and shape, reaching a maximum length of 20-25/a, usually 16-20 x 5-7/*, spherical or ovoid cells about 4-6/x, longer cells often cylindric or nearly so, often united into groups of 2 or 3, or more rarely into short simple or branched hyphae up to 200/x long. Attempts to isolate were unsuccessful on most media tried, due to an ac- tively growing Streptococcus, but cultures on potato and potato agar were successful. On potato, colony shining, dirty gray becoming brownish in age. On potato agar, whitish, bright, then graying as the colony thickens. Cells spherical to slightly ovoid, 3-4/a in diameter in 15 days. Characters intermediate between Saccharomycopsis guttulatus and Zymo- nema farciminosum. EREMASCACEAE IMPERFECTAE 299 Oidium sp. Coley & Tracy, Jour. Med. Res. 16: 237-249, Pis. 18-20, 1907. Perhaps Zymonema dermatitidis. TJiis organism shows much mycelium, somewhat suggesting Madurella or IScopulariopsis. It is not yeastlike. Monilia sp., Flu & Woensdregt, Meded. Burgerl. Geneesk. Dienst Nederl. Indie 6: 1-26, 6 figs., 1918. Isolated from blastomycosis of the central nervous system. Pathogenic to guinea pigs. In spinal fluid, cells 6-7/x in diameter, spherical, in chains of 3-8. Organ- ism gram-positive. No spores found in culture. On Sabouraud agar in 3 days, at 26° C, colony opaque, circular, smooth, shining, moist, 1-2 mm. in diameter, china white, hemispheric, becoming yel- lowish white or yellowish brown. In broth, liquid remains clear with slight sediment and pellicle in about 10 days. Pellicle crumbles and settles after a time. No fermentation of sugars. Parendomyces sp. Launois et Pinard, Soc. Path. Comp. 1912. Monilia de Launois et Pinard Sartory, Champ. Paras. Homme Anim. 708, 1922. [Monilia sp.] Williams, Med. Jour. Australia 2: 185-187, 4 figs., 1922. Isolated from cerebrospinal fluid in case of systemic blastomycosis. Patho- genic for rabbit. Cells in tissue somewhat elongate, sprouting from ends. In cultures only spherical cells observed. Colony yellowish white, sticky, coalescing and running down surface of slant. Broth cultures turbid, appearing as if a gel were present. This organism is possibly Zymonema capsulatum, but was transferred too frequently to secure hyphal formation. Not described fully enough to permit of definite placing. CHAPTER XII SACCHAROMYCETACEAE The Saeeliaromycetaceae, or true yeasts, may be regarded as direct de- rivatives of the Eremascaeeae where growth of the thallus by sprouting has become almost universal. Some species, such as Saccharomy codes Ludwigii, Deharyomyces Kloeckeri, and Zygosaccharomyces Priorianus, are still able to form true hyphae on gelatin substrates, but the hyphae are unstable and with a slight alteration of the medium break up into sprout mycelia. The sprout cells are generally spherical or ellipsoid and hyaline, the size varying according to medium and age. Under unfavorable conditions they store fat and glycogen and form a double membrane. These resting cells, which are very resistant to environmental changes and carry the organism over unfavorable periods, are probably chlamydospores. At germination each cell ruptures the outer fragile wall and grows into a sprout mycelium. On the basis of their method of cell multiplication, the yeasts are often divided into two tribes. In the Schizosaccharomyceteae, each cell elongates, abjoints two equal daughter cells which round off, separate, and again abjoint. At present only a few species are known. In the Saccharomyceteae, sprout cells begin as small lateral protrusions of the motlier cell ; they are abjointed, increase in size, and finally attain the appearance of the mother cell. This tribe contains most of the true yeasts. The tribes are not absolutely distinct since Saccharoniycodes Ludivigii usually divides as in the Schizosaccharomyce- teae but occasionally multiplies by sprouting, although the sprout cells are produced only at the poles of the mother cell, never laterally. If sprouting or division occurs rapidly, the cells may cling together in small colonies or, in occasional old cultures, in filaments. Under certain conditions, as in age, with the exhaustion of nutrient, or on solid substrates as gypsum blocks, asci with ascospores are formed. In the wild yeasts the number of ascospores varies from 1 to 12 ; in many industrial yeasts, certain numbers predominate; e.g., in Schizosaccharomyces octosporus 4 or 8, in Saccharomyces cerevisiue 4, and in S. Pastorianus 2. The spores are spherical to ellipsoid and either smooth or rough. Ordinarily the spore wall has only one layer, but in SaccJiaromycopsis guttulatus, isolated from feces, there are two layers, the outer of which ruptures on germination. In 3-day cultures of Schizosaccharomyces octosporus, the cells copulate in pairs by short tubes (Fig. 57, 1-6). Where the cells are in short chains, the separating wall between sister cells may dissolve (adelphogamy). Two nuclei migrate into the copulation canal and fuse; the copulation canal broadens and each of the two individuals develops into a barrel-shaped struc- ture, where, within approximately half an hour, after three or more, rarely 300 SACCHAROMYCETACEAE 301 tAvo, nuclear division, there appear 8 — rarely 4 — spores (Fig. 57, 7-11). Often the eopulatory canal is narrow so that the young ascus is shaped like a dumb- bell between whose ends are divided the 8 spores. At germination, the spores in the ascus swell, rupturing the ascus wall. Each spore thus liberated is divided by a septum into two daughter cells. These cells continue to divide similarly, and in some liquid media they form short branched filaments. Under unfavorable conditions, copulation may occur earlier, so that al- ready in the ascus the ascospores copulate with ascospores of the same or a neighboring ascus. In these cases the resulting zygote becomes an ascus di- rectly without first forming a sprout mycelium. Other strains have a tendency to become asporogenous; although they may still form eopulatory processes, which may be functionless. In very rare instances, there seems to be a par- Fig. 57. — Schisosaccharomyces octosporus. Copvilation and development of asci (X750). (After Guilliermond 1905, 1917.) Fig. 58. -Zy gosacchuromyces Chevalieri siiowing stages of copulation. 1913.) (After Guilliermond thenogenetic development of spores — 4 to an ascus. Copulation is normally isogamous and the gametangial copulation is replaced by pseudogamy. This also disappears and the cells, after unfruitful attempts at copulation, may change parthenogenetically to asci. In 8chizosaccliaromyc.es Pomhe, an Afri- can beer yeast, and in S. mellacei, from rum factories of Jamaica, many aspor- ogenous strains are known, and in S. asporus, from arrack in Java, spores have never been observed. In the Saccharomyceteae, we have two series of degeneration, one isog- amous characterized by a smooth ascospore, the other heterogamous with a rough one. In the simpler forms of the smooth-spored series, as in Zygosac- charomyces, 2 cells form eopulatory processes toward each other, the nuclei migrate into the bridge and fuse (Fig. 58, 1-3). This diploid nucleus divides, both daughter nuclei migrate back into the copulating cells, there dividing a second time forming 2 spores in each copulating cell. The whole cell, similar 302 MEDICAL MYCOLOGY in shape to a dumb-bell, is an aseus (Pig. 58, 4-5). In some cases, at least in Zygosaccharomyces Priorianus, copulation is absent, and the individual sprout cells develop parthenogenetically to asci. Rarely, in small colonies where the mother cell lacks other cells, it may copulate with a daughter cell in pseudogamy. That which is anomalous in Zygosaccharomyces Priorianus becomes increas- ingly common in other species ; as sexual tendencies weaken, copulation becomes more and more difficult. Thus in some species, such as Z. niongolicus, although copulation does occur, most of the asci develop parthenogenetically after put- ting forth copulation tubes which fail to function. Fig-. 59. — Torulaspora Rosei. Development of asci (XI, 800). (After Guilliermond 1912.) Fig. 60. — Saccharomyces cerevisiae. 1-i, formation of spi-out cell with amitosis of nucleus (XloOO); 5-10, development of sprout cell to ascus and g;ermination of ascospore (X750). (After Guilliermond 1902, 1904.) Torulaspora shows still further degeneration toward parthenogenesis. On favorable media, the sprout cells of T. Delhrucki form numerous sprout cells which attempt fusion (Fig. 59). Occasionally this is successful and ascospores are formed, but generally the separating wall is not dissolved, and each cell forms 1-4 ascospores parthenogenetically. AVhile copulatory processes are formed in T. Rosei and T. fermentaii, they ntver fuse, and parthenogenesis is complete. Finally complete parthenogenesis witli no morphologic suggestion of sex is reached in the large genus Saccharomyces (Fig. 60). This genus furnishes most of the common yeasts used in alcoholic fermentation, such as S. cerevisiae, SACCHAROMYCETACEAE 303 the beer yeast, S. ellipsoideus, the wine yeast, etc. A few species have been re- ported as pathogenic, but generally they are rather imperfectly described. The heterogamic series producing rough ascospores is less known and has not degenerated quite so far. In Nadsonia fulvescens ascospore formation does not generally occur within the larger fusion cell; on the side opposite the copulatorj^ canal, a sprout cell receives the united contents of both fusion cells and develops 1 or 2 brown ascospores (Fig. 61). Thus we find a trace of ascus formation characteristic of the Eremascaceae, as also cell division intermediate between that of the Schizosaccharomyceteae and normal sprout- Fig. 61. — Nad^miia fulvescens. 1-3, copulation of mother and daughter cell; i, 5. development of ascus; 6, ascus set free with ascospore. (After Guilliermond 1928.) Fig. 62. — Debaryomyces Kloeckeii, development of asci. (After Guilliermond & P6ju 1920.) ing. In N. Richteri occasionally the male cells are not present on liquid media. In this case long copulation tubes are formed. In Deharyomyces, traces of a separate ascus formation have disappeared. While copulation is typically heterogamous, we find increasing parthenogenesis. In D. glohosus, from earth in the Virgin Islands, about 75% of the asci are parthenogenetic. Occasionally copulatory processes are formed, but they grow past each other, apparently for lack of sexual attraction. Copulation occurs in about one-fourth of the cases. This may take place between two mature individuals, where the zygote nucleus either divides in the bridge and returns a daughter nucleus to each fusion cell or migrates before division into one fusion cell and there forms a spore (Fig. 62, 3, 4). Copulation may also 304 MEDICAL MYCOLOGY occur between a mother cell which is still sprouting- and a young daughter cell, in which case the contents of the daughter cell migrate back into the mother cell and form spores. In other species this becomes the rule as in D. mucosus (Sartory, Hufschmitt & Meyer 1930). Debaryomyces frequently oc- curs in cutaneous infections. In Saccharoniyces (Fig. 60) all trace of sexuality has been lost. Finally, we should describe a phenomenon whose phylogeny is not clear. It seems to be present in several genera which otherwise do not appear closely related. When Zygopichia Chevalieri of the Pichiaceae is insufficiently nour- ished, 2 ascospores may copulate after they have swollen and ruptured the ascus wall, or an ascospore may copulate with a sprout cell and change to a l-spored ascus, or the ascospore may function without copulation, as an ascus and form more ascospores. In Williopsis Saiurnus, Saccharoniyces ellipsoideus, 8. Chevalieri, 8. Mangini, and 8. annulatus, this phenomenon is common. On germination some spores develop the normal sprout mycelium, while others copulate in pairs. The zygote begins to sprout, forming an apparently diploid sprout mycelium in the copulatory canals, occasionally also on the whole sur- face of the copulating cells. Later, Avithout further sexual processes, the asco- spores develop in the vegetative cells. Finally, we have the curious genus Saccharomy codes, whose sole species produces asci with 4 spores which lie in pairs at the two poles (Fig. 63, 1, 2). The spores at a given pole result from the division of one nucleus, thus they are sister cells and are connected by a protoplasmic layer remaining from the periplasm. At germination they swell much in the ascus and form small copulatorj'- canals (Fig. 63, 1-4). Occasion- ally several spores fuse. Two cells of the same sexual tendency attract a third of a different tendency. Copulation may be retarded and the tubes continue to grow, perhaps rupturing the ascus wall. The copulatory processes may fuse with spores from the other pole of the ascus ; or some spores may abort, in which case fusion with spores of another ascus is possible (Fig. 63, 9). Very rarely in old cultures, spores may germinate parthenogenetically and develop special strains which continue to reproduce parthenogenetically through several generations at least. After fusion of the copulatory processes, the nuclei migrate into the bridge and fuse (Fig. 63, 5, 7) . Occasionally fusion may occur in one of the spores instead of in the canal or may be retarded and occur only in the germ tube which grows out of the copulatory canal, ruptures the ascus wall, and germinates to a sprout mycelium. Guilliermond (1931) suggests that these forms in which copulation between ascospores occurs, may be derived from the Taphrinaceae where Juel and others observed copulation between sprout mycelium from germinating ascospores. It is un- certain whether this copulation of ascospores has phylogenetic significance, since we find it among widely divergent groups, such as Ascoidea, Williopsis, and Saccharoniyces. In Ascoidea, Walker believes it is purely a vegetative fusion. SACCHAROMYCETACEAE 305 Determination of Species. — Two media have been widely used iu studies of yeasts : malt extract and yeast decoction. The former is prepared as fol- lows (Guilliermond 1928) : Barley is germinated on moist blotting paper on plates. When the grain has swollen and the radicle has begun to emerge, it is dried in an oven at 30° C, and ground in a mortar. Two hundred grams of powder are stirred into a liter of cold water which is slowly heated to 60°, and this temperature is maintained for 45 minutes, with occasional stirring of the mixture. Four grams of hops are added and the whole boiled for about an hour. The result- ing decoction is then sampled, the amount of maltose determined, and the whole diluted to bring the concentration of maltose to 3%. It is then sterilized in the autoclave at 120° C. The decoction is usually used to prepare either 1.5% agar or 6-15% gelatin. The malt extracts now on the market seem suitable and have yielded good results in our laboratory. The preparation of media from them is much less tedious. Yeast decoction is prepared by boiling and stirring 100 gm. fresh yeast in 1 liter of distilled water, filtering, and autoclaving. The liquid furnishes a small amount of available nutrient so that some sugar is often added. j?ig. 63. — Saccharomy codes Ludioigii. Copulation and development of asci (X750). (After Guilliermond 1905.) The macroscopic characters on malt extract incubated for 24 hours at 25° C. are studied. Some yeasts form a white sediment in the bottom of the test tube, while others grow mostly at the surface, forming a pellicle or, if they creep up the sides of the tube, a ring. If the tube is not handled care- fully, the pellicle may break up and settle as a sediment, but if left alone a new pellicle will soon form. Often the pellicle is characteristically folded, etc. Fermentation should also be observed if present. The microscopic characters are more variable. One should observe the general shape and size of the cells, the method of budding, whether only terminal or lateral, whether a pigment is characteristically produced. For variation in appearance of cells of different ages, the following account is summarized from Shrewsbury (1930) : The adolescent cell (Fig. 40, 1-3) is spherical to allantoid in shape with a thin cell wall and refractile, homogeneous cytoplasm. Sprouting is usually most active at the poles, but may occur anywhere. The mote cell is larger than the adolescent cell, and the cytoplasm more granular. A single vacuole, 306 MEDICAL MYCOLOGY rarely more, fills about half the cell, and highly refractile corpuscles or motes (probably metachromatic granules) showing Brownian movement are seen in the vacuole. After these appear in culture, growth slows down, and reserves of fat and glycogen are stored in preparation for formation of ascospores or of resistant cells. The adult or durable cell (Fig. 40, 4 and 5) often persists unchanged in cul- tures for months. This cell is larger with a vacuole, empty or containing a single oil globule, occupying three-quarters of the volume. Fat is usually stored in a large polar globule, occasionally as small globules around the vacuole. The globules consist of an outer layer of albuminous substance en- closing a semifluid emulsion of fatty substances. Shadow or senescent cells are empty cell walls from which the contents have disappeared by rupture or otherwise (Fig. 40, 6-9). The permanent cell is often still larger with a dark-colored wall thinner than that of the normal cell. Vacuoles are usually absent, though occasionally present, dark granules are present and, usually, fat globules also. Sometimes there results a pseudomycelium (Fig. 40, 10) composed of adolescent cells, nonvacuolated cells with fat globules, and cells having finely granular cytoplasm — the latter cells never seen save in pseudomycelium. Oc- casional bizarre forms occur whose shapes are usually the result of multiple abnormal budding. In the rare red yeasts even conidia are produced, especially in the genus Sporoholomyces, whose method of conidial discharge sometimes causes it to be placed in the Basidiomycetes, a very distantly related group of fungi. Physiologic characteristics should be noted, such as temperature limits of sprouting, optimum temperature for growth as determined by colony size, the temperature limits of formation of pellicles, etc. Sporulation should be induced if possible. The media most commonly in use are gypsum blocks and Gorodkova's agar. The yeast is cultivated for 24 hours on malt extract agar; the growth is then scraped off and placed on a short cylinder of plaster of Paris by means of a platinum spatula. The cylinder is placed in a sterile Petri dish, filled to about half the height of the cylinder with sterile water. Spores often appear in 1-8 days. Gorodkova's* agar may also be used. It consists of: distilled water 1,000 c.c, agar 10 gm., peptone 10 gm., beef extract 10 gm., sodium chloride 5 gm., glucose 2.5 gm. This enables the yeast to start growth, but the nutrients are quickly ex- hausted and sporulation often results in 1-8 days. Some species which do not form spores on either of these media will produce them on carrot in very old cultures. Maneval (1924) reports spore formation in cultures kept in the ice box. Once the asci have been obtained, their form, method of formation, and the shape and size of their spores should be observed. It is often very difficult •Maneval (1924) omits the peptone and reduces the Liebig meat extract to 3 gm. in this formula. He also recommends 15-20 gm. of agar. SACCHAROMYCETACEAE 307 to distinguish spores from the large oil globules found in many yeasts. Spores are usually much more uniform in size and do not stain readily with Sudan III or osmic acid. The method of IMoeller (Hufschmitt, Sartory & Meyer 1931) is also sug- gested : Treat for 10 seconds to 5 minutes in 1% sulphuric acid, wash, stain with carbolfuchsin, heating for 1 minute, differentiate with 5% sulphuric acid for 5 seconds ; wash, counterstain with aqueous methylene blue for 3 minutes. Also the method of Schumacher (Buschke & Harry 1923) : Fix, stain for 1 minute in carbol-methylene blue, rinse with distilled water, stain 1% minutes with slow movement or slide with 1% phosphin (diamidophenylacridin). Spores also stain with Ziehl-Neelsen acid-fast procedure if the sulphuric acid is replaced by 1% nitric acid in alcohol. Fatty acids may be stained blue and neutral fats red by the technic of Smith (1907). For further methods see Shrewsbury (1930). The germination of the ascospores should be noted, if possible, especially whether there is copulation between them. Many species form very characteristic giant colonies, while others do not. Some also produce rather characteristic liquefactions of gelatin. Among the pathogenic yeasts, Fontoynont reports very good results in dressing the lesions with methylene blue ; he also recommends 0.1 gm. methy- lene blue per diem administered per os, stating that it is tolerated for several months at a time. Key to Genera Cells dividing by septa, sprouting absent or rare, asci usually 4-8-spored. „„,..,. , ,. ^ , , Schizosaccharomyces. Cells dividing by sprouting, septa absent. Spores rough-walled, copulation usually lieterogamic. Ascus sprouting from the zygote, cell division sometimes suggesting that of Schizo- saccharomyces. Nadsonia. Ascus developing directly in the zygote. Debaryomyces. Spores smooth, copulation usually isogamic or absent. Ascospores thick-walled with a double membrane. Saccharomycopsis. Ascospores thin-walled without a double membrane. Copulation between vegetative cells prior to ascosporc formation, parthen- ogenesis rare. Zygosaccharomyces. Copulation produced but usually not functional, ascospores commonly resulting from parthenogenesis. Torulaspora. Copulation absent. Soccharomyces. Copulation between ascospores while still in the ascus. Saccharomy codes. SCHIZOSACCHAROMYCES Schizosaccharomyces Lindner, Woch. Brauerei 10: 1298, 1893. The type species is Schizosaccharomyces Pomhe Lindner. Cells cylindric or elongate ellipsoid; vegetative division by fission not by sprouting; asci arise after isogamous copulation; spores spherical or ovoid 308 MEDICAL MYCOLOGY with smooth wall which is colored blue by KIJ, ; 4-8 spores per ascus. Sedi- ment in malt extract, occasionally also a ring is formed. Various sugars strongly fermented ; nitrate not assimilated, no growth with ethyl alcohol as a substrate. So far there has been no case of pathogenicity for mammals which has not been shown to be wrongly referred here. Apparently Bacillus megathe- rium has been frequently picked up as a contamination and referred here, the granule or oil globule being mistaken for a small spore. All reports of this genus with spore number fewer than four, especially if the cell size is small, should be studied very critically before admission to this group. Schizosaccharomyces hominis Benedek, Centralbl. Bakt. I, 104: 291-303, 3 pis., 1927. See also other papers by Benedek. Dorrepaal (1930) shows conclusively that this organism is a bacillus either identical with or closely related to Bacillus megatherium BarJ^ DEBARYOMYCES Deharyomyces Kloecker, C. R. Trav. Lab. Carlsberg 7: 273, 1909. The type species is Deharyomyces glohosus Kloecker. Cells predominantly spherical but also ovoid, mostly small ; vegetative reproduction by sprouting. Asci following isogamous or heterogamous copula- tion occasionally parthenogenetic ; spores spherical mostly with a large central oil globule, wall verrucose (warts often difScult to see), usually only 1 spore per ascus. In malt extract, producing sediment, usually a ring and occasion- ally a pellicle. In most species no fermentation, very few species having a good fermentation ; nitrate assimilation negative ; with ethyl alcohol as a substrate, growth good, often producing a thin pellicle on this medium. Key to Species Glucose and sucrose strongly fermented, slight ring but no pellicle. Colonies becoming bister or darker on agar. D. Hitdeloi. Colonies remaining wliite or light colored. D. mucosus. Colony color unknown. Asci, 2-5m in diameter. D. emphiisematosus. Asci, 5-7ai in diameter, fermentation questioned. D. Gruetsii. No fermentation or a weak fermentation of glucose only. [Ota reports slight fermentation of some other sugars, probably either due to impurities or to the nietliods used.] Eing formed on malt extract but no pellicle. Sucrose not inverted, cells 2-3.5 x 3-7jU,, ovoid, occurring in chains in old cultures. D. KloecTceri. Sucrose slightly inverted, cells 2-8 x 2-6/a in groups springing from the same mother cell. D. Nadsoni. Sucrose inverted. Cells 3.6-5.4 x 1.8-3ai, cells single or in pairs. D. Matruchoti. Cells 2-6iJi rarely 8-10^, rarely also in chains. D. Fabryi. SACCHAROMYCETACEAE 809 No ring or pelliclo on malt cxtruct, sodinKMit brownish white, sucrose inverted. D. Leopoldi. D. Lundsgaardi. D. Laedcgaardi. D. Hildegaardi. Pellicle on malt extract, sucrose inverted. D. G^Ulliermondi. Fermentation unknown, probably close to D. Fahryi. D. Burnirri. In the following descriptions I have tried, to translate and bring together the salient points of each organism nnder the original name. Ota's descriptions of numerous new species seem to have been too hurriedly written without a sufficient background to appreciate the stability of characters and with too little biometric data to give much validity to his cell measurements. Since most of the cultures had been isolated many years previous to his studies and been carried in stock cultures for so long, it is possible that they may not be the original organism isolated from the cases mentioned. Quite probably Dekker is right in referring most of his species to D. Matruchoti. On the other hand, it seems strange that a single species of yeast should produce such dif- ferent types of lesions as Ota's A'arious species have been reported to do. Debaryomyces Hudeloi (Gougerot) Fonseca, Brasil Med. 36: 101-102, 1922. Cryptococcus sp. Hudelo, Duval & Laederich, Bull. Mem. Soc. Med. Hop. Paris 23: 723-734, 1906. Atelosaccharoniyces sp. Duval & Laederich, Arch, de Parasitol. 14: 224- 318, 1911. Atelosaccharomyces Hudeli Gougerot, Paris Med. 1: 462, 1911. Debaryomyces KloecJieri var. Hudeloi Dekker, Verhandel. K. Akad. Wet- ensch. Amsterdam Afd. Natuurk. 28: 473, 1931. Isolated from blastomycosis with multiple foci, clinically more or less similar to that produced by Atelosaccharomyces hominis. Pathogenic for mice, less so for rats and guinea pigs, not for rabbits, dogs, or hens; subcutaneous inoculation in guinea pig produced abcesses, which healed spontaneously in a fortnight. Cells usually spherical, occasionally ellipsoid, isolated or in pairs, very rarely in short chains of 4-5 cells, mostly 2-4/* in diameter, with ellipsoid cells up to 8/* and giant cells 16 x 6/x [original description states cells 2-20/*] mostly uniguttulate (Fig. 64, 1-3). Copulation heterogamous, asci 1-spored, asco- spores 2-3^ in diameter, sparingly echinulate with a large fat globule (Fig. 64,4). Optimum temperature 22° C, growth very good at 38° C, not anaerobic. Ota reports that after 18 years oplnnum temperature is 30°, growth poor at 37°, good at 12° C. On Sabouraud glucose agar, colonies opaque, smooth, moist, shining, margin regular becoming undulate, consistency mucous, viscid, strongly ad- herent to the agar but not penetrating it, 1.5 mm. thick, creamy, growth not spreading, finally becoming yellow bister, then cafe-au-lait, and deep brown 310 MEDICAL MYCOLOGY after 8-10 months. Similar, but less luxuriant on lactose, maltose, and sucrose agar; remaining pale yellow, small, punctiform on glycerol and peptone agar. On malt agar, Ota reports cells larger, chains sometimes branched, cells up to Sfj. in diameter. On potato and potato with glycerol, growth good, creamy, white, smooth, shining, moist, 2-3 mm. thick, mammillate, the upper portion dry and efflorescent, below in contact with the glycerol, colony moist, diffluent with viscid or translucent streaks forming abundant mucous deposit. Finally, the colony is ochre yellow to sepia, with new pale yellow colonies dotting the surface of the old colony. After 6-8 months, the colony forms a thin dry pellicle, almost scaly, gray or reddish above and almost black below. On car- rot, a thin diffluent streak which never forms a thick colony. Ota reports Fig. 64. — Deharyomyces Hudeloi. 1-S, sprout mycelium ; !i, asci with ascospores. (After Ota 1923.) occasional chains, with branches several cells long. On gelatin, growth slow, colonies grayish white, mucous. No growth on coagulated serum. On broth, growth slow, with whitish flocculent or pulverulent deposit, no turbidity. On sugar broths, limpid, ring formed but no pellicle, deposit a white clot which disintegrates on shaking, producing turbidity which soon settles out. Ota reports no ring in 30 days on malt extract, sediment brownish white. Gelatin not liquefied ; sucrose inverted, glucose, maltose, and sucrose fermented, lactose not fermented. Debaryomyces mucosus A. & R. Sartory, Hufschmitt & Meyer, C. R. Acad. Sci. 191: 281-283, 1930. Case history and pathology in Hufschmitt, Sartory & Meyer, Ann. Derm. Syphiligr. VII, 2: 850-876, 11 figs., 1931. SACCHAROMYCETACEAE 311 Isolated from abscess on neck, not paint'nl, in some cases healing spon- taneously. No organism seen in the tissue, discharge showed a yeast and a streptococcus. On guinea pig caused abscess which healed spontaneously. Growth slow on solid media. In liquid media showing some budding cells, cells 3-4/A, filled with dense protoplasm and asci 4-5 (-12)/x in diameter with 4 spores, and at the bottom a zoogloea of round cells. Cells more elongate on repeated subculture. With beginning of ascospore formation in the culture, sprouting largely ceases. Copulation heterogamic, often between a sprout and an adult cell. Parthenogenesis occasional. Ascospores spherical, slightly rough, with one to several oil drops, 2-3/a in diameter. Colonies on sugar agar somcAvhat hilly, shining, viscid, with a brownish mucous deposit in water of condensation. Ascospores appear after the twelfth day, their number finally, after repeated transfer, being reduced to one. On usual media, colonies dirty white. Giant colony on potato agar attains a diameter of 5 cm. with elevated mounds at the center and a few radiating striae, margin crenulate. On potato juice, colony is humid and shining, later becoming ochraceous. Growth best on potato media, giving colonies of a cream color which later darkens to chamois yellow. Optimum temperature 37° C, growth slow at room temperature. In liquid media, there is a brownish yellow sediment, no pellicle, but a slight ring. Sucrose inverted, sucrose, and glucose fermented, no action on other sugars. No effect on gelatin or coagulated serum. Debaryomyces emphysematosus Ota, Derm. Woch. 78: 284-289, 312-316, 1924. Hefe emphysematosus Sasakawa, Centralbl. Bakt. I, 88: 273, 1922. Organism rtceived from Hautklinik, Bern, marked Sacc. emphysematosus. It is nonpathogenic for laboratory animals. On malt extract agar after 24 hours appear cells, usually solitary, very rarely in chains, 2-5/t in diameter, no fat globules. After 3 days on carrot, asci appear. Both heterogamous and isogamous asci smaller than in other species. Spores very slightly verrucose. Glucose, fructose, sucrose, and raffinose fermented. Debaryomyces Kloeckeri Guilliermond & Peju, C. R. Soc. Biol. 82: 1343, 1919; Bull. Soc. Myc. France 36: 164-172, 10 pis., 1920; Konokotina & Kras- silnikov, Jour. Microbiol. Leningrad 9: 93, 1929; Bachinsky, Jour. Microbiol, Leningrad 9: 14, 1929. Originally isolated from the throat of a patient with angina. Later iso- lated from fluid obtained in lumbar puncture by Bachinsky, 1929, in Leningrad. Cells ovoid, 2-3.5 x 3-7/^ in young malt extract cultures, with sediment and ring. Asci 1-spored. heterogamous, rarely isogamous, copulation. Spores spherical, verrucose, with oil droplet. Colony on malt agar, after 75 days at 15° C, gray yellow (Klinsieck Code 103A), moist, shining, center elevated and granular, with radial folds and wavy margin.' Growth on ethyl alcohol good, but organism cannot assimi- . late nitrates. Slight fermentation of glucose. 312 MEDICAL MYCOLOGY Debaryomyces Nadsoni Guilliermond & Peju, Bull. Soc. Myc. France 37: 35-38, 1921; Rev. Med. de I'Est [Nancy] 50: 342-347, 1922. Isolated from a case of sycosis, probably a secondary invader. Cells spherical or ovoid, often united in groups consisting of a mother cell and the cells sprouted from it, the older cells containing a single oil globule, 2-8 X 2-6/i,. Asci arising from heterogamic copulation between a large and small cell, often between mother and daughter. Usually one (rarely more) ascospore per ascus, 2-3/x in diameter, wall rough, containing one oil globule. At germination, the ascospores swell and lose the verrucosities of the wall. ]\Iinimum temperature for sporulation 8'^-9° C, optimum 20°-25° C, maximum 34°-35° C. Growth between 3° and 46° C, optimum 25°-30° C. On malt extract, whitish deposit visible after 36 hours ; a ring formed after 5 or 6 days but no pellicle. By Lindner's microfernientation method, no fermentation was observed, su- crose slightly inverted. Debaryomyces Matruchoti Grigorakis & Peju, C. R. Soc. Biol. 85: 459-462, 1921. Isolated from feces of patient suffering from helminthiasis. After 24 hours at 25° C, on malt extract, cells spherical or ovoid, 3.6-5.4 K 1.8-3fi. In old cultures, cells usually ovoid Avitli large fat globule. On Gorodkova agar, sporulation is common, usually heterogamous but occasion- ally isogamous. Asci are 1-spored. Spores verrucose with large oil globule. On malt agar, after one month at 25° C, colony attains size of a one-franc piece, white and shining, center slightly rosaceous. On Gorodkova agar, old colonies are chocolate brown. In malt extract liquid, a slight sediment ap- pears after 48 hours at 25° C, finally a ring, but no pellicle. On malt gelatin, colony slight yellowish white, center raised, margin lobed, rays present. The gelatin is not liquefied. Upper limit of growth is 37°-38° C, sporulation limit 30°-32°. Sucrose inverted, mannose alone fermented. Dekker (1931) claims there is no fermentation at all. Debaryomyces Fabrii Ota, Derm. Woch. 78: 286-289, 312-316, 1924. Hefe Fabry, Munchen. Med. Woch. 64: 1557, 1558, 1917. Debaryomyces Fabryi Dekker, Verhandel. K. Akad. Wetensch. Amsterdam, Afd. Natuurk. 28: 475, 1931. Isolated from interdigital mycosis in Dortmund. Nonpathogenic to mouse. On malt agar, after 24 hours at 25° C, cells spherical or ovoid, 2-6/a in diameter, occasionally 8-10/i,, in groups or pairs, very rarely in chains. At this time the oil drop is very small, but after 10 days the cell is larger and the oil drop large. On carrot, morphology is the same as on malt agar, but the oil drop is larger, and there is more ascus formation. On Gorodkova agar, no mycelium, heterogamy and sometimes isogamy, ascus ovoid or polyhedral with thick membrane 5-6ju. Spores ovoid or polyhedral, verrucose. Oil drop colors readily with Sudan III. SACCHAROMYCETACEAE 313 On malt extract a slight ring appears. Growth is good at 38° C, but there is none at 44° C. Fermentation of glucose, denied by Dekker (1931). Var. tremoniensis (Ota) Dodge, n. comb. Deharyomyces tremo7iiensis Ota, Derm. Woch. 78: 284-289, 312-316, 1924. Isolated from a case of interdigital mycosis in Dortmund, Germany. Morphology same as that of B. Fahrii. Ferments glucose and mannose. Debaryomyces Leopold! Ota, Ann. Parasitol. Hum. Comp. 1: 128-130, 1923. Saccharomyces neofonnans Leopold, Arch. Gynaekol. 61: 77-120, Pis. 1-6, 1900, nonal. Blastomyces {Leopold) Sasakawa, Centralbl. Bakt. I, 88: 273, 1922. Isolated from carcinoma of the ovary. Originally reported pathogenic to laboratory animals, but is nonpathogenic to guinea pig. Fig. 65. — Deharyomyces Leopoldi. (After Ota 1923.) After 24 hours on malt agar, at 25° C, cells are ovoid or spherical, the smaller having a diameter of 3-5/a, the larger 6 x 7/*. Cells solitary or in pairs with a tendency to occur in masses, uniguttulate, chains not observed (Fig. 65). After 9 days cells 6-8/a. In 40 days, membrane thickens, oil globules larger and more numerous. Growth on carrot similar. After 9 days on Gorodkova agar, spores appear, asci scarce, but more abundant than in D. Hudeloi, 1-spored. Spores 2-3;u,, usually echinulate, sometimes smooth. On plaster, asci appear in 4 days. Copulation usually heterogamous, but isogamj^ has been observed. Giant colonies, after 30 days, show white rays from center with definite margins. In malt extract, brownish white sediment appears in 30 days. No growth at 40° C, feeble growth at 36° C, growth still good at 12° C, and there is still some at 5° C. No liquefaction of malt gelatin in 35 days. Sucrose 314 MEDICAL MYCOLOGY inverted, slight fermentation of glucose and sucrose. Trace of fermentation of fructose, galactose, mamiose reported. None with maltose, lactose, and raffinose. Debaryomyces Lundsgaardi Ota, Ann. Parasitol. Hum. Comp. 1: 130-132, 1923. [Original case by Lundsgaard, Klin. Monatsh. Augenheilk. 38: 13-19, 1900.] Hefe Lundsgaard Sasakawa, Centralbl. Bakt. I, 88: 274, 1922. Isolated from hypopyokeratitis in 1900. Found nonpathogenic to guinea pig in 1923. After 24 hours at 25° C, cells spherical or ovoid, 2.5-3. 5/t, solitary or in pairs, rarely in colonies, occasionally showing oil globules. In 3 days, some cells are as large as 6-8/* with more abundant oil globules. In 9 days, an occasional chain may be seen, but never such as in D. Hudeloi. After 40 days, occasional cells as large as 12 x Idfi. On carrot, form and size of cells as on malt agar, but with more giant cells, thicker membranes, and larger fat globules. 8porulation is rare but heterogamous, the male cells being small and elongate. Asci spherical or ovoid, 3-5/x in diameter, with thick wall. Asco- spores 2-S/x, slightl}^ cchinulate, containing small oil globule. Giant colony attains diameter of 5 cm. in 35 days, is brilliant white with fine radial rays, margin definite with rare indentations. In malt extract, a brownish white sediment appears. Growth at 35° C, but not at 36° C, good growth at 12° C, and some as low as 5° C. No liquefaction of malt gelatin in 37 days. Organism inverts sucrose, but does not ferment any of the usual sugars. Debaj-yomyces Laedegaardi Ota, Ann. Parasitol. Hum. Comp. 1: 132-134, 1923. Isolated from skin infection (Sasakawa). Also in Krai collection, with- out record of pathogenicity. Nonpathogenic to guinea pig. On malt agar, after 24 hours at 25° C, cells appear ovoid, spherical or, occasionally, cylindric, solitary or in groups, 2.5-5/^ in diameter. After 3 days, cells a little larger. After 9 days, a few chains of long slender cells appear. Asci very rare. On carrot as on malt agar, with cells in old cultures up to 10/x in diameter. On Gorodkova agar, in 12 days appear asci, ovoid or poly- hedral with thick membrane, 3.5-4/i,, or, if oblong, 5-6/a broad. Ascospores 2-2.5)u,, veri'ucose, with small oil globule. Copulation heterogamous. Giant colony attains diameter of 3 cm. in 25 days, yellowish white with fine rays from center to periphery, edge definite. In malt extract, there is a brownish white deposit without pellicle or ring. No liquefaction of gelatin in one month. Sucrose inverted, glucose somewhat fermented, hence sucrose also. No fermentation with other sugars. Debaryomyces Hildegaardi Ota, Ann. Parasitol. Hum. Comp. 1: 134-136, 1923. Blastomyces Hildegaard Sasakawa, Centralbl. Bakt. I, 88: 273, 1922. SACCHAROMYCETACEAE 315 Deharyomyces Matruchoti race Hildegaardi Dekker, Verhandel. K. Akad. Wetensch. Amsterdam Afd. Natuurk. 28: 471, 1931. Organism came from Krai collection. Nothing is known of its origin. It is nonpathogenic to guinea pig. Grown on malt agar at 25° C, for 24 hours, it produces spherical, ovoid, and sometimes elongate cells, solitary or in chains, most often in pairs. In 8 days, cells may be as large as 7/a in diameter. In 40 days, membrane is thickened and large oil globules appear. On carrot, cells are large, up to lOju, m diameter. Heterogamy usual, occasionally isogamy. On Gorodkova agar, after 9 days, appear 1-spored asci. Spores are 2.5-3.5/x, uniguttulate, echinulate. In malt extract, a brownish white sediment appears. Organism grows at 36° C, but not at 37° C. Growth is fair at 12° C, but at 5° is almost absent. Malt gelatin is not liquefied in 35 days. Reported to invert sucrose, and slightly ferment glucose, fructose, maltose, and sucrose, but not galactose, lactose, mannose, or raffinose. Dekker (1931) denies any fermentation. Debaryomyces Gruetzii Ota, Derm. Woch. 78: 283-289, 312-316, 1924. Deharyomyces Matruchoti race Gruetzii Dekker, Verhandel. K. Akad. Wet. ensch. Amsterdam Afd. Natuurk. 28: 477, 1931. Isolated from mycosis of a nail of a six-year-old child in Kiel. Fatal to mouse in 26 days. Organism recovered. In 24-hour-old cultures on malt agar, cells are spherical or ovoid, solitary or in pairs, sometimes in groups, 2-4/a in diameter. On fourth day these have increased to 7-11/a. Month-old cultures contain many polyhedral cells. Asci appear in 10 days and are ovoid or polyhedral, 5-7/*. Asci appear earlier on carrot. Ascospores are spherical, verrucose, 2-3/a. On malt extract, yellow brown sediment appears, but no ring. The or- ganism was reported to ferment glucose, fructose, mannose, sucrose, but Dek- ker (1931), using quantitative methods, denied this. Debaryomyces Guilliermondii Dekker, Verhandel. K. Akad. Wetensch. Amsterdam Afd. Natuurk. II, 28: 1 : 457-459, 1931. Cryptococcus Fari^iae Pollacci & Turconi, nom. nud. Originally isolated from spoiled sausages by Cesari & Guilliermond. Re- ported by Agostini (1934) from blastomycosis of the Gilchrist type (case of Farina, 1929). Cells 3-8/x in diameter; asci 4-8/i,, 1-spored, copulation isogamous or heter- ogamous ; spores 2.5-4.5/1 in diameter, spherical, rough, with a large oil drop. On Pollacci agar, colonies white, round, margin subsinuous, plane, be- coming confluent, creamy white, minutely verrucose. On Sabouraud and malt agar, growth similar but sloAver, brownish yellow on maltose agar. On carrot and potato, colonies similar but Avhite and powdery. On malt gelatin, colony almost smooth. In liquid media with sugars, pellicle Avhite, thick, elastic, sediment yellowish white. On ethyl alcohol, pellicle thin, dull. No fermenta- tion of sugars, sucrose inverted. Gelatin liquefied. 316 MEDICAL MYCOLOGY Debaryomyces Bumieri Ota, Ann. Parasitol. Hum. Comp. 2: 35-37, 1924. This organism was isolated from epidermomycosis of eczematous aspect. Grown on malt agar, at 25° C. for one day, it shows spherical or ovoid cells, 4-5/x in diameter or 8-10 x d/x, occasionally up to 15/a long, solitary or in twos, occasionally in chains. There are large vacuoles and fat globules. The cell wall is very thin. After 7 days, cells solitary or in twos, membrane thicker. On carrot, cells may be solitary or in groups, but not in chains, walls thick, 1-12/* in diameter. On malt extract, cells often in chains. Gorodkova agar shows sporulation on the third day. Asci spherical, ovoid, or polyhedral, 4-7/i,. Ascospores spherical or ellipsoid, 2-4/a, verrucose, uniguttulate. On malt agar, colony chalky white, smooth, even edges. In malt extract a whitish deposit appears in 24 hours, and on the tenth day a ring which thickens and yellows on the fortieth day. Fermentation not reported. SACCHAROMYCOPSIS Saccharomycopsis Schionning, C. R. Trav. Lab. Carlsberg 6: 103, 1903. The type species is Cryptococcus guttulatus Robin. Cells ellipsoid or elongate, sprouting only from the poles ; on glucose broth, a thick powdery pellicle; asci parthenogenetic, 1-4-spored, ascospores Pig. 66. — Saccharomycopsis guttulatus. l-J/, stages in the formation of sprout cells ; 5-9, nuclear phenomena in the formation of ascus. (After Buscalioni 1896.) smooth and thick-walled, exospore rupturing on germination ; only glucose fermented, sucrose inverted ; gelatin not liquefied, milk clotted. Saccharomycopsis g-uttulatus (Robin) Schionning, C. R. Trav. Lab. Carls- berg 6: 124, 1903. Cryptococcus guttulatus Robin, Veg. qui Croiss. sur les Animaux Vivants, 52, 1847. Saccharomyces guttulatus Winter in Rabenhorst, Kryptog. Fl. Deutschl. ed. 2, I, 1: 72, 1881; Buscalioni, Malpighia 10: 281-327, PI. 7, 1896; lUd. 12: SACCHAROMYCETACEAE 317 59, 1898; Ann. Ig. Sperim. 8: 229-243, Pis. 2-3, 1898; Wilhemi, Diss. Bern [Cen- tralbl. Bakt. II, 4: 305-309, 353-361, 412-417, 713, 1898J. Atelosaccharomyces guttulaUis Beurmann & Gougerot, Bull. Mem. Soc. Med. Hop. Paris III, 28: 251, 1909. Remak, Diagnostisehe und Patliogenische Untersuchiingen, 221-225, 1845. Frequently isolated from feces, especially those of guinea pig (Remak 1845), also from digestive tracts of rabbits. Pathogenic to both guinea pig and rabbit in subcutaneous or intraperitoneal injections. Cells ellipsoid or oblong-ovoid, 6-8 x 15-20/t (Fig. 66, 1-4). Asci 1-4-spored (Fig. 66, 5-9). Exospore ruptured on germination of spore. Spores have not yet been produced on artificial media but are obtained by alternate wetting and drying of the feces. On glucose agar, colony is thin, moist, dirty white, regular margin with branching in old cultures. On glycerol a^ar, colony small, center elevated, margin branched and areolate. On potato, colony is slightly elevated, dirty white with elevated white points, margin also elevated, sinuous, slightly powdery. In acid glucose broth, a thick, powdery white pellicle forms. Su- crose inverted, glucose and sucrose fermented. On gelatin, colony circular, moist, whitish, margin irregular. Gelatin not liquefied. Casein precipitated in milk without acid formation. SACCHAROMYCES Saccharomyces Meyen, Arch. Naturgeschichte [Wiegemann] 4: 2: 100, 1837; emend. Reess, Unters, ii.d. Alkoholgarungspilze 80, 1870. The type species is Saccharomyces cerevisiae Meyen. Cells spherical, ovoid, or elongate, often in small chains, vegetative repro- duction by sprouting ; in malt extract a sediment, often a ring and very rarely a pellicle ; asci parthenogenetic, spores 1-4 per ascus, spherical, ovoid, rarely kidney-shaped, smooth-walled ; occasionally spores copulate before germina- tion ; glucose always fermented, usually also the other sugars ; nitrate not assimilated; poor growth or occasionally good growth on ethyl alcohol as a substrate but never producing a pellicle on this medium. Key to Species Colony rose to vermilion. *Si. granulatus. Colony grayish rose, grayish white on peptone gelatin. S. anginae. Colony yellowish, ochraceous, or brown. Growth only on potato, colonies ochraceous yellow with chocolate center. Growth on many media. ^- Ferrani. Fermentation with most sugars, colony brownish gray. S. Hagiwarae. Lactose not fermented, colonies white at first becoming chocolate brown in one month. Gelatin liquefied. S. Lemmonieri. Gelatin not liquefied. S. Zimmeri. Lactose and maltose not fermented, colony brownish. S. tumefaciens. No fermentation, colony yellowish becoming brown, cells often thick-walled, held, together by mucous sheath. S. Blanchardi. 318 MEDICAL MYCOLOGY Colony white or whitish. Gelatin liquefied. Gelatin not liquefied. Pellicle formed, sucrose inverted. Maltose fermented. Milk not coagulated. Milk coagulated. Maltose not fermented. Pellicle evanescent, maltose fermented, milk coagulated. S. Lemmonieri. 8. Sternoni. S. Zimmeri. S. labialis. S. Jadini. Only a ring formed, sucrose not inverted, maltose fermented. S. annulatus. Neither ring nor pellicle formed. s. pseudotubercolaris. Saccharomyces granulatus Vuillemin & Legrain, Arch, de Parasitol. 3: 237-268, 1900 [condensed by Pellier, Ann. Derm. Syphiligr. V, 1: 191-196, 1910] . Case of Pontoynont & Salvat, 1922, referred here belongs in Cryptococcus Fontoynonti Vuillemin. Fig-. 67. — Saccharomyces granulatus. 1, Z, sprout cells showing- sculptured membrane ; .?, elon- gate sprout cell; /,, 7, thlck-walled cells; 5, G, asci. (After Vuillemin & Legrain 1901.) Isolated from an abscess (?) of pigeon's egg size on the jaw of a patient with malaria. When lanced, the abscess exuded yeast cells and leucocytes. Organism cultivated. Pathogenic for rabbit. Cells ovoid or short ellipsoid, 4-5 x 3.3-4/a, solitary, budding. Atypical forms elongate (Fig. 67, 3), solitary or in simple branched chains with a rose pigment, which is insoluble in alcohol, acid, or alkali, collecting in the oil globules of the old cells. Membrane sculptured with granules occasionally arranged in lines or reticula (Fig. 67, 1, 2) . Vegetative cells sometimes encyst, forming chlamydospores (Fig. 67, 4, 7). Asci 2-4-spored (Fig. 67, 5, 6). Growth is good on all media tried — solid or liquid. In dark or light, pigment varies from rose to vermilion. No pellicle or cloudiness on liquids. Gelatin not liquefied. Saccliaromyces anginae Vuillemin, Rev. Gen. Sci. Pures Appl. 12: 735, 1901. Saccharomyces sp. Troisier & Achalme, Arch. Med. Exp. 5: 29-37, 1893. SACCHAROMYCETACEAE 319 Cryptococcus anginae Clerc & Sartory, C. R. Soc. Biol. 64: 136, 2 figs., 1908. Isolated in exudate from sore throat following typhoid fever. Lesions had general appearance of thrush. Cells in exudate 8-9/x in diameter, in groups of 6-8 among epithelial cells. On peptone gelatin, at 20° C, ascospores produced abundantly, usually 4 to an ascus, spherical or ovoid (Fig. 68). On malt extract gelatin, asci more elongate, the 4 ascospores being in a row (Fig. 68, 6-8). No chlamydospores on Nageli mineral medium. On agar, colony thick, rounded, grayish rose. On carrot (cooked), the same. Colonies, on peptone gelatin, both deep and superficial, the latter slightly grayish white and shining, the former brownish, spherical, surrounded by rays. On malt extract, no pellicle, turbidity clearing in 3 days, sediment thick, glutinous, brownish, having the odor of beer yeast. Sucrose fermented with a pleasant odor resulting. No acetic acid or aldehyde demonstrated. Saccharomyces Ferrani Froilano de Mello, Paes & Sousa, Arq. Hig. Pat. Exot. 6: 17-40, 1918. Fig. 68. — Saccharomyces anginae. (After Vuillemin & Legrain 1901.) Isolated from the pus of large cold abscess in the axilla with metastases. In this pus Avere yeast cells varying from 1-5/x in diameter. Cells spherical, with 1-2 refringent granules, solitaiy or in groups of 2-80 individuals, with pseudomycelium but no true mycelium. Asci numerous, 2-4-spored, ascospores spherical or subreniform. Grows on potato only, of the large number of media tried. Colonies ochraceous yellow, with center becoming chocolate colored and margins yel- lowish brown. Saccharomyces Hagiwarae Dodge, n. sp. Hefe Hagiwara, Jap. Zeitschr. Derm. Urol. 22: 941-980 [77-78], Pis. 19 A- 19B, 1922. Original lesion followed a bee sting behind the ear, later lesions on arms and axilla. Pathogenic to mouse, not to rabbit. Yeast cells spherical or ovoid, 2.8-6.4 x 1.9-5.6/x. On gypsum block after 53 hours, asci produced, 2-4-spored. No mycelium figured or described. 320 MEDICAL MYCOLOGY Colonies round, brownish gray, moist, shining, center slightly elevated, with radiations at the margin. Optimum temperature 17-20° but growth pos- sible between 0° and 55° C. Fermentation of maltose, dextrin, dextrose, fructose, galactose, lactose, sucrose, etc. The figures are poorly reproduced, but those showing spores are quite plausible. Saccharomyces tumefaciens Castellani & Chalmers, Man. Trop. Med. ed. 2, 768, 1913. Saccharomyces subcutaneus tumefaciens Curtis, Ann. Inst. Pasteur 10: 449- 468, 1896. Isolated from myxomatous tumor and lumbar abscess. Pathogenic for rat, mouse, rabbit, guinea pig, and dog. On acid peptone broth, cells are ovoid or spherical, 3-6/i, in diameter, wall thick, one to two refractive granules to a cell, no capsule. On sugar media, sells larger and in chains of 3-4 cells. Spores are spherical, 1-4 per ascus. Colony on potato, creamy, becoming brownish. On potato and glycerol, colony creamy, abundant. Growth from gelatin streak in creamy elevated spots. Feeble development in broth, with light floccose sediment. No growth on serum. No veil on liquid media. Glucose and sucrose fermented, not lactose or maltose. Gelatin not liquefied. Saccharomyces Blanchardi Guiart, ? Precis Parasitol. 1910; Castellani & Chalmers, Man. Trop. Med. ed. 2, 768, 1913. Saccharomyces sp. Blanchard, Swartz & Binot, Arch, de Parasitol. 7: 489- 507, 1903. Isolated from a gelatinous mass weighing a kilogram which was taken from the peritoneum in the vicinity of the appendix. Cultures from this mass gave principally this species. A rabbit, inoculated with this organism, died in about a week from diarrhea, but showed no peritonitis. Masses of the yeast were found in the kidneys. Also fatal to rats, mice, guinea pigs, and marmots, but not to hens or pigeons. Cells spherical, 4-10/* rarely up to 20/a in diameter, rarely in chains, often with several cells united in a common capsule whose gel is frequently as thick as the cells themselves. After a week on sugar agar, asci appear. These have thick membranes, are 8-spored, the spores 3/a in diameter. On sugar agar, at 22° C, growth slow, colony appearing after 5-6 days, attaining size of a lentil after 3 weeks, and at one month diameter of only 2 cm., opaque yellowish white or clear gray, surface granular with irregular contour, edges thick; with age, colony gradually becomes browner. On ordi- nary agar, growth is scanty and even slower, otherwise similar. Growth on potato slow but abundant, being several millimeters thick in a month, colony yellowish white, becoming verrucose and clear brown. On carrot, colony abundant and viscous. On potato glycerol, growth more rapid than on potato, colony gelatinous, clear yellowish white. On gelatin plates, colony whiter, more regular, round or festooned, and of a varnished appearance. Streaks SACCHAROMYCETACEAE 321 on gelatin grow more slowly and less abundantly than on agar, are grayish white and of a mucous consistency. Almost no growth on coagulated serum. After 4 days at 37° C, in bouillon, there is a sediment, but the liquid is clear. No fermentation with sugars. Gelatin slowly liquefied. Saccharomyces Lemmonieri Sartory & Lasseur, C. R. Soc, Biol. 78: 48-49, 1915; Bull. Sci. Pharmacol. 23: 151-157, 1916. Isolated from a case of bronchitis and pulmonary congestion. Pathogenic to guinea pigs, white mice, and rabbits. Cells 3.1-3.5/A in diameter, in age slightly elongate and clinging together in short chains. In deposits only spherical cells found. On plaster block and filter paper soaked in lactose solution, asci formed. These are 4-spored, the spores being 2.5-3/^ in diameter. On Sabouraud agar, colonies punctiform, nonconfluent, white, dull. Cre- nate margins 2-3 mm. high. At thirtieth day, colonies become chocolate brown. On potato, colonies white, punctiform, confluent, center elevated and acumi- nate, margins smooth. In age, colony cream white, becoming chalky on desic- cation. On carrot, thick creamy colonies cover whole surface in a week. On gelatin, growth is slow. On egg albumen, smooth punctiform colonies with smooth borders appear on the ninth day. In broth with peptone, glycerol, and glucose, growth rapid, pellicle quite thick, ring rather fragile, liquid tur- bid, clearing with a sediment 2 mm. thick in 3 weeks. Sucrose inverted, glu- cose, fructose, sucrose, and maltose fermented ; no action on lactose, galactose, inulin, starch, or mannite. Milk coagulated on tenth day, curd digested by the eighteenth. Liquefaction of gelatin begins on the eighth day. Saccharomyces Zimmeri Dodge, n. sp. Saccharomyces sp. Sartory, Sartory & Meyer in Zimmer, "Weil, Sartory, Sartory & Meyer, Arch. Derm. Syphilis 167: 211-221, 7 figs., 1932. Isolated from abscesses from the gluteal region and upper thigh of woman in Alsace. Pathogenic for guinea pigs. Cells ellipsoid, 4-5 x 2-3/i, forming chains up to 30/a long. Ascospores in pus and on gypsum block, cultures smooth, 1.25/* in diameter. Asci formed without copulation, and ascospores germinate directly. On potato agar and Sabouraud glucose agar, colonies white, homogeneous with smooth margin, center becoming verrucose with yellowish Brockelchen and radial furrows, finally grayish. On gelatin, colony spreading. On Meyer's liquid medium, gray flocculent sediment and a pellicle after 3 days at 37° C. ; this later sinks and a new one forms. Optimum temperature 27°-35° C, optimum pH 5.8. Glucose, maltose, and sucrose strongly fermented and acidi- fied, no action on fructose and lactose. Coagulated albumin unchanged. Milk coagulated but not digested in 8 days. Gelatin not liquefied. Saccharomyces Sternoni Sartory, Sartory, Sternon & Meyer, BuU. Acad. Med. Paris 107: 120, 121, 1932. Isolated from lesions in interdigital mycosis. Lesions reproduced on skin of guinea pig. 322 MEDICAL MYCOLOGY Cells spherical, 2-2.5/x in diameter, thin-walled, budding. Asci spherical to ovoid, 3.75-4/t in diameter, each containing 4 ascospores, 1.25-1.5/i. in di- ameter. No copulation observed. In the scales, the intracellular parasite shows thin-walled cells. On beef agar, potato agar, and Eaulin agar, growth is good but less abun- dant, colony dirty white. Growth poor on senim and egg albumen. On potato juice, growth is good, pellicle shining white, thick. On Raulin acid medium, a pellicle forms after 12 hours and falls to the bottom the next day. Growth not luxuriant. A pellicle forms in milk on second day. Invertase secreted, maltose fermented with evolution of gas. No action on lactose and levulose. Milk not coagulated. Neither gelatin nor coagulated serum nor egg albumen liquefied. Saccharomyces labialis Ribeyro, Anal. Fac. Med. Lima. 2: 122-127, 2 ph., 1918. Isolated from lesions on the lips of a young woman. Lesion on lower lip worst, involving region adjacent to mucous membrane. Abscess in guinea pig reproduced by inoculation. After growth for 24 hours at 37° C, in ordinary or glucose agar, cells were predominantly 4-6/* in diameter with some smaller ones, 3-4/i. After 12-15 days, ovoid asci, 7-8/a with 4 ascospores to each. Ascospores fairly thin- walled. No mycelial forms seen. On agar, growth abundant, colony dull white. Growth on potato good, colony elevated, opaque, white. On beet, growth good, colony dark and oily. Growth also good on carrot, colony oily and white. On SAveet potato, growth poor, colonies white, slowly yellowmg. On gelatin, growth is mediocre, colo- nies white, elevated, folded. Colonies on beef serum white and of the appear- ance of porcelain. These do not darken with age. On hay infusion agar, growth is scarce, colonies yellow white. In broth an abundant sediment forms. In broth, to which maltose has been added, growth is good, the liquid is turbid, and there is some sediment. In broth and glucose, levulose, sucrose, or man- nite, growth is abundant, pellicle and sediment formed. With glucose and levulose, acid is formed, with the other sugars slight. Medium is turbid. In broth plus dextrin, inulin, or lactose there is less growth, medium remains clear and sediment settles out, no acidity. In milk, growth is fair, sediment forms. In a 1% aqueous solution of nutrose, growth is fair, liquid clear with some sediment. Glucose and sucrose fermented, not the others. Milk not coagulated, gelatin not liquefied. Saccharomyces annulatus Negroni, Ann. Parasitol. Hum. Comp. 7: 303- 308, 1929. Isolated from an abscess of the epididymis in Roumania. Pathogenicity not known. Cells ovoid or ellipsoid, 3 x 4-6.5 x 11.5/x, on carrot (Fig. 69, 1). On both carrot and Gorodkova agar, cells develop asci without copulation. Asci 1-4- SACCIIAROMYCETACEAE 323 spored, most frequently 2. Spores spherical, smooth, tlattened at the poles, about 3.5/x in diameter with a slight equatorial ring suggestive of Williopsis Saturnus (Fig. 69, 3). Ascospores copulate in pairs within the ascus (Fig. 69, 3-7). The nuclei usually fuse and sprouting begins from the copulation canal. Parthenogenesis is occasionally observed. The sexuality is at about the same stage as in Sacchaiomycodes Luflwigii. Optimum temperature for budding about 40° C, optimum for sporulation 28°, although both sprouting and sporulation occur at room temperature (14-17° C). On malt agar, colony is dirty white, creamy in consistency, shining, moist. Giant colony attains diameter of 4 cm. in 1 month, grayish with some small, crateriform elevations in center. Malt extract becomes cloudy with a deposit after a few days and a ring at the surface. Glucose, maltose, lactose, galac- Fig-. 69. — Saccharomyces annulatus. 1, sprout ceU ; 2, ascospores ; S-t, stages in copulation. (After Negroni 1929.) tose, and raffinose fermented, sucrose and xylose slightly, without inversion in the case of sucrose. Fermentation of lactose denied by Dekker (1931). Gelatin not liquefied. Dekker (1931) denied the existence of the equatorial ring on the asco- spore, although she admitted she had observed one after staining with iron hematoxylin. This she claims to be an artefact and reduces this species to synonymy with the common beer yeast (V) ; eventually attacking the brain. C. psicrofilicus. Capsule well developed; cells spherical, mostly 3-4/^ (in pus up to 13/i) ; attacking the brain. C. histolyticus. Oryptococcus Neveuxii Vmllemin & Lasseur aput Servet, These de Nancy 108 pp., 1922; Lasseur & Servet, Rev. Med. de I'Est 50: 357-370, 1922. Torulopsis Neveuxii Almeida, Annaes Fac. Med. Sao Paulo 9: 10, 1933. Isolated from a case resembling tuberculosis, which yielded quickly to medication with KI; Grand Bassani, Ivory Coast. On potato, colony white, moist, then showing dull islets which spread, giving colony a chalky appearance, margins sometimes festooned. No asci on carrot. On glucose agar, colony as on potato except it is finally folded. On artichoke, colony thick, granular, green. In glucose peptone, pellicle thick, folded, white, climbing up the tube for 4-5 mm., sediment white and liquid clear. No fermentation. Acid production with glucose, fructose, maltose, galactose, lactose, and xylose. Cryptococcus neoformans (Sanfelice), Vuillemin, Rev. Gen. Sci. Pures Appl. 12: 747-750, 1901. Saccharomyces sp. Sanfelice, Centralbl. Bakt. 17: 113-118, 625-634, 1895. Saccharomyces neoformans Sanfelice, Ann. Ig. Sperim. 5: 239-262, Pis. 9-10, 1895. Torula neoformans Weis. Jour. Med. Res. 7: 280-311, Pis. 21-24, 1901. Blastomyces neoformans Arzt. Arch. Derm. Syphilis 145: 311, 1925. Torulopsis neoformans Almeida, Annaes Fac. Med. Sao Paulo 9: 10, 1933. Isolated from the surface and juice of a peach but found pathogenic for laboratory animals upon inoculation. [Weis gives additional information based upon subculture received from Sanfelice.] Lodder (1934) referred here a culture received from Voss as C. liominis. In young cultures cells spherical, 4-18ja, capsule very delicate with occa- sional vacuoles and numerous oil drops. In older cultures, cells spherical or SACCHAROMYCETACEAE IMPERFECTAE 331 ovoid, 2-22/i,, with irregular and distorted forms. Large vacuoles present, capsules more conspicuous. Cell division in all planes; no ascospores. On malt agar, colon^^ whitish, translucent, elevated, becoming opaque, yellowish white, margins beveled, finally yellowish brown and up to 6 mm. thick. On malt gelatin, colonies confluent, flat margins elevated, centers slightly depressed, surface dull, smooth, waxy, yellowish white. On blood serum, colonies milk white, elevated, slightly slimy, glistening becoming yel- lowish, drying chalky, resembling daubs of white paint. On potato, colony elevated, margins regular, verrucose, white or slightly yellowish becoming gray or drab. On malt extract, liquid clear, sediment abundant, granular to flocculent, ring well developed, pellicle very finely granular or cheesy, be- coming thick. On glucose broth, little or no pellicle, sediment grayish white. No action on milk. (lelatin not liquefied (Sanfelice) or slowly liquefied after a week, becoming complete in 5-6 weeks (Weis), Cryptococcus Plimmeri (Costantin) Neveu-Lemaire, Parasitol. Anim. Domest. 60, 1912. Saccharomyces sp. Plimmer, Centralbl. Bakt. 25: 805-809, 1899. Saccharomyces Plimmeri Costantin, Bull. Soc. Myc. France 17: 147, 1901. Torula Plimmer Weis, Jour. Med. Res. 7: 280-311, Pis. 21-24, 1902. Torulopsis Plimmeri Almeida, Annaes Fac. Med. Sao Paulo 9: 10, 1933. Found in about half of a series of cancer cases examined by Plimmer. Pathogenic to guinea pig in both intraperitoneal and subcrural injections. In pellicle of young cultures, cells 4-8/i,, without capsules, vacuolate with many oil globules. In sediment, cells larger, 2-10/a, capsules on very large cells, not on small ones. In old cultures, cells 2-16/* with 1-3 large vacuoles, capsules highly developed; cells in chains of 3 cells, the middle cell usually smaller; cell division in one plane only; no ascospores. On malt agar, colony whitish, translucent, elevated, becoming radially furrowed and zonate, center pure yellow Math white margin. On malt gelatin, colonies confluent, smooth, center not depressed, margins not elevated, white, then yellowish. On blood serum, growth slight, whitish, flat, giving the ap- pearance of daubs of paint. On potato, colony yellowish brown, surface con- voluted, margin regular. In malt extract broth, liquid clear, pellicle delicate not settling to the bottom, sediment cheesy to coarsely granular, highly de- veloped, ring heavy, opaque. In glucose broth, pellicle well developed at 25°, little or none at 37° C; sediment very slight. Growth good in litmus milk but no change in medium. No fermentation of sugars. Gelatin not liquefied (Plimmer), very slowly liquefied after one week, becoming complete in 5-6 weeks (Weis). Cryptococcus lithogenes (Sanfelice) Vuillemin, Rev. Gen. Sci. Pures Appl. 12: 739-741, 1901. Saccharomyces litogenes Sanfelice, Centralbl. Bakt. I, 18: 521-526, 1895. Toriila Sanfelice Weis, Jour. Med. Res. 7: 280-311, Pis. 21-24, 1902. Blastomyces lithogenes Sasakawa, Centralbl. Bakt. I, 88: 274, 1922. 332 MEDICAL MYCOLOGY Torulopsis Uthogenes Almeida, Annaes Fac. Med. Sao Paulo 9: 10, 1933. Isolated from the lymphatic ganglia of an ox, which died of generalized carcinoma. Pathogenic for guinea pig and mouse, but not for sheep. (Strain studied by Weis was isolated from adenocarcinoma of human ovary by San- felice.) In tissues cells spherical, variable in size, containing rounded or angular masses of calcareous material. In pellicle, cells spherical, 1-8/x,, oblong and misshapen cells observed during active sprouting ; capsules absent or ex- tremely delicate. In sediment, cells 3-12/a with a capsule, protoplasm granular with oil drops and vacuoles. In old cultures capsules 1-1. 5yu, thick, cells often embedded in a mass of gel, many resting cells present ; cells dividing in all planes, no ascospores present. On malt agar, colony at first whitish, elevated, becoming opaque, yellow, center folded, margins even, sharply beveled. On malt gelatin, colonies con- fluent, flat, somewhat elevated at the margin Avith slightly depressed centers, surface dull, smooth, waxy, yellowish white. On blood serum, colonies white, elevated, slimy, glistening, yellow, growth scant, drying chalky, suggesting daubs of white paint. On potato, colony very dark brown, margin very ir- regular, surface glistening and corrugated. In malt extract broth, liquid clear, pellicle at first very delicate, later heavy and membranous; sediment slight until after fall of first pellicle, then heavy, cheesy, and flaky; ring well developed. On glucose broth, pellicle developed, white becoming yellowish, little sediment grayish white. No fermentation, no action on milk, gelatin not liquefied (Sanfelice) or slowly liquefied after one week, the action com- plete in 5-6 weeks (Weis). Cryptococcus nasalis (Harrison) Dodge, n. comb. Torula nasalis Harrison, Trans. R. Soc. Canada, Biol. Sci. 22: 207, 1928. Torulopsis neoformans race nasalis Lodder Anaskosporogenen Hef en 1 : 176, 1934. Isolated from nasal tumor in horse by K. F. Meyer (1912). In young colonies, cells spherical, average diameter 4/a. In 57-day-old cultures, cells still spherical, varying from 3.5 to 7/a in diameter (average 5/t), oil globules present. Growth good between 25° and 37° C. On malt agar, growth is elevated, shining, spreading, and white, later darker in color and more massive. On potato, colony white, raised, dull, spreading, becoming darker in age. On malt gelatin, colony elevated, white, waxy, radiately striate, forming a funnel-shaped depression in 57 days. Giant colony is dull, white with slightly elevated and lobate margin and depressed center. Slight growth under olive oil. In malt extract, a creeping pel- licle appears, then a heavy ring with cloudiness and thick sediment. No fermentation of sugars. Acid produced in glucose, fructose, mannose, man- nite, xylose, maltose, sucrose, and dextrin; forming a pellicle and turbidity which clears as the sediment forms in all sugars except rhamnose, glycerol, SACCHAKOMYCETACEAE IMPERPECTAE 333 and inulin. Sucrose inverted. Milk becomes alkaline without other change. Malt gelatin slightly liquefied in 57 days, ordinary gelatin in 26 days. Cryptococcus meningitidis Dodge, n. sp. Torula histolytica Harrison, Trans. R. Soc. Canada, Biol. Sci. 22: 187-225, 5 ph., 1928. Isolated from a case of cystic blastomycosis of the cerebral gray matter. Cells spherical, even in size, averaging 3.5/4 in diameter; on 56-day-old cultures 3-6/x in diameter, averaging 4.2ju,; oil globules present, the older cells having a mucinous capsule. On malt extract agar, growth is Avhite and shining, slightly spreading, in age becoming yellow then brown, the growth more massive. In malt gelatin, colony white, waxy, elevated, with radiate markings, later forming a funnel- shaped depression. Giant colony is whitish, shining, slightly rugose, margin lobate, center slightly depressed. On potato, colony elevated, white, shining, and spreading. In malt extract, a ring and pellicle develop with turbidity and heavy sediment. Ring in solutions of maltose, lactose, sucrose, raffinose, salicin, and dextrin, none in glycerol, sorbite, or inulin. Under olive oil, capitate colonies form, growth slight. Grows well between 25° and 37° C. Sucrose not inverted. In acid sugar media (pH 5.2), pellicle, turbidity and sediment in glucose, mannose, galactose, fructose, and xylose. In milk, ring forms, reaction slightly alkaline, no coagulation. Gelatin liquefied in 26 days. Lynch & Rose (1926) report slight liquefaction of gelatin after 3 months, also slight acidity in maltose (and after 3 months in sucrose). Rappaport & Kaplan (1926) report their organism to be gram -positive when young. Growth best on carbohydrate media and under low oxygen tension. Cryptococcus rotundatus (Redaelli) Dodge, n. comb. Torulopsis rotundata Redaelli, I Miceti come Associazione Microbica nella Tubercolosi Polmonare Cavitaria 46, 1925 (fide Lodder, Anakosporogenen Hefen 1: 170-172, 1934). Isolated from a case simulating tuberculosis of the lung. Cells spherical 5-6.5/a in diameter, giant cells to 10/t. On malt agar, cells slightly ovoid. On carrot agar, colonies white to slightly pinkish, surface dull. On malt agar, colonies yellowish, slightly reddish, humid, smooth, slimy, margin smooth. On malt gelatin, colonies yellowish, dull, center elevated, margin somewhat sinuous and slightly shining. On malt extract, a ring and sediment, the whole fluid becoming slimy. Cryptococcus Gotoi Dodge, n. sp. Hefe Goto, Mitt. Med. Fak. Univ. Tokyo 15: 75-101, Pis. 9-10, 1915. Isolated from a case of blastomycotic meningitis in Japan. Fatal for mice, rats, guinea pigs, and rabbits, but not for dogs. Cells 3-6/x, walls thin when young, becoming very thick-walled when old. No mycelium or ascospores. Capsule well developed. 334 MEDICAL MYCOLOGY Colonies on blood agar very small, white, punctiform, moist, by transfer gradually accustomed to all the usual media. On malt (mizuame) agar, colo- nies moist, white, round, opaque, finally confluent and yellow, especially at room temperature. GroAvth poorer on ordinary agar, blood, glycerol, and ascites agar. On potato, colony moist, white, confluent. On broth and pep- tone solution, no pellicle, sliglitly granular sediment. On malt extract and sugar solutions, a ring develops, also turbidity and a yellowish white sediment. No fermentation ; acid in arabinose, xylose, glucose, fructose, galactose, man- nose, sucrose, lactose, maltose, raffinose, inulin, starch, dextrin, mannite, and erythrite. Milk coagulated in 10 days. Gelatin not liquefied in several weeks. Cryptococcus hondurianus (Castellani) Dodge, n. comb. Torulopsis hominis var. Jwnduriana Castellani, Med. Press Circular 136: 438-443, 1933; Jour. Trop. Med. Hyg. 36: 311, Fig. 36, 1933. Torulopsis neoformans Lodder, Anaskosporogenen Hefen 1: 176-178, 1934. Isolated from blastomycosis of the skin with pulmonary complications in Honduras. Cells spherical, 3.5-10. 5ja, occasionally ovoid. In malt extract, cells 3.9-5 X 4.5-5.5/i,. On glucose agar, colony smooth, mucoid, yellowish. No pigmentation of mannitol agar. On malt agar, colony yellowish, humid, smooth, slimy, margin smooth. On malt gelatin, colony dull, almost smooth, margin smooth. On malt extract, broad yellowish ring, well-developed sediment, and a few slimy floating islets. No fermentation and no acidity with carbohydrates except glucose, sucrose, xylose (and in one strain on glycerol). Serum not liquefied; gelatin liquefied very slowly after 2 weeks in one strain, in other not liquefied. Cryptococcus cong-lobatus (Redaelli) Pollacci & Nannizzi, Miceti Pat. deiruomo e degli Anim. 7: No. 63, 1928. Torulopsis conglohata Redaelli, I Miceti come Associazione Microbica nella Tubercolosi Polmonare Cavitaria, 1925. Isolated from lungs with much destruction of tissue, in Italy. Later iso- lated by Carnevale-Ricci from tonsillar crypts. Subsequently found in several other cases in lesions of the lung. Cells spherical or slightly ovoid, 1.5-3/x, occasionally up to 4.5/^, in diam- eter, at first homogeneous then vacuolate and uniguttulate, solitary or in groups of 3-6 cells ; on carrot, cells somewhat larger, 5-6ju, ; thick-walled chlamy- dospores, 5.5-6.5/*, present in old cultures but no mycelium. Chlamydospore wall rupturing on germination. No ascospores. On Pollacci agar, colony thick, luxuriant, spreading, white, smooth, shin- ing, margin smooth. Colonies on Sabouraud agar similar. On carrot agar, colony white, creamy, opaque, margin smooth or slightly striate. On Gorod- kova agar, colony brown, opaque, easily separable from the substrate. On carrot, colony creamy. Avhite, margins irregularly lobed. On potato, colony grayish, surface slightly irregular, thick and opaque, small. On malt gelatin, colony plane, yellowish, opaque, 3 cm. in diameter in 30 days, margin finely SACCHAROMYCETACEAE IMPERFECTAE 335 and irregularly dentellated. On malt extract, producing fragments of a milky white ring with traces of a pellicle floating on the liquid, with dirty yellow sediment. Milk coagulated. Cryptococcus Kleini (Weis) Colin apud Gueguen, Champ. Paras. Homme 114-115, 1904. Torula sp. Klein, Jour. Ilyg. 1: 90-94, 1901. Torula Klein Weis, Jour. Med. Res. 7: 280-311, Pis. 21-24, 1901. Isolated from milk along with other organisms. Pathogenic for guinea pig, rabbit, and dog, producing tumor in guinea pig in original case. Cells spherical, 2-8/* in diameter, homogeneous in content, finely granular, thin-walled when young, finally surrounded with a hyaline capsule occupying about one-fourth the total diameter. On solid media cells secrete a mucinous sheath. Growth from 20° to 37° C. On glycerol, malt, and sugar agars, colonies thick, viscid, yellowish. Colo- nies on nutrient agar remain white. On alkaline nutrient gelatin, colony thick, rounded, moist, center raised, white in reflected light, brown and granu- lar in transmitted light. On sugar gelatin, growth more rapid, colony light yelloAv. After 5-6 weeks, gelatin slowly liquefied into a thick, turbid, syrupy mass. On blood serum, colony white gradually becoming yellowish, slimy, growth very slow. On potato, colony yellowish, surface convoluted, margins ir- regular. On malt extract, pellicle delicate, sediment flocculent to granular, ring well developed. In glucose broth, a slight pellicle and a powdery white sedi- ment appear, the liquid remains clear. Growth on milk good, medium not coagulated. No fermentation of lactose, sucrose, maltose, or glucose. Cryptococcus Guilliermondi Beauverie & Lesieur [name with reference to previous description]. Jour. Physiol. Path. Gen. 14: 998, PI. 11, 1912. Levure Guilliermond & Lesieur, C. R. Soc. Biol. 70 : 952-954, 1911. Isolated from the lungs of a man dying with cancer of the kidney. Lung symptoms resembled tuberculosis without accompanying fever. Inoculations into guinea pig negative. Cells spherical or ovoid, 3-5 x 2.8-4.3/x, solitary or in pairs, with a thick membrane but no capsule, possessing one or two vacuoles and numerous oil droplets. In fruit juices, cells 3.5-6.4/j, in diameter, even larger in Raulin solu- tion. In old cultures elongate cells may be seen. Sometimes, though rarely, these are branched. Giant cells common. No spores found in cultures on plaster blocks, Gorodkova agar, or slices of caiTot. On carrot, growth abundant, white even in old cultures, viscid. On potato, growth feeble, colonies size of pinhead, white, dry. On agars, grape juice, broth, peptone solution, growth is good, with viscid, yellow colonies ; grayish in age. In other fruit juices, growth poor, sediment produced but no pellicle. Sugars are inverted, but there is no alcoholic fennentation. Crjrptococcus psicrofilicus Nino in Speroni, Llambias, Parodi & Nino, Bol. Inst. Clin. Quiriirg. Univ. Buenos Aires 5: 94-155, 35 figs., 1930. Isolated from a nodulo-ulcero-scabrous dermatitis originally, with subse- quent involvement of the prostate, gastritis, and meningoencephalitis, this 336 MEDICAL MYCOLOGY latter the immediate cause of death about 3 months after the appearance of the first cutaneous lesion. Patient a native of Spain, resident twenty years in the Argentine. Pathogenic to rabbits, guinea pigs, white rats, and mice, with reproduction of lesions both microscopically and macroscopically. Examination of pus from abscesses showed spherical yeast cells, very variable in size, 7-30/a, sprouting or not, possessing a thick, smooth double- layered wall, quite visible and enclosing a somewhat granular substance. Some cells have 2-4 granules, 1-2/^ in diameter. Some large cells with thick, gelatinous, hyaline wall, 5/x or more thick. Cells solitary or in chains of 2, 3 or 4. Some- times vacuolate with protoplasm along wall in ring or half moon and showing a few black corpuscles. Occasionally spore wall ruptures, allowing cell con- tents to escape. Yeast cells were also encountered in the tissues. Under culture the or- ganism always has appeared with spherical cells, 3-7/* in diameter, single or sprouting, with thick external membrane and somewhat granular content. Hyphae never seen. Gelatinous sheath never seen in culture. While in the pus, the organism was hard to stain, being gram-negative and only faintly tinted with Leishman and May-Griinwald-Giemsa stains ; the cultivated organ- isms stain readily. Optimum temperature 25°, little growth at 37° C. On Sabouraud agar, abundant growth in 24 hours, colonies cream white, hemispheric, not confluent, humid above, not adhering to medium below. When growth is very abundant, culture appears dry and oily. Growth in plain agar similar, but less abundant. On poor nutritive agar, abundant growth in 24 hours. On potato, with 8% glycerol, good growth in 48 hours, colonies confluent, moist, grayish white, liquid without turbidity, with sedi- ment at bottom of tube. On carrot with 8% glycerol, regular growth after 48 hours, colonies isolated, white, hemispheric, moist, sediment at bottom of tube with liquid clear. On coagulated human serum, growth scarce after 24 hours and poor thereafter, no color production or liquefaction. When inocu- lated by stab, growth scant at first. After 20 days infundibuliform growth and liquefaction at bottom. Scant growth also on coagulated albumen, with neither liquefaction nor pigment formation. On Gougerot gelatin (stab), sur- face growth at 48 hours, colonies flat, white with scalloped edge and moist surface. On Drigalski-Conradi medium, no growth in 76 hours. In plain broth or peptone solution or acid Raulin solution, after 48 hours poor, pow- dery growth along the walls of the tube and forming a scant sediment at the bottom, no turbidity and no pellicle. In Sabouraud broth, after 3 days, medium becomes turbid with abundant sediment and white pellicle which falls to the bottom after being shaken. Does not ferment any of the usual sugars, does not liquefy gelatin or coagulate milk. It seems probable that the following organism should be referred here. Torula sp. McGehee & Michelson, Surgery, Gyn. Obstet. 42: 803-808, 6 figs., 1926. SACCHABOMYCETACEAE IMPERFECTAE 337 Isolated from pus of an inguinal abscess with symptoms of gonococcemia in a negro woman, but all attempts to isolate Neisseria go7iorrhoeae were unsuc- cessful. Highly pathogenic for guinea pigs and rabbits. Yeast cells in pus 7-15/* in diameter, spherical or ovoid, walls thick, en- veloped in a capsule. Gram-negative or nearly so, not acid-fast. Each cell with 1-5 spherical refractile granules. In cultures, morphology similar to that in pus, the gelified matrix appearing only in old cultures and holding the individual cells together in short chains. No ascospores. On Sabouraud agar, colonies in 24 hours small, white, opalescent, round, becoming coarsely granular and hemispheric, then thickly mucoid and heaped up in the center, finally confluent and of varying shades of yellow and brown. On blood agar, colonies sparse, discrete, small, grayish white. On Loffler's blood serum, colonies similar but smaller and more watery. In broth, growth poor, forming a thick adherent sediment with a tendency to grow up the sides of the tube. No fermentation of sugars, no liquefaction of gelatin, no action on milk, no indol formation. Crjrptococcus histolyticus Stoddard & Cutler, apud Castellani Amer. Jour. Trop. Med. 8: 393, 1928. Torula sp. Frothingham, Jour. Med. Res. 8: 31-42, Pis. 3-4, 1902; Tiirck, Deutsch. Arch. Klin. Med. 90 : 335-366, 1907. Torula histolytica Stoddard & Cutler, Monog. Rockefeller Inst. Med. Res. 6: 1-98, 9 pis., 1916; Ota, Ann. Parasitol. Hum. Comp. 2: 41-43, 1924. Torulopsis histolytica Castellani & Jacono, Jour. Trop. Med. Hyg. 33: 316, Fig. 44, 1933. Isolated from the lung of a horse by Frothingham (1902). Experimental inoculation by Stoddard & Cutler (1916) produced brain lesions similar to those in human brains caused by C. meningitidis. Perhaps organism of Voss (1923) should be referred here. Cells spherical, 1-6/* in diameter (average 3-4/*), sprouting from the medium-sized cells. Capsule 0.5-1.0/i thick. No spores observed. On glucose agar, colonies at first white, then more or less yellowish, heaped up, smooth, pasty, shining, thick. In broth, slight turbidity, no pellicle, fine white sediment. No fermentation of lactose, sucrose, dextrin, or glucose. No liquefaction of gelatin. Doubtful Species The following species of pathogenic yeasts have been too poorly described to place definitely. Cryptococcus Breweri (Verdun) Castellani & Chalmers, Man. Trop. Med. ed. 2, 771, 1913. Blastomyces sp. Brewer & Wood, Ann. Surgei-y 48: 889-896, Pis. 1-7, 1908. Atelosaccharomyces Breweri Verdun, Precis Parasitol. 1912; Froilano de Mello, Paes & Sousa, Arq. Hig. Pat. Exot. 6: 34, 1918. 338 MEDICAL MYCOLOGY Saccharomyces Breweri Neveu-Lemaire, Precis Parasitol. 97, 1921. Torulopsis Breweri Almeida, Annaes. Fae. Med. Sao Paulo 9: 10, 1933. Cryptococcus Copelli Neveu-Lemaire, 1921, referred here as synonym by Vuillemin, Champ. Paras. 96, 1931. Isolated from two tumors on spine involving spinal processes, on Russian who had been in America for six months. Tumors removed and recovery complete. Inoculation into guinea pigs reproduced the human lesions. Reproduction by sprouting only, with some chains. Growth slow, small grayish punctate colonies after 48 hours on glycerol agar. On agar slants, heaped up, yellowish, creamy growths formed. On potato, growth heavy and white. Organism grows on litmus milk but causes no change. Gelatin not liquefied, perhaps because of slow growth. Cryptococcus Costantini Froilano de Mello & Gonzaga Fernandes, Arq. Hig. Pat. Exot. 6: 294, 297, 1918. Cryptococcus Jiominis var. Costantini Vuillemin. Saccharomyces hominis Costantin, Bull. Soc. Myc. France 17: 145-148, 1901. Torulopsis Costantini Almeida, Annaes Fac. Med. Sao Paulo 9: 10, 1933. Organism isolated from a tumor. Does not become brown on potato. Membranes not thickening on ordi- nary media. Cryptococcus Cooperi Dodge n. nom. Cryptococcus quasi linguae pilosae Cooi)er, Am. Jour. Trop. Meil. 12: 97- 100, 1932. Isolated from a single case of lingua nigra out of 102 studied. Patho- genicity not reported. Cells ovoid, 2-9/x, long, producing only one sprout at a time. Cultural characters not given. Acid produced with glucose, maltose, fructose, and galactose, also litmus milk. No fermentation. Pellicles pro- duced in liquid media. Since the name as published hy Cooper is a polynomial, it is invalid. To shorten it to linguae-pilosae is impossible as that name already exists for an- other species. To hyphenate all three words makes an unnecessarily long name. Hence, I take pleasure in dedicating it to its author. Cryptococcus Corsellii Neveu-Lemaire, Precis Parasitol. Anim. Domest. 60, 1912. Blastomicete Corselli & Frisco, Ann. Ig. Sperim. 5: 433-447, Pis. 12-15, 1895. Isolated from a sarcoma. Colonies small, more or less circular on gelatin, glycerol agar, or sugar agar. No growth on potato or fruit. Cells spherical, sprouting. Ascospores reported, but figures not very convincing. Description too inadequate for further identification. Notes on use of various stains in organism and in tissue. SACCHAKOMYCETACEAE IMPERPECTAE 339 Cryptococcus degenerans Viiillemin, Rev. Gen. Sci. Pures Appl. 12: 740, 1901. Blastomyces vitro simile degenerans Roneali, Centralbl. Bakt. 18: 353-368, 1895. Zymonema degenerans Froilano de Mello, Paes & Sousa, Arq. Hig. Pat. Exot. 6: 33, 1918. Tortdopsis degenerans Almeida, Annaes Fac. Med. Sao Paulo 9: 10, 1933. Pathogenic to guinea pig. Cells spherical, isolated or united in groups in the tissue, thick-walled. On potato, colonies are small, dry, elevated, grayish white, margins in- dented and wavy, cells double the size of those on other media. On gelatin, superficial colonies are yellowish gray and granular; those deeper are small, rounded, brownish. In glycerol or glucose broth, a powdery deposit appears. No fermentation of sucrose. Gelatin not liquefied. Cryptococcus gracilioides Castellani, Jour. Trop. Med. Hyg. 28: 220-223, 1925; Am. Jour. Trop. Med. 8: 394, 1928; Mallardo, Jour. Trop. Med. Hyg. 32: 145-147, 1929. Monilia sp. Castellani, Jour. Trop. Med. Hyg. 28: 8, 9, 1925. Cells ovoid to flask-shaped, 3-4.5 x 2-3ju,; gram-positive, not acid-fast. On dextrose and on McConkey lactose agar, colonies small and similar to those of Streptococcus. Very scanty growth and no liquefaction of serum or gelatin. No fermentation. Darkening of lead agar. Cryptococcus granulomatog-enes Vuillemin, Rev. Gen. Sci. Pures Appl. 12: 741, 1901. Saccharomyces granulomatogenes Sanfelice, Zeitschr. Hyg. 29: 498-501, 1898 (spelled granulomatosus in heading). Torulopsis granulomatogenes Almeida, Annaes Fac. Med. Sao Paulo 9: 10, 1933. Isolated from granuloma of lung of swine. Not pathogenic to laboratory animals but reproduced the disease when inoculated into swine. Cells smaller than in Cryptococcus 7ieoformans. No ascospores. Colonies round, white on usual media, gray on potato, rose color on pear and honey. Thin pellicle and turbidity in glucose broth. Gelatin not liquefied. Cryptococcus Jeanselmei Burnier & Langeron, Congres Derm. Syphiligi*. Langue Franc. 1 : 40, 41, 1922. Torulopsis Jeatiselmei Almeida, Annaes Fac. Med. Sao Paulo 9: 10, 1933. Isolated from a case of interdigital pruritus with exfoliation. Cells about 4/a in diameter, in age attaining 6fi. No mycelium or asco- spores observed. Colonies on maltose agar whitish, shining, smooth or mammillate at the center. Cryptococcus mutilans Breuil, 1915 ; Neveu-Lemaire, Precis Parasitol. Hum. 79, 1921. 340 MEDICAL MYCOLOGY Reported as the cause of gangosa or rhinopharyngitis mutilans in the Pacific Islands. Not cultivated and inoculations negative. Cryptococcus myrmeciae Chalmers & Christopherson, Jour. Trop. Med. Hyg. 17: 129-135, 1914. Torulopsis myrmeciae Almeida, Annaes Fac. Med. Sao Paulo 9: 10, 1933. Isolated from extensive warts on the face. In lesions, cells yeastlike, sprouting, 1.4 x 2.1/a in diameter. Not obtained in culture. Cryptococcus psoriasis Rivolta, Paras. Veg. 464, 469, 1873 (fide Kraus). Torulopsis psoriasis Almeida, Annaes Fac. Med. Sao Paulo 9: 10, 1933. Seen in scales from psoriasis patients. Cells spherical, thick-walled, sprouting, not producing mycelium. Cryptococcus radiatus Sartory, Sartory & Meyer, C. R. Soc. Biol. 106. 597, 598, 1931. Saprophyte isolated during an epidemic in Alsace, the disease being char- acterized by falling hair. Cryptococcus septicus (Gaetano) Dodge, n. comb. Saccharomijces septicus Gaetano, Rif. Med. 13: 3:590-593, 1897. Isolated from septicemia in guinea pig. Cells mostly spherical, occasionally ovoid. No hyphae observed. Ger- mination by sprouting. Optimum temperature for growth 20-30° C. On gelatin, small, round, shining, milky white colonies. About the same on Casagrandi agar. Slower growth in neutral or alkaline media. In liquid cultures, no pellicle or turbidity, although sediment appears at the bottom of the tube. Cryptococcus Burnieri Nannizzi, Tratt. Micopat. Umana 4: 305, 1934. Cryptococcus de Burnier (cas 8) Ota, Ann. Parasitol. Hum. Comp. 2: 47, 48, Fig. 7, 1924. Isolated from case of epidermomycosis. On malt extract, at 25° C, after one day, cells ovoid or slightly elongate, rarely spherical, slightly pointed at the ends, 4-5 x 2//,, solitary or in pairs, walls thin, slightly larger after a month, still solitary but becoming thick- walled, and on carrot reaching 7/a in diameter. On Gorodkova agar, cells elongate, 13 x 3/x, in small chains of 3-4 cells, no asci; sprouting bipolar. On malt agar, colony cream color or grayish, surface smooth, shining, moist, margins definite. On malt extract, a slight ring and thick sediment. Cryptococcus sp. Refe, Ota, Festschr. Keizo Dohi 335-350, Tls. 32, 33, 1917. Isolated from two cases of cutaneous abscesses. Pathogenic to guinea pig, rabbit, and white rat. Cells 1.4-4. 5/x in diameter when spherical, or 4.9-6.2 x 3.1-2.9/^ when oblong, thick-walled, granular. SACCHAROMYCETACEAE IM PERFECT AE 341 On agar, colony thin, grayish white, slimy. On potato, a grayish white, thick membrane appears. Membrane consistency of "mizuame, " with fine hyphae. Peptone solution, with or without sugar, remained clear. No acid or gas formed in presence of sugars, but alcohol was detected by iodoform test. Perhaps this was the strain later sent to the Centraalbureau voor Schim- melcultures under the name Cryptococcus hominis and referred by Lodder (1934) as C. neoformans. Cryptococcus sp. Blastomyces sp. Escomel, Bull. Soc. Path. Exot. 8: 90-92, 1915. Isolated along with Leishmania in the disease called "espundia" in Peru. Started as cutaneous ulcers on neck and limbs and persisted for a long time. Secondary lesions, nodular ulcers suggesting a mulberry, develop in mucus of nose, pharynx, tonsils, pillars of fauces, soft palate, tongue, cheeks, gums, larynx, and sometimes even extend to the external surface of the head. The ulcers gradually extend, accompanied by great salivation, ending after 20-30 years in a cachectic state. The disease appears somewhat analogous to that in Brazil described by Splendore & Lutz. Cells ovoid, rarely spherical, 1.9-8/t in diameter, with granules staining purple with Giemsa stain while protoplasm stains blue. Cells solitary or in pairs, reproducing by sprouting. Organism grows well on Sabouraud media, glucose and sucrose solutions, milk, potato, carrot, Arracacha esculenta, Ipomoea datatis and Oxalis tuherosa. Cryptococcus sp. Irons & Graham, Trans. Chicago Path. Soc. 6: 445-448, 1906; Jour. Infect. Dis. 3: 666-682, 1906. Coccidioides immitis strain 2 Hamburger, Jour. Infect. Dis. 4: 201-209, 1907. The characters as given by Irons & Graham show a close similarity to the mycelium of Zymonema capsulatum and not to Coccidioides. Hamburger does not figure his strain. PSEUDOSACCHAROMYCES Pseudosaccharomyces van Laer, Bull. Ass. Beige Chimistes No. 3, 1893 ; Klocker, Centralbl. Bakt. II, 35: 375-388, 1912, not Samberger, Sbornik Klini- cky 5: 466-485, PI. 6, 1904, nor Briosi & Fameti, Atti 1st. Bot. R. Univ. Pavia II, 10: 31, 1906. Eansenia Zikes, Centralbl. Bakt. II, 30: 145-149, 1911, not Zopf, Zeitschr. Naturw. 56: 539, 1883, nor Lindner, Jahrb. Vers. Lehranstalt Brau. Berlin 7: 448, 1904. Kloeckeria Janke, Centralbl. Bakt. II, 69: 310, 1923 (spelling corrected to Kloeckera by Janke, Centralbl. Bakt. II, 76: 161, 1929). I have been unable to locate a copy of van Laer's publication and hence I am not sure as to the synonymy of Haiisenia Zikes and Kloeckera Janke. The type species of the two latter is Saccharomyces apiculatus Reess 342 MEDICAL MYCOLOGY (asporogenous strains only). Bamberger uses Pseudosaccharomyces for any asporogenous yeast without reference to spore shape. His use became the type of Parasaccharomyces (see p. 265). Briosi & Farneti follow Samberger in their definition but apply it to an organism of wholly different affinities. Pseudosac- charomyces must either be dropped as a permanent source of confusion or used in the original sense of van Laer. If the former is done, Kloeckera is the next valid name for the group. Klocker (1912) restricted Pseudosaccharomyces to asporogenous yeasts with citriform cells and described 16 new species. Cif erri & Redaelli, Ann. Myc. 27: 243, 1929, and Lodder, Anaskosporogenen Hefen 1: 232, 1934, characterize Kloeckera as follows: Cells usually apiculate or citriform, with occasional ellipsoid cells; not producing ascospores, single or only slightly clinging together, smooth, hyaline or bright colored; usually fermenting and producing acid in sugars; assimilat- ing peptone only ; no growth on alcohol ; gelatin liquefied. This genus is saprophytic, usually isolated from soils. No mammalian pathogens so far reported. ATELOSACCHAROMYCES Atelosaccharomyces Beurmann & Gougerot, Tribune Med. 42: 502, 1909. The type species is Atelosaccharomyces Busse-Buschki (Cryptococcus homi- nis). To illustrate the morphology of the genus the authors figure the Crypto- coccus of Gotti & Brazzola {A. Gottii) and Cryptococcus guttulafus (Saccharo- mycopsis guttulatus) . Cells always thick-walled in tissue, usually so on media in the adult stage ; sediment usually slimy, pellicle rare; gelatin not liquefied, sugars fermented. This genus was originally conceived as a name for all the asporogenous yeasts, in which sense it is synonymous with Cryptococcus, a much older name. By emending the description, however, to include only those forms which ferment sugars and do not liquefy gelatin, we have a very serviceable genus to cover a group of pathogens. Unless one is to disregard all the work of most of the older authors and develop a wholly new set of criteria of genera, this seems the best arrangement. Even so several species are of doubtful position because of brief descriptions. There is practically not a single char- acter of strictly generic or specific rank which has been recorded for every species so far described in the imperfect yeasts. It is to be hoped that by calling attention to the older literature, that some of these species may be found again and described in sufficient detail to show their place in the clas- sification. Key to Species Ferments glucose and fructose only. Colony white on most media, becoming grayish brown on potato. A. hominis. Colony white on most media, becoming yellowish on potato; almost no growth on blood serum. A. Catanei. SACCHABOMYCETACEAE IMPEBFECTAE 343 Colony yellowish white on Sabouraud agar, brownish in age on potato glycerol. A. Conori. Colony white, becoming brownish on malt agar, grayish white and dry on potato, light yellow and moist on blood serum. A. membranogenes. Ferments sucrose and maltose, milk coagulated. Cells spherical, 4-8/i, colony on carrot white, smooth, humid. A. laryngitidis. Cells ellipsoid, 7-10 x dfi, colony on carrot white, finally becoming pale rose, granular pulverulent. A. clericus. Ferments sucrose, maltose, and galactose. Cells spherical, 6-9fi. A. pyogenes. Atelosaccharomyces hominis (Viiillemin) Proilano de Mello, Paes & Sousa, Arq. Hig. Pat. Exot. 6: 34, 1918. Saccharomyces sp. Busse, Centralbl. Bakt. 16: 175-180, 4 figs., 1894; Arch. Path. Aiiat. Physiol. 140: 23-46, Pis. 1-2, 1895; Ihid. 144: 360^372, 1896. Cryptococcus hominis Viiillemin in Gedoelst, Champ. Paras. Homme 50- 51, 1902. Atelosaccharomyces Busse-Bvschki Beiirmann & Goiigerot, Nonvelle Mycoses 29, 1909 (cf. Hudelo, Benrmann & Gougerot, 1911). Torulopsis hominis Castellani & Jaeono, Jour. Trop. Med. Hyg. 36: 315, 1933. Not Saccharomyces hominis Klein & Gordon (1904). Isolated from subperiosteal suppuration of the tibia with purulent osteitis terminating in a generalized infection. Must be differentiated from syphilis or tuberculosis. Pathogenic to rabbits and mice. Cells ovoid or spherical, thick-walled, elements united in a common homo- geneous gelatinous sheath. Multiplication by sprouting. Neither mycelium, asci, nor spores seen. Optimum growing temperature 37° C. Growth good on glycerol agar. On potato, colony is dirty white, later becoming grayish brown, viscous. On gelatin, colonies small, white, shining, rounded after 24 hours, later elevated. Colonies on coagulated serum dirty white, confluent in a thick layer. In broth, a thick whitish sediment appears. Prune decoction turns dirty white with thick sediment. Gelatin not liquefied. Atelosaccharomyces Catanei Dodge, n. sp. Cryptococcus sp. Catanei, Arch. Inst. Pasteur Algerie 3: 384, 385, 1925. Isolated from cases of black pilose tongue along with CasteUania linguae- pilosae. Not pathogenic for guinea pig or rabbit. Cells in liquid media small, 2-4. 5/a in diameter, spherical or ovoid ; on solid media, slightly larger, 4-5.3 x 2-2. 6/i,, budding but not more than 3 cells cling- ing together at one time. Rarely slightly longer cells appear on potato and Sabouraud conservation agar. On Sabouraud glucose, colonies round, white, creamy, shining ; on con- servation agar, colonies very small and opaline. On gelatin, colonies small, white, growing along line of stab without arborescence ; practically no growth on serum. On potato, colonies slightly yellowish, dull. On potato glycerol, colonies lighter and creamy; on carrot, colonies very small. On broth and 344 MEDICAL MYCOLOGY peptone solution, no pellicle or ring, sediment light. Glucose and fructose fermented with acid production, no action on other sugars. No action on milk, serum, or gelatin. Atelosaccharomyces Conori Dodge, n. sp. Cryptococcus sp. Conor, Arch. Inst. Pasteur Tunis 1912: 215-218, 1912. (Case history of Conor & Bruch, Bull. Soc. Path. Exot. 4: 360-368, 1911.) Isolated from ulcer with small nodules, giving a cheesy matter, at base of skull and neck. Ulcer failed to respond to treatment and infant died in about 6 months from onset. Organism produced lesions on liver, spleen, and lungs of experimental animals. Cells spherical or ovoid, 4-10/x in diameter, no trace of hyphae, cells oc- curring singly or in pairs, very rarely in three's. No change of morphology on desiccation or aging of culture (up to one year). No ascospores found. Growth very slow at 15° -18°, optimum at 37° C. Colonies on Sabouraud agar, yellowish white, creamy, slightly elevated, and regularly rounded. On Sabouraud with tartaric acid, 1 :1,000, growth much more rapid with same appearance as the preceding. On potato, growth thick, whitish, mammillate, becoming pebbled on drying. On glycerol potato, growth moist, shining, with sediment on bottom of tube, brownish in age. On carrot, growth quite scanty, whitish. Peptone broth clear with slight sediment. In acid glucose broth, growth more rapid with floccose sediment. Glucose and fructose fermented. No action on maltose, lactose, or mannite. Atelosaccharomyces membranogenes (Steinhaus) Dodge, n. comb. Saccharomyces membranogenes Steinhaus, Centralbl. Bakt. I, 43: 49-69, 1907. Cryptococcus membranogenes Vuillemin, Champ. Parasit. Homme Anim. 94, 1931. Pound in diphtheric false membranes. Pathogenic for rabbit, guinea pig, and mouse, producing an acute inflammation with hemorrhagic exudate. Per- haps Strain II of Gentzsch (1908), isolated from diabetic urine drawn from the bladder under sterile conditions, should be referred here. Cells spherical, the size of a red blood cell, multiplying by sprouting from any portion of the cell, no capsule in cultures. On 3% malt agar, colonies circular, moist, white, soon confluent, finally becoming brownish. On potato, growth dry, gray Avhite, circular, confluent. On Loeffler's blood serum, colony mcist, circular, light yellow. On gelatin, colonies white, shining, granular, at times surface suggests fat. Only glucose fermented. Atelosaccharomyces larjmgitidis (Sartory, Petges & Claque) Dodge, n. comb. Cryptococcus laryngitidis Sartory, Petges & Claque, C. R. Soc. Biol. 84: 179, 180, 1923. Monilia laryngitidis Vuillemin, Champ. Parasit. Homme Anim. 84, 1931. Torulopsis laryngitidis Almeida, Annaes Fac. Med. Sao Paulo 9: 10, 1933. SACCHAROMYCETACEAE IMPERFECTAE 345 Producing pharyngolaryngitis. Symptoms are a tickling sensation in throat, burning sensation in air passages, continual cough with abundant ex- pectoration. Free edge of epiglottis and upper face of arytenoid cartilage had principal lesions which appeared as large, protruding, opaline plaques separated by narrow bands of healthy tissue. No lesions in larynx. No tuberculosis or syphilis. Lesions finally cured by medication with potassium iodide. Cells 4-8/i in diameter ; ascospores not seen. On carrot, optimum tempera- ture for growth was 28°-32° C, growth ceases at 39°-40°, but colony lived for 15 days at least at this temperature. Pellicle on broth at 25°-28° and 32°, but not at 35°-37°. On agar and gelatin plates, colonies round and white. On potato, colo- nies white, turning gray later. On carrot, colonies white, smooth, humid. On gelatin stab, growth in depths as well as at surface. Pellicle in broth. In sugar broths, growth is active, with the production of a uniform cloudiness. Glucose and maltose fermented, but not lactose or galactose. Sucrose is in- verted. Milk coagulated on twelfth day but casein not peptonized. Gelatin not liquefied. Atelosaccharomyces clericus Dodge, n. sp. Cryptococcus sp. Clerc & Sartory, C. R. Soc. Biol. 64: 135-137, 1908. Isolated from a case of chronic angina with small yellowish concretions on pillars of fauces, tonsils, and posterior wall of pharynx. Pathogenic to guinea pig. Cells ovoid, 7-10 x 5/a, isolated or in groups of five or six, sprouting at one pole. No mycelium or spores on any medium tried. Gram-positive. Optimum temperature 30° C. Colony on agar white, on potato dirty white. On carrot, colony at first smooth and pure white, later becoming granular, pulverulent, and pale rose in color. Poor growth on serum. Alcoholic fermentation of sucrose and maltose but not galactose. No aldehyde formation. Sucrose inverted. Starch not digested. Gelatin not liquefied, milk coagulated but no digestion of curd. Atelosaccharomyces pyogenes (Mattlet) Dodge, n. comb. Cryptococcus pyogenes Mattlet, Ann. Soc. Beige Med. Trop. 6: 13-14, 1926. On potato decoction, at 37° C, appear yeast forms, spherical, rarely ovoid, 6-9/* in diameter, multiplying only by sprouting. No fat granules. On Sabouraud agar, colonies circular, dull, slightly yellowish at the center and crumpling with fine radial striations, giving the margin a lacy appearance. On potato decoction, there is formed a coherent, membranous sediment Avhich breaks up on shaking. Acid formation and very active fermentation with glucose, fructose, maltose, galactose, and sucrose. No action on gelatin or milk. This organism is close to Cryptococcus hominis Vuillemin, but differs in its active fermentive power. 346 MEDICAL MYCOLOGY Perhaps the unnamed species of Cryptococcus described by Brazzola be- longs here. Cryptococcus sp. Brazzola, Mem. K. Accad. Sci. Bologna V, 6: 303-310, 1 pi., 1897 (Sez. Med. Chirurg. 43-50). Isolated from a case of gangrenous pharyngitis with infiltration of all the glands of the neck. Pathogenic to guinea pigs. Cells of variable size and irregular groupings, walls thick. On agar, glycerol agar, and gelatin, colonies round, margins slightly fimbriate, center thicker, yellowish white, becoming darker on aging. On potato, growth rapid, colony thick, mammillate, margin fimbriate, pearl white becoming dark yellowish, especially after exposure to rather intense light. On broth, a slight pellicle and no turbidity. On milk, growth luxurious with- out coagulation. Fermentation of glucose, maltose, and lactose. Optimum temperature between 30° and 35° C. Doubtful Position Atelosaccharomyces Gottii (Neveu-Lemaire) Froilano de Mello, Paes & Sousa, Arq. Hig. Pat. Exot. 6: 34, 1918. Cryptococcus sp. Gotti & Brazzola, Mem. R. Accad. Sci. Bologna V, 6: 721-754, 2 ph., 1897 [Sez. Med. Chirurg. 257-290]. Atelosaccharomyces sp. Beurmann & Gougerot, Bull. Mem. Soc. Med. Hop. Paris III, 28: 251, 1909. Isolated from lesions in tJie nasal passages of a horse. Pathogenic for guinea pigs. Cells from the lesion encapsulated, large, slightly ovoid or spherical, sprouting. In cultures capsule absent or slightly developed. On glycerol agar, white to mother-of-pearl colonies, creamy to almost gelatinous, margins slightly fringed, browning in age. On potato, growth rather rapid, colony rather thick, creamy, at first white, then grayish and finally gray brown, or even tobacco brown if exposed to light. On gelatin, colonies mother-of-pearl white, margins fringed, becoming granular and ir- regular, yellowish gray but not brown. No growth on coagulated serum. Growth poor on liquid media, no turbidity and no pellicle, with a slight sedi- ment at the bottom of the tube. Gelatin not liquefied. The position of this species is uncertain since no fermentation is recorded, and it is quite likely it belongs in Cryptococcus or even in Zymonema, since it was not observed long enough to be sure that no hyphae were produced. Sprouting occurs only at the ends of the cells, and occasionally the cells cling together in short chains. Cryptococcus niger (Mafucci & Sirleo) Gedoelst, Champ. Paras. Homme 59, 1902. Saccharomyces niger Mafucci & Sirleo, Policlinico, Sez. Chirurg. 2: 138- 144, 245-254, 1895. Torulopsis niger Almeida, Annaes Fac. Med. Sao Paulo 9: 10, 1933. SACCHAROMYCETACEAE IMPERFECTAE 347 Isolated from pulmonary myxoma of a guinea pig. Cells small, sprouting, dark. Colonies on agar, circular, whitish, covering the whole surface. On potato, colonies grayish and chocolate color. In glucose broth, sediment but no pellicle. Good growth in milk. Alcoholic fermentation of malt extract. Gelatin not liquefied. Little growth on gelatin, less on blood serum, and very little on beef broth. Quite probably this species should be transferred to the Toruleae. Cryptococcus sp. Breed, Arch. Int. Med. 10: 108-121, Figs. 1-6, 1912. Isolated from the respiratory tract of man, also from one case of severe vaginitis. Pathogenic for rabbits, white rats, guinea pigs, and monkeys. Cure effected by medication with sodium iodide. Cells size of a red blood corpuscle. No spores found. Grows between 18° and 69° C, though slight at room temperature. Killed in one hour at 71° C. Viable dry for 2 years. In 12-24 hours at 37° on glucose and glycerol agars there is a profuse creamy growth. On potato slant under similar conditions, a slight film forms. Grows on plain or glucose broth with production of heavy sediment (and foam with latter), on shaking. Good growth on plum and grape decoctions but no foam. Growth on litmus milk creamy white with soft curd in 24 hours. Glucose fermented. Growth but no fermentation on lactose, raffinose, inulin, and mannite. Cryptococcus de Bumier (cas Th) Ota, Ann. Parasitol. Hum. Comp. 2: 44-45, Fig. 5, 1924. Isolated from a case of epidermomycosis. On malt agar, at 25° C. in day-old cultures, cells ovoid or slightly elon- gate, solitary or in chains of a few cells, 6-8 x 4-5/i., wall thin. After a week, cells 12 X 10/t, spherical or ellipsoid, wall thicker. On malt agar, colony grayish white becoming yellowish, smooth, humid, margin definite. On malt extract, forming a thin ring and sediment. TORULOPSIS Torulopsis Berlese, Giorn. Viticoltura ed Enologia 54, 1894. I have been unable to locate the original publication of this name, but it was designed to end the confusion caused by the application of Torula to two different groups of organ- isms (see p. 327). This has been applied to the asporogenous yeasts, following the traditional application of Torula by Turpin, Mem. Acad. Sci. 17: 1-88, 1838, Pasteur, fitudes sur la Biere 73, 1876, Hansen, Medd. Carlsberg Lab. 2: 87-93 [47-52], 1888, Will (1916), etc. Turpin prob- ably included both asporogenous and sporogenous strains and emphasized fermentation; Pasteur held the same concept, but denied fermentation. Hansen was the first to use the name in the modern sense, applying it to asporogenous yeasts without regard to fermentative power. In proposing this genus name, Berlese gave a good resume of the characters given by Hansen, limiting its application definitely to the forms without mycelium, and excluding the species with citriform cells (see Pseudosaccharomyces) . Will (1916, 1917) still further re- stricted the generic concept (although still using the name Torula), defining it as follows: 348 MEDICAL MYCOLOGY Young cells with refractive protoplasm, no oil globules, pellicle slimy, usually arising as separate floating islets which are confluent, glassy but tough, sediment more or less gelatinous. Ciferri So Redaelli (1929) still further modify it by excluding species with agglomerations of cells or pseudomycelium. Since so many of the species are pink or reddish, these authors have discarded Berlese's Tarulopsis rosea as the type species in favor of Torula gelatinosa Will, since they claim that the adoption of such a poorly described siiecies would be a perma- nent source of error. They characterize their genus as follows: Cells ellipsoid, spherical, or irregular, never citriform or catenulate, rarely, and this particularly in liquid cultures, with formation of irregular sprouting agglomerates of crowns and elongated cells, cells continuous, hyaline or light colored, not forming endo- spores, mycelium or pseudomycelium. Cell wall thin, young cells without the oil globules which may appear in old age ; superficial vegetation with giant colonies according to Will's Type III; little or no fermentative power, usually forming acids on sugar media ; producing hydrogen sulphide and coagulating milk ; producing a carotin pigment, hence colors some shade of orange or red. Key to Species Cells large, 8-12;tt, spherical to elongate. Glycerol assimilated. T. cavicola. Glycerol not assimilated. T. bronchialis. Cells smaller, rarely over 8fi. No ring to liquid media. Gelatin liquefied, colony rose becoming the color of eosin powder. T. mena. Gelatin not liquefied. Milk coagulated, colony orange yellow. T. Sangiorgii. Milk not coagulated, colony reddish orange. T. mitis. Ring and often floating islets but no pellicle on liquid media. Gelatin liquefied. Nitrate assimilated. T. glutinis. Nitrate not assimilated. Cells long ellipsoid, often curved, milk not clotted. T. rubra. Cells spherical or short ellipsoid, milk clotted. Cells 4-5 X 5-7fi. T. Sanniei. Cells 2.5-4 X 4-6/x, more slender. T. mucilaginosa. Tomlopsis cavicola (Artault) Almeida, Annaes Fac. Med. Sao Paulo 9: 10, 1933. Cryptococcus cavicola Arthault, Arch, de Parasitol. 1: 259-265, 1898. (Vuillemin, Rev. Gen. Sci. Pures Appl. 12: 740, 1901, suggests that this is close to or a strain of Saccharomyces Fresenii or 8. roseus.) Isolated from pseudotuberculosis in lung of guinea pig. Only slightly pathogenic to rabbit on subcutaneous injection. Inoculated into eye of rab- bit, it produced lesion from which organism was reisolated. Cells ovoid, 8-12yM, solitary or, rarely, in chains of three. On agar, growth slow, suggesting Serratia marcescens (Bacillus prodigi- osus). On potato, colony similar to that on Raulin's, but thicker, redder, SACCHAROMYCETACEAE IMPERPECTAE 349 and moister. On Raulin's liquid, colonies rose to clear vermilion, margin irregular, surface shining to oily, becoming 1 mm. thick and spreading over the whole surface of the medium. On 5% or 10% glucose solution no pellicle appears, very slight pigment, medium turbid, no fermentation (?). Organism attacks oil used to secure anaerobiosis, apparently assimilating glycerol and leaving floating masses of stearic acid crystals at interface of oil and medium. Torulopsis bronchialis Ciferri & Redaelli, Atti 1st. Bot. R. Univ. Pavia, ni, 2: 245-248, 283, 1925. Bhodotorula hroncliialis Lodder, Anaskosporogenen Hefen 1: 91-93, 1934. Isolated from sputum of a patient with bronchopneumonia, not pathogenic for laboratory animals. Cells ovoid, at first hyaline, then guttulate, 7-7.5/a in diameter up to 9-9.5 X 10/x, often forming short chains. On malt extract, 3-4 x 5-8/t, singly or in pairs. No spores or mycelium. On carrot agar, colonies dense, creamy, radially folded, opaque, shining, oehraceous to reddish. On malt agar, colony intense orange, moist, smooth, margin pellucid. On gelatin, colonies red orange, verrucose radially folded. On carrot, colonies small, uniform, surface irregular, reddish. On potato, colonies spreading, flesh red, uniform, margins smooth, becoming irregularly granulose. On malt extract, a pulverulent pellicle formed by confluence of single colonies with granular sediment. On glucose broth, similar, pellicle pink. Methylene blue reduced slowly. On milk, pellicle almost white, casein precipitated after one month and very sowly digested. No alcoholic fermenta- tion ; acid with glucose, fructose, galactose, sucrose, lactose, and inulin. Lac- tose not assimilated (Lodder). Torulopsis mena (Fontoynont & Boucher), Dodge, n. comb. Cryptococcus mena Fontoynont (nom. nud.), Bull. Mem. Soc. Chirurg, Paris 48: 439-442, 1922; Fontoynont & Boucher, Ann. Derm. Syphiligr. VI, 4: 213-235, 3 figs., 1923; Ota, Ann. Parasitol. Hum. Comp. 2: 45-47, 1924. Isolated from ulcerations on great toe and legs, with severe lesions on chest due to earlier unsuccessful surgical interference when lesion was mis- taken for cold tuberculous abscess. Medication with potassium iodide, etc., wholly unsuccessful ; in fact, KI made lesions worse. Final cure effected by internal and external applications of methylene blue. Even in pure cultures, 0.0015-0.002% methylene blue checked growth. Organism secretes a toxin fatal to rat and guinea pig. No agglutination of Cryptococcus with patient's blood nor with other blood, but agglutinations prompt in presence of meth- ylene blue. Yeast cells ellipsoid, 4-5ju. in diameter, sprouting on second to fourth day. By thirtieth day cells become more rounded, cell walls smooth. Optimum temperature 20° but growth good also at 37° C. On glucose agar, colony rose, finally becoming brick red, attaining di- ameter of 2.5 cm. in 5 days. On maltose agar, colony rose brown. On poor agar, colony develops very slowly with production of bad odor. On potato. 350 MEDICAL MYCOLOGY colony rose, terra cotta, then red brown. On glucose gelatin, colony rose by reflected light, grayish white beneath and by transmitted light, later becom- ing color of eosin powder. In glucose broth, liquid remains clear with sediment which is white at first, slowly turning rose. Gelatin slowly liquefied. Pigment production varies with temperature, being less at higher temperature, also less when organism is freshly isolated. Var. Mekimdu (Mattlet) Dodge, n. comb. Cryptococcus Mekundu Mattlet, Ann. Soc. Beige Med. Trop. 6: 14, 15, 1926. Isolated from sputum in case of bronchitis in Belgian Congo. Patient had been weakened by malaria, bronchitis accompanied by a low continuous fever with periodic exacerbations suggesting undulant fever, but no aggluti- nation with Micrococcus melitensis. Patient removed to Europe shortly after organism was isolated and not further studied. In sputum, cells 3-4/* in diameter. On potato decoction at 25°, cells spheri- cal or ovoid, walls thin homogeneous on third day, sprouting, 2-6/a in diameter. Older cells have thick walls. On Sabouraud agar, colony red, circular, soon flowing to bottom of slant. On potato decoction, rose sediment on bottom and walls, clots disappearing on agitation. Gelatin liquefied, optimum temperature 25° C. Slight acid pro- duction in, and coagulation of, milk. Acid with glucose, fructose, galactose, sucrose, and lactose. Close to T. mena, but interior of cells less granular. Torulopsis Sang^orgii Dodge, n. sp. Cryptococcus sp. Sangiorgi, Centralbl. Bakt. I, 63: 58-62, 1912. Isolated from the peripheral blood stream and from the liver of a dog after 5 months of fever. Pathogenic for dog, slightly for rabbit. Cells in liquid media ovoid or ellipsoid, on solid media similar or very rarely spherical. Extreme dimensions 4-6 x 3-5/*. Sprouting from the ends of cells on solid media producing chains of 2-4 cells but no true hyphae. Growth at 20° and 27° C, the higher temperature best for acid media. On Sabouraud agar, colony rosy, moist, creamy. On lactose agar, colony very superficial. Growth slight on plain agar, blood agar, and coagulated horse serum. On potato, colony thick, spreading, creamy, moist below, and powdery above. Colonies orange yellow on most media but grayish white on blood agar. On gelatin, growth slow% penetrating the medium. In plain broth, liquid remains clears, no pellicle, and slight sediment. In glycerol broth, sediment on bottom and walls but no pellicle. No indol formation in peptone solution. No fermentation of glucose, maltose, lactose, dextrin, or inulin, no inversion of sucrose. Milk coagulated in 27-30 days. Gelatin not liquefied in 90 days. Torulopsis mitis (Mazza, Stabile de Nucci & Canal Feijoo) Dodge, n. comb. Cryptococcus mitis Mazza, Stabile de Nucci & Canal Feijoo, Bol. Inst. Clin. Quirurg. Univ. Buenos Aires 5: 293-308, 21 figs., 1930. I SACCIIAROMYCETAC'EAE IMPERFECTAE 351 Producing pustules, origiiiall}^ appearing on tiie face, subsequently spread- ing to all external parts of the body, in a 10-year-old Argentine boy. Lesions secreting a serous, sanguinolent liquid. ^Medication with increasing doses of potassium iodide caused eventual complete cicatrization of the lesions. Emul- sion of yeast isolated and reinjected into dermis of patient caused local reac- tion. Emulsions of organism cultivated on ISabouraud agar injected intra- peritoneally into Cebus monkey, rabbit, guinea pig, two mice and two rats, nonpathogenic. Scarifications and rubbing with organism caused no lesions on monkey. Subcutaneous injection into white rat caused appearance of nodule v\diich, removed and examined, showed presence of organism. On Sabouraud agar, cells round, averaging 5/x. in diameter, possessing a thick membrane with an unevenly granular content. No hyphae or asci seen after long observation. Reproduction by sprouting. Gram-positive. On Sabouraud agar, at 37° C, no growth. At room temperature after 3 days, orange red colonies, slightly elevated at center, scalloped margin. After 10 days colonies elevated, margin undulate, with a few radial furrows. Organism grows well on potato or carrot and glycerol, also on plain agar and Gougerot gelatin and always at room temperature. In plain broth, slight cloudiness in 48 hours. In Sabouraud broth, general turbidity. Fermenta- tion after 48 hours with raffinose and fructose, none with sucrose, inulin, lactose, mannite, maltose, galactose, sorbite, glucose, dextrin, or arabinose. No indol formation. Milk not coagulated. Torulopsis glutinis (Fresenius ampl. Harrison) Dodge, n. comb. Cryptococcus glutinis Fresenius, Beitr. Mykol. 2: 77-78, 1852. Saccharomyces glutinis Cohn, Beitr. Biol. Pflanzen 1: 187, 1875 (cf. Hansen, Medd. Carlsberg Lab. 1: 253-264 [81-88] 1879, etc.). Torula glutinis Pringsheim & Bilewsky, Beitr. Biol. Pflanzen 10: 118, 1910. Rhodotorula glutinis Harrison, Trans. R. Soc. Canada 22: 187, 1928 [cf. Lodder, Anaskosporogenen Hefen 1: 65-68, 1934]. The following pink yeasts rather imperfectly described are often referred here but may belong to other species of this genus, Saccharomyces roseus Engel, Bull. Soc. Sci. Nancy 6: 1, 1877 (cf. Mag- giora & Gradenigo, 1896). Saccharomyces rosaceus Crookshank, Introd. Pract. Bact. 224, 1886 [cf. Vuillemin & Legrain, Arch, de Parasitol. 3: 260-266, 1900]. Saccharomyces Fresenii Schroeter, Kryptog. Fl. Schlesien 3: 2: 208, 1893. This species seems to be primarily a saprophyte, not pathogenic for labo- ratory animals, although from time to time reported as isolated from otitis media, etc., perhaps as a contaminant, as this group of pink yeasts is fre- quently found in old laboratory cultures. The following description is based on culture of Pringsheim & Bilewsky isolated from air, amplified by Har- rison and Lodder who studied the same strain. 352 MEDICAL MYCOLOGY Cells ellipsoid, 3-4.5 x 5-7/u,, mostly single or in pairs, rarely in small chains. On malt agar, colony red with orange tone, moist, smooth, slimy. On malt gelatin, colony shining, rose, punctate, margin dentate. On malt extract, slimy islets, a broad ring, with thick slimy sediment, liquid turbid, rose color. No fermentation ; acid with glucose, fructose, and mannose ; potassium nitrate assimilated ; gelatin liquefied. Tonilopsis rubra (Demne) Almeida, Annaes Fac. Med. Sao Paulo 9: 10, 1933. Saccharomyces ruher Demne, Ann. Microgr. 2: 555, 556, 1889 [1890] ; Mitt. Naturf. Ges. Bern 1889: ix-xi, 1890; Festschr. E. Kenoch Berlin, 1890; Casa- grandi, Ann. Ig. Sperim. 8: 535-545, 1897-1898. Cryptococcus ruher Vuillemin, Bull. Seances Soc. Sci. Nancy III, 1: 61, 1900. Rhodotorula rubra Lodder, Anaskosporogenen Hefen 1: 69, 1934, Originally isolated from a red cheese and later from feces of children Avith diarrhea who drank some of the milk from which the cheese was made. Thought by Demne not to be j)athogenic, although when pure cultures were fed to dogs in large doses, digestive disturbances resulted. Casagrandi iso- lated the organism from diabetic urine which had been exposed to the air. Recently Bagnacci (1926) reports this species in glossitis of an infant. Patho- genie in subcutaneous injections to guinea pigs and rabbits, also by ingestion of milk cultures which produce diarrhea. Cells 3.8-4.5-6.8/x in diameter, in short chains of 3-4 cells. Chlamydospores 7.5-8.5 X 5-5.5/x. Grows on glucose agar, better on glycerol agar, producing pale rose colo- nies which become reddish chestnut in color and are elevated, shining, with margin lobed. On Pollacci agar, colony at first whitish becoming red, plane or with center elevated, moist, shining, with margin lobed. On malt agar, colony orange, moist, smooth, margin pellucid. Growth on potato even bet- ter, colony red, 2-3 mm. thick. No growth in agar with more than 5% tartaric acid. On gelatin, colonies yellowish at first becoming clear raspberry red, nailhead size, with whitish lobed margin. In glucose broth, there is a whitish pellicle, reddish sediment, and no turbidity. Casagrandi reports fermentation of glucose and sucrose, others report no fermentation. Sucrose inverted. Milk not acidified or coagulated. Gelatin liquefied in 2-4-10 months. Tonilopsis Sanniei Ciferri & Redaelli, Atti 1st. Bot. R. Univ. Pavia III, 2: 245-248, 283, 1925. Rhodotorula Sannei Lodder, Anaskosporogenen Hefen 1: 99-101, 1934. Isolated from lungs. Not pathogenic for laboratory animals. Cells ovoid or ellipsoid, often spherical, single, at first hyaline then gut- tulate, 4-5 x 6-7.5ft, no ascospores or mycelium. On carrot agar, colonies peach blossom red, margins somewhat serrate, then the surface wrinkled and brick red. On must gelatin, colonies round with fine concentric furrows, blood red. On malt gelatin, colonies moist, saccharomYcetaceae imperfectae 353 almost, smooth, slightly punctate in the middle, margin smooth. On malt ex- tract agar, colonies red to slightly orange, shining, with small warts and sinuous margin. On malt extract, ring thin, thick sediment. No fermentation of sugars. Milk coagulated, gelatin liquefied (denied by Lodder). Torulopsis mucilaginosa (Jorgensen) Ciferri & Redaelli, Atti 1st. Bot. R. Univ. Pavia III, 2: 147, 1925. Torula mucilaginosa Jorgensen, Die Mikroorg. d. Garungsindustrie ed. 5, 402, 1909. Bhodotorula mucilaginosa Harrison, Trans. R. Soc. Canada 22: 187, 1928. Cryptococcus ruhrorugosus Castellani, Arch. Derm. Syphilol. 16: 403, 1927. Bhodotorula mucilaginosa race ruhrorugosa Lodder, Anaskosporogenen Hefen 1: 104-105, 1934. Originally isolated as a laboratory contaminant. Castellani reports it from scrapings in the axilla. Cells 2.5-3.8 x 3.2-5/i,, single or in pairs. On malt agar, colony intense red, dull, smooth with small warts in the center, margin smooth. On malt gelatin, colony red, center elevated with radial ridges, margin smooth. In malt extract, a broad ring and thick, slimy sediment. Good growth in alcohol. No fermentation; gelatin not liquefied. Var. pararosea (Lodder) Dodge, n. comb. Cryptococcus pararoseus Castellani, Arch. Derm. Syphilol. 16: 402, 1927; Amer. Jour. Trop. Med. 8: 393, 1928. Bhodotorida mucilaginosa var. pararosea Lodder, Anaskosporogenen Hefen 1: 102-104, 1934. Isolated from sputum in a case of chronic bronchitis. Cells ovoid to long ellipsoid, 2.5-4 x 5-9/x, single or in pairs in malt extract ; a little shorter, 2.5-3.5 x 4-7.5/a, in malt agar. Colony orange red, moist, elevated in the middle with radial folds, margin pellucid, ragged, somewhat slimy. On malt gelatin, colony red, with central knob, radial furrows, and sinuous margin. In malt extract, a ring and sedi- ment produced. No fermentation, acid only with glucose and fructose. Growth good in alcohol. Gelatin not liquefied. Var. plicata (Lodder) Dodge, n. comb. Bhodotorula mucilaginosa var. plicata Lodder, Anaskosporogenen Hefen 1: 108-110, 1934. Cryptococcus corallinus A. Sartory, R. Sartory, Hufschmitt & J. Meyer, C. R. Soc. Biol. 104: 1316-1318, 1930. Torulopsis corallina Ciferri & Redaelli, Arch. Protistenk. 71: 430, 1930 not T. corallina (Saito) Ciferri & Redaelli, Atti 1st. Bot. R. Univ. Pavia III, 2: 147, 1925. Isolated from hair in pruriginous nodular lesions suggestive of tricho- phytie kerions. In malt extract, cells short, ovoid, 2-3 x 4.5-6. 5/x, solitary or in groups of 1-3 cells. On malt agar, cells 2-4.5 x 4-5.5/a. On Sabouraud agar, 1.5-2.5/x. 354 MEDICAL MYCOLOGY On Sabouraud agar, colonies cream white, viscous, moist, furrowed in drying- (37° C.)- At 27° C, colonies pink, finally salmon or coral red, much wrinkled, almost cerebriform. On malt agar at 15° C, colonies deep red to slightly orange, shining, wrinkled, margin sinuous, pellucid. On malt gelatin at 15° C, colonies smooth, moist, slightly elevated in the center, margin smooth, dirty yellow. On Eaulin solution, slight cloudiness and rose-colored pellicle at 27° C. On malt extract at 25° C, ring and sediment. Glucose and maltose fermented (denied by Lodder), no action on lactose, fructose, ga- lactose, and inulin. Lodder claims that fructose is assimilated, both Ciferri and Lodder that galactose is slightly assimilated. Milk coagulated on third to fourth day, pellicle rose color, not digested. Egg albumen not liquefied, serum not coagulated, gelatin liquefied. Var. Carbonei (Lodder) Dodge, n. comb. Saccharomyees glutinis Carbone, herb. nom. not Cohn, 1875. Blast odendrion Carhonei Ciferri & Redaelli, Atti 1st. Bot. R. Univ. Pavia III 2: 147, 1925. Torulopsis nitritophila Ashford & Ciferri, Zentralbl. Bakt. II, 81: 63-67, 1930. Cryptococciis radiaius Sartory, Sartory & Meyer, C. R. Soc. Biol. 106: 597, 598, 1931. Probably first isolated as a contaminant, reported by Ashford & Ciferri from human feces and by A. Sartory, R. Sartory & J. Meyer as a saprophyte isolated during the study of an epidemic in which the disease was character- ized by falling hair. Cells spherical to short ovoid, 3.5-4.5 x 3.5-6/a, single or in pairs, slightly narrower on malt agar. On must gelatin, giant cells 6-10/4 in diameter, cells aggregated in mucilaginous masses. Colonies shining, bright salmon color, not folded. On Sabouraud agar, colonies English red, confluent, margin scalloped, with projections suggesting the pseudopodia of Radiolaria. On malt agar, colonies flame scarlet to orange chrome, moist, slimy, smooth, margin smooth. On malt gelatin, pale red,' humid, flat, punctate, margin sinuous. On malt extract, a thin ring and a thick, slimy light salmon orange, sediment. No fermentation ; no liquefaction of gelatin. Growth extremely good in neutral Raulin's solution with potas- sium nitrite (whence the name of Ashford & Ciferri 's strain from human feces). EUTORULA Eutorula Will, Centralbl. Bakt. II, 46: 226-281, 1916, non Euforula Sae- eardo as subgenus. Eiitorulopsis Ciferri, Atti 1st. Bot. R. Univ. Pavia II, 2: 141-143, 1925. The type species is Eutorula vulgaris Will. This renaming by Ciferri was unnecessary. He defines the genus as fol- lows : Cells ellipsoid, spherical, or irregular, not apiculate nor catenulate, SACCIIAROMYCETACEAE IMPERFECTAE 355 thin-walled, continuous, hyaline or light coJorod; no mycelium or pseudomy- celium ; no endospores, young cells nni- or plui-jguttulate ; pellicle dry, chalky white, often folded at tirst, becoming- smooth, citron yellow to brownish yel- low, clinging to the walls, ductile, or breaking, and portions floating on the sur- face. Sediment loose, often powdery. Gelatin liquefied, sugars fermented. Key to Species Ring, but no pellicle on glucose broth, colonies on potato glycerol dry, white, beginning to yellow about twenty-fifth day. E. fusca. Neither ring nor pellicle on glucose broth. Colony white or grayish, sediment white. E. Bernasconi. Colony ochraceous, sediment yellowish. E. irritans. Eutorula fusca (Fontoynont & Boucher) Dodge, n. comb. Cryptococcus fuscus Fontoynont & Boucher, Ann. Derm. Syphiligr. Yl, 4: 325-330, Fig. 6, 1923; Fontoynont (nomen nudum), Bull. Mem. Soc. Chirurg. Paris 48: 439-442, 1922. Torulopsis fusca Almeida, Annaes Fac. Med. Sao Paulo 9: 10, 1933. Isolated from ecthyma of the leg and mycetoma of the foot, with black grains (according to testimony of patient, but grains not seen by the authors). Pathogenic to rat, rabbit, and pigeon. Yeast cells ovoid, 6-7/a in diameter, thin-walled. On maltose or glucose agar, colonies white, becoming cafe-au-lait in color on eighth day and, finally, caramel colored. Elevated as in other slants. On "poor" agar, colonies small, not elevated, beginning to brown on the fifteenth day. Colonies on potato with glycerol, white and dry, 3 mm. thick, beginning to yellow on the twenty-fifth day. On sweet potato, colonies thick, creamy, colorless, shining, turning cafe-au-lait by the eighth day. Gas bubbles appear on thirteenth day. Growth begins to yellow on twenty-fifth day and by the third month turns a caramel color. On gelatin w^ith glucose, colonies thick, white, creamj'. Slow liquefaction after twenty-fifth day with liquefaction com- plete on the fifty-seventh day with a brown sediment. On glucose gelatin slant, colony hemispheric with a thin edge, turning caramel after a time. Glucose broth becomes turbid with large white sediment, ring but no pellicle. Liquid clears by the fiftieth day, with ring and deposit tui'ning cafe-au-lait in color. Optimum temperature for growth 20°-22° C, growth very slow at 37° C. Eutorula Bernasconi (Fontoynont & Boucher) Dodge, n. comb. Cryptococcus Bernasconi Fontoynont & Boucher, Ann. Derm. Syphiligr. VI, 4: 318-325, Figs. 4-5, 1923; Fontoynont, Bull. Mem. Soc. Chirurg. Paris 48: 439-442, 1922 (nomen nudum). Torulopsis Bernasconi Almeida, Annaes Fac. Med. Sao Paulo 9: 10, 1933. Cause of extensive ulceration of the skin, with multiple suppurating ade- nitis. Slightly pathogenic for rabbit. In pigeon secretes a soluble toxin which is fatal. 356 MEDICAL MYCOLOGY Yeast cells ovoid or pyrif orm the first day, mostly 4-5/1. in diameter but some as high as T/x, few sprouting forms. On second day, cells all pyriform. No ascospores observed. On glucose agar, colonies white, creamy. On maltose agar, the same, but with tendency to become thicker in the middle. Colony grayish on poor agar. On potato, colony dry and chalkj- white. In broth to which glucose has been added, white sediment, liquid clear, no pellicle or ring, some gas evolved. Gelatin is very slowly liquefied. This organism grows as well at 37° as at 20°-22° C. Eutorula irritans (Mattlet) Dodge, n. comb. Cryptococcus irritans Mattlet, Ann. Soc. Beige Med. Trop. 6: 15, 1926. Isolated from sputum of a patient in Belgian Congo, who had long attacks of coughing and dry rales. Greenish yellow purulent clots showing yeast cells were abundant in the sputum. Treated with potassium iodide and novar- senobenzol Billon. After 3 weeks, the symptoms and the yeasts disappeared from the sputum. Yeast cells in sputum 2-4/* in diameter. On potato decoction at 25° C, in 3 days, yeast cells 2-6/i, thin-walled. Old cells thick-walled with fine radial striations. On Sabouraud agar, colonies ocliraceous, circular, soon flowing. On potato decoction, deposit of yellowish flocci, disappearing on shaking, no pellicle. On gelatin, liquefaction of upper portion in shape of a cone. Optimum tem- perature 25° C. No action on milk. Organism ferments glucose, fructose, maltose, galactose, and lactose, not sucrose, mannite, dextrin, or inulin. ASPOROMYCES Asporomyces Chaborski, Recherehes sur les Levures Theromphiles et Cryo- philes, These Geneve 627: 26-30, 1918. The type species is Asporomyces asporus Chaborski. Reproduce only by sprouting, the yeast cells put forth copulation canals from time to time but no fusion occurs and no spores are produced. Asporomyces Mug-era (Mattlet) Dodge, n. comb. Cryptococcus Mugera Mattlet, Ann. Soc. Beldge Med. Trop. 6: 12-13, 1926. Frequently isolated from stools of dysentery patients in Belgian Congo. In potato decoction, small spherical yeast cells with a small oil globule, about 3/x in diameter in 3 days, very old cells up to Ifi, spherical with the wall thick and somewhat warted. Copulation forms observed between cells about 4/x in diameter, but no ascospores seen. On Sabouraud agar, colonies white, dull, circular, smooth, center of colony slightly yellowish in age. In potato decoction, slight turbidity with granular sediment, pellicle and ring formed. Optimum temperature 37° C. Milk, acid then neutral. Gelatin not liquefied. Acid formation in glucose, fructose, galactose, and sucrose. I SACCHAROMYCETACEAE IMPERFECTAE 357 MIOROBLASTOSPORION Micr'ohlastosporion Ciferri, Arch. Protistenk. 71: 420-424, 1930. The type species is Torula hotryoidea Chaborski. Sprout cells separate early from parent cell by wall, no polarity, no my- celium. Grown only on media used to secure spore production. Colony radially striate, lobed. In liquid media, no pellicle, slight ring and sediment, gelatin liquefied. TRIGONOPSIS Trigonopsis Schachner, Zeits. f. d. ges. Brauwesen 52: 137, 1929. The type species is Trigonopsis variabilis Schachner. Cells partly ellipsoid, partly appearing triangular, occasionally suggest- ing a crescent with rounded ends. Islets, a thin ring and sediment in liquid cultures ; no fermentation, no liquefaction of gelatin. Excluded Genera Schizotorulopsis Ciferri, Arch. Protistenk. 71: 431-435, 1930. The type, Schizotorulopsis Alfonsecai Ciferri, Arch. Protistenk. 71: 431- 435, 1930, was shown by Verkaik (1931) to be a culture of Bacilhis mega- therium Bary. As the following species have been so poorly described or even proposed as nomina nuda, they should be dropped until more adequate descriptions are available. Cryptococcus albus Hlava cited by Vuillemin, Rev. Gen. Sci. Pures Appl. 12: 741, 1901. Cryptococcus epidermidis Castellani & Chalmers, Man. Trop. Med. ed. 3, 1075, 1919. Torulopsis epidermidis Almeida, Annaes Fac. Med. Sao Paulo 9: 10, 1933. Cryptococcus Lowi Castellani, cited by Sartory, Champ. Paras. Homme Anim. 243, 921. Tropical blastomycosis. Cryptococcus Tonkini Castellani & Chalmers, Man. Trop. Med. ed. 2, 771, 1913. Blastomyces Tonkini Legendre, Bull. Soc. Path. Exot. 4: 24-26, 1911. Torulopsis Tonkini Almeida, Annaes Fac. Med. Sao Paulo 9: 10, 1933. Isolated from two cases of blastomycosis in Tonkin. In pus, cells in short chains of 4-5 cells. CHAPTER XIV MALASSEZIA Malassezia Baillou, Traite de Bot. Med. Cryptog. 243, 1889. Pityrosporum Sabouraud 1895; Castellani & Chalmers, Man. Trop. Med. ed. 2, 836-837, 1913. The type species is Microsporon furfur Robin. The members of this genus are obligate parasites confined to the outer layers of the epidermis and the sebaceous glands, not easily isolated and very easily dying out after the first transplant. In the scales, small yeast cells and occasional mycelium are shown. Colony dry and chalky. The organisms of this group have been too infrequently cultivated and too poorly described to be placed definitely, but their very difi'erent cultural requirements and their very specialized habitat suggest that they belong to a different genus. One can only speculate whether they represent a very de- generate and highly specialized dermatophyte related to Achorion or whether they are related to yeasts. Their habitat on the host, suggests the former alter- native, especially as the organism starts in the homy layer of the epidermis, and invades the hair follicle without attacking the hair shaft. The yeasts on the whole show much less narroAv specialization to a habitat, being rarely localized on the outer layers of the skin, and when in this situation produce much more acute lesions. Malass«zia furfur (Robin) Baillon, Traite Bot. Med. Cryptog. 243, 1889. Microsporon furfur Robin. Hist. Nat. Veg. Parasit. 436-439, 1853. Epidermophyton sj). Bazin, Lecons . . . sur les Affections Parasites de la Peau, 1862. Sporotrichum (Microsporon) furfur Saccardo, Syll. Fung. 4: 100, 1886. Oidium {Microsporon) furfur Zopf, Die Pilze, 257, 1890. Oidium subtile Kotliar, Vratche No. 12: 2055, 1892 [in Russian]. Monilia furfur Vuillemin, Champ. Parasit. Homme Anim. 89, 1931. This is the organism of pityriasis versicolor, isolated and described from the scales. Cultures reported by Schmitter, Jour. Trop. Med. Hyg. 26: 190- 194, 1923, and by Macleod & Dowling, British Jour. Derm. Syphilis 40: 139- 148, 1928. From his description, Kotliar probably had an Actinomyces species, Hyphae 2-3/* in diameter. Spores 3-8/* in diameter, spherical or ovoid, very refractive, often budding, striated with spiral striae formed by budding, often in clusters, not easily stained. Method of staining: fix scales in absolute alcohol, stain with light green, eosin, or even zinc chloriodide. On glycerol agar, there is a free, glistening, yellowish growth along streak. After a week, tiny white drops appear on the surface. These enlarge to 2 mm. in diameter and appear crateriform. Broth clouds with heavy sedi- 358 MALASSEZIA 359 ment settling, indol produced. Acid is formed in lactose broth. Litmus milk clotted in 2 days, decolorized and peptonized in 10-20 days. Gelatin liquefied. The organism cultivated and studied by Kambayashi (1932) is described as follows : Spores mostly ovoid or elongate ovoid, 5-6 x 4/^., occasionally nearly spherical in liquid media. Hyphae short, either pointed or rounded ends, 2.4-2. 6/i in diameter, straight or partially curved. In much older cultui-es, hyphae longer, richly branched, abjointing the ends of short lateral branches on a slightly differentiated conidiophore. Conidia sometimes in short chains of 2-3 cells, 3-5-6 x ifx. On Sabouraud, growth of primary culture very slow, about 9 months to produce a colony the size of a rice grain, whitish yellow, slightly verrucose. The first subculture attains about this size in 20 days. Colonies yelloAvish white, confluent, surface grooved and knotted, with a tendency for super- ficial grooves to be radial at the margins. The yellow of the young colony becomes lighter to grayish, then with age the yellowish color reappears and passes on to brown. Colony at first moist, becoming dryer and more chalky. The giant colony is a conical mass, about 3.5 cm. in diameter, with fine irregu- lar radial folds cut irregularly by cross or circular folds, mostly moist, some- what shining, margin whitish gray, central mass 1.5 cm. high. A study of the authors' figures seems to show a conidiophore suggesting an Acremonium or, in some cases, even a suggestion of Verticillium or Clado- hotryum. The fact of the very long time of incubation and the fact that the colonies occurred near the edge of the plate suggest that the fungus described may be a contaminant. This work needs further confirmation. Malassezia ovalis (Bizzozero) Acton & Panja, Indian Med. Gas. 62: 603- 614, 10 pis., 1927. Pityrosporum Malassez, Arch. Physiol. II, 1: 203-212, PI. 11; 451-464, PI. 20, 1874. Saccharomyces ovalis Bizzozero, Arch. Path. Anat. Physiol. [Virchow] 98: 441-459, PI. 13, 1884. Saccharomyces sphericns Bizzozero, Arch. Path. Anat. Physiol. [Virchow] 98: 441-459, PI. 13, 1884. Saccharomyces capillitii Oudemans & Peckelharing, Nederl. Kniidk. Arch. II, 4: 502-562, 1885. [Contr. Fl. Pays Bas 11.] Microsporuni Malassezi Baillon. Bot. Med. Cr\'ptog, 1889. Flaschenbacillus Unna, Monatsh. Derm. 13: 225, 1891. Microsporum Audouini Vuillemin, Bull. Soc. Myc. France 11 : 94-103, 1895 non aliorum. Pityrosporum Malassezi Sabouraud, 1895. Pityrosporum ovale Castellani & Chalmers, Man. Trop. Med. ed. 2, 836-837, 1913. Cryptococcus ovalis Vuillemin. 360 MEDICAL MYCOLOGY Cryptococcus capillitii Vuillemin. Cryptococcus Malassezii Benedek, Centralbl. Bakt. I, 116: 47-67, 1930. Torulopsis ovalis Almeida, Annaes Fac. Med. Sao Paulo 9: 10, 1933. f Pityrosporum Cantliei Castellani & Chalmers, Man. Trop. Med. ed. 2 837, 1913. Saccharomyces Cantliei Castellani, 1908. Originally described by Malassez from the superficial homy layer and the mouths of the hair follicle in pityriasis sicca of the scalp, afterward reported from seborrhea capillitii. First named by Bizzozero, who also reported it from the heads of normal persons along- with his Saccharomyces sphericus. Later it was confused with spores of various dennatophytes seen in scrapings [for care- ful review of this literature see Kraus 1913]. Unna (1891) claimed that the structures reported by earlier workers were swollen remains of a bacillus, which he referred to as the Flaschenbacillus, and which he described as follows: cells short cylindiic, 1 x 2/a, either swelling to spherical cells, 2-3 times normal, or becoming flask-shaped, rarely pyriform or the shape of a dumb-bell, or even elongating to 2-3 times normal length, 1.5-3 x 2-4/^. Hoorn recognized P. sphaericum as a slowly growing white yeast similar to Pekelliaring's Saccharo- myces capillitii and S. ovale as a large flask-shaped bacillus with swollen forms. He also regularly isolated a small bacillus from seborrhea. Lomery returned to the hypothesis that it was a yeast and not a bacillus, and Tieche found these organisms in only 2 out of 50 eases of seborrhea. Hodara apparently first succeeded in cultivating the organism but, as has been the experience of most subsequent workers, it died on attempted sub- culture. Acton and Panja are apparently the next ones who were successful with this organism. The following accounts of the organism, as well as of pityriasis and seborrhea, are largely taken from their paper. Both the terms "pityriasis" and "seborrhea" have been used in different senses. In the original use, pityriasis was applied by Willans to the delicate pellucid scaling of the epidermis, without obvious signs of inflammation, of which dandruff is the most characteristic. This term has been variously ex- tended by later authors to include scaling of wholly different origin, such as pityriasis rosea, etc. Seborrhea was originally used to designate an excessive flow of sebum, considered to be due to any functional disturbance of the sebaceous glands, without thought of disease. Hebra introduced confusion by teaching that the scales of pityriasis were dried flakes of sebum. When the flow is excessive, the flakes are oily, when normal, the flakes are dry. Since there, are no sebaceous glands on the palmar and plantar surfaces, although they are often oily, Unna concluded that the oil was produced in the sweat glands. The French school, under the leadership of Sabouraud, advocated the theory that seborrhea was a mild local inflammation of microbic origin. Sabouraud recognized the yeast of Malassez and a gray coccus (morococcus) , which Acton and Panja have since shown to be different stages of the same organism. Sabouraud also recognized the acne bacillus as a complication in MALASSEZIA 361 many lesions producing acne (comedones or blackheads). Sabouraud (1932) recognizes pityriasis simplex capitis (dandruff), pityriasis steatoid (seborrheic eczema of Unna) usually a later stage of dandruff, which eventually leads to seborrheic alopecia (baldness), acne vulgaris (comedones or blackheads), papular seborrheas, in connection with acne rosacea and, finally, seborrheic warts of old age. Acton and Panja, using a dry, fatty medium, show that all conditions are due to the same organism, although secondary infections may occur. Two other kinds of scaling are common, pityriasis versicolor caused by Malassezia furfur, and pityriasis flava (tinea flava of Castellani) caused by Malassezia tropica. Probably pityriasis capitis is the most important, as the scalp forms a con- tinual source of scales for the transfer to other portions of the body, and until baldness begins to appear in middle life, it is easily overlooked and seldom treated by a physician. Also these scales usually contain Staphylococcus alhus or S. aureus, thus forming a reser\^oir for these organisms to spread to other parts of the body, producing pimples, boils, etc. When Malassezia grows on delicate skin, as that of the face, flexure of joints, etc., of children, the desquamation leaves the basal cell layer exposed, allow- ing the invasion of the corium by streptococci with the production of seborrheic eczema and eczema rubrum of children. The synonymy may be tabulated as follows : General terms: seborrheic dermatitis, seborrhagia, steatidrosis, seborrhea of Hebra, hyperidrosis oleosa of Unna, flux sebacea of Eayer. Scalp lesions: dandruff, pityriasis capitis, seborrheic dermatitis of the scalp, seborrhea- sicca, seborrhea oleosa, seborrheic alopecia. Glabrous skin: seborrheic dermatitis, pityriasis circinata, pityriasis steatoides, flannel rash, seborrhea corporis, lichen circinatus. Acne: comedo, blackhead, grouped comedo, acne vulgaris, acne disseminata, acne necrotica. Lips: exfoliative cheilitis. With streptococcus as secondary invader: seborrheic eczema and eczema rubra. For the benefit of the botanist who is unfamiliar with the structure and development of the sebaceous glands, I give the following summary from the work of Acton and Panja (1927) and of Sabouraud (1932). The embryonic skin consists of two layers: a basal layer of columnar cells and Rauber's layer, consisting of swollen cells which stain badly, and form a fatty substance which waterproofs the fetus against the maceration of the skin by the liquor amnii. About the fingers and toes, these cells are swollen, hemispheric, polyhedral, and are called "the bladder cells of Zander." After birth, the superficial layer begins keratinization. In the early stages of keratinization (first three years), Malassezia usually infects the scalp. The sebaceous glands, which are scat- tered over the surface of the body in fishes and amphibia, become associated with the hair follicles in the mammals. Some of the glands which are not associated with the follicles have taken on specialized functions. Of those associated with the hair follicle, those in the scalp, beard, etc., are simple or 362 MEDICAL MYCOLOGY bilobed and relatively small, and are the sites of pityriasis, while those asso- ciated with the area of lanugo hair are larger, multilobed, often racemose and are found in the areas which furnish the sites of acne. Those associated with hair follicles seem to function in oiling the hair shafts. Some of those not associated with hair follicles, are associated with the sexual glands. Of these the mammary glands are the most conspicuous, but smaller ones are found in the axilla, scrotum, prepuce, and labia which not only lubricate these regions but also produce volatile fatty oils which impart a distinct odor to the individual or to the species, the odor being especially marked during the mating season. These glands may be infected with various organisms which give rise to inflammation and abscess formation. Later in life, retention cysts may form in them as a result of previous irritation. Other sebaceous glands are concerned with the lubrication of the various orifices of the body, e.g., the ceruminous gland of the ear, the meibomian glands of the eyelids, the glands on the lips, those of the anal margin, etc. These glands are often the seat of staphylococcal inflammation, the organisms being derived from the scales in the scalp, resulting in abscesses of the external auditory meatus, styes, retention cysts, etc. In the palmar and plantar surfaces, their function is assumed by the sweat glands which produce small oil drops at the extreme ends of the coiled glands. Sebum is an oily secretion, becoming cheesy on exposure to air. In chemical nature, it is similar to lanolin (sheep's w^ool fat). Besides the fatty acids, such as stearic and palmitic, there are volatile acids which impart the characteristic odor. Sebum is not a true secretion like milk of the mammary glands, but it is produced by the failure of the superficial layer of cells to undergo keratinization. Instead, a fat vacuolization occurs until the whole layer of cells is converted into sebum. In infections by Malassezia, the super- ficial cells of the scalp undergo this same degeneration, producing the greasy scales of dandrufi^. When Malassezia invades the sebaceous gland, it greatly hastens the production of sebum. Since the sebaceous glands are not provided with a nerve supply, as are the sweat glands, the regulation of flow is dependent on heredity, race, age, sexual activity, customs and habits, diet, temperature, humidity, and perhaps acidosis. Heredity probably determines the number of these glands and their relationship to hair follicles and the function of the endocrine systems in gen- eral, for often greasy skins with tendencies to acne and baldness are common to several members of a single family. Little data regarding race have been ac- cumulated that are not also explicable in connection with their habits and cus- toms. Baldness is rare in women but, whether this is due to endocrine func- tion or to the habit of wearing the hair long, thus preventing infection of the scalp, or the development of the organism by the higher humidity, is un- known. The fashion of wearing short hair is too recent in women to have yielded conclusive evidence on this point. If baldness is linked with the MALASSEZIA 363 gonadal function, one Avould not expect an increase of baldness in women following the adoption of bobbed hair, while if long hair prevents infection, one would expect an increase. When developmental processes in the fetus are at their height, the seba- ceous glands are very active, producing a desquamating layer of fatty cells, the vernix caseosa. A few days after birth the vernix caseosa is shed, and the superficial laj^er begins to keratinize gradually. Here the Malassezia attacks the scalp and the pilosebaceous glands about the face, giving rise to the condi- tion known as miliary sebaceous acne. More rarely it attacks the sebaceous glands of the front or the back of the chest, giving rise to grouped comedones. After the child is three years old, there is little chance of further infection until puberty, as the functions of these glands and of the gonads are in abey- ance. At puberty, both glands again become active, the sebaceous glands, often predisposed to the infection by Malassezia, giving rise to pityriasis and acne. The complexion is muddy, the skin is coarse, greasy and thickened, the mouths of the sebaceous glands are large and those about the nose blacked by comedones. This condition (keratosis) may persist throughout the period of sexual activity, but it usually disappears in early adult life (about 25). Women are not prone to pityriasis but, in those suffering from acnes, there is often a relationship with the menstrual cycle and gonadal activity. In men, at the height of their sexual activity (between 30 and 40 years old), baldness appears on the scalp, and the infection extends to other portions of the body. Later in life we find sebaceous cysts and still later seborrheic warts. Sabou- raud has attempted to correlate baldness with excessive activity shortly after puberty but the difficulty of measuring this activity and of securing reliable data is so great that his results are inconclusive. Customs and habits often play a large part in the spread of the disease. Customs which tend to keep the skin very oily and in low humidity seem to favor its spread, such as anointing the scalp with oil, infrequent bathing, etc. The wearing of hats, often decried, seems to be unimportant ; for example, the bareheaded, short-haired, middle class Bengali is very apt to be bald while the Pathan, with long hair, covered with a hard kulor and surrounded by a turban is practically free from baldness. Excessive amounts of carbohydrates and fats or of alcohol tend to increase the greasiness of the skin and pre- dispose to infection as do warm, dry atmospheres. In infancy, the lesions commence on the scalp as a greasy heaping up of scales, which are often colored dark from accumulated dirt. Thence the in- fection spreads to the face, typically over the flush area of the cheeks. Mild lesions consist of slightly desquamating areas with erythema. In severe cases there is usually secondary infection by streptococci. The lesions of the head and face frequently become infected by a secondary impetigo and the disease spreads rapidly over the scalp, face, behind the ears, neck, and even on the body. In some children the fungus may involve the forehead, producing tiny vesicles caused by blocking the sebaceous glands of this region. It may also 364 MEDICAL MYCOLOGY extend to the flexures of the arms, the popliteal space, and the front of the legs. Since infants under three months are unable to scratch themselves, they relieve the irritation by rolling the back of the head and the sides of the face on the pillow, thus spreading the infection. The sebaceous glands are rarely infected in infancy and when they are, it is probably due to stoppage of their mouths by too liberal application of oils by the mother, with subsequent in- vasion by staphylococci. From puberty until the twenty-fifth year, the organism growing on the scalp produces little or no symptoms beyond slight dandruff. During this period, the invasion of the pilosebaceous glands not connected with hairs is more common. The glands of the nasolabial folds, between the scapulae and those of the front of the chest are most frequently attacked, rarely those of the forehead (acne frontalis). In the mild cases, the skin becomes coarse and blackheads are formed, occasionally these are infected by staphylococci, giv- ing rise to superficial inflammation (with or without pustules and papules). If the staphylococcal infection extends deep into the corium, it gives rise to deep abscesses, the skin over them is bluish and the abscesses are slow in coming to a head. Very rarely the suppuration may end in a localized necrosis some- what similar to the formation of a carbuncle (acne necrotica). Such lesions are seen only in debilitated persons. The scars left by the rupture and ab- sorption of these pustules also vary in different individuals. In superficial lesions, the scars are difficult to see after the inflammation has subsided. In deep lesions, the scars often leave little pitlike areas scattered over the cheeks and sides of the neck. Sometimes these scars undergo keloid formation, pro- ducing ugly raised keloids. More rarely the scar is deeply situated and gives rise to superficial atrophy of the skin overlying it, so that these atrophic scars are white, suggesting the morphea spots. In the last named conditions, the individual shows a lower basal metabolism, suggesting connection with hypo- function of the thyroid [hypophysis or gonad?]. In this period the dandruff scales are small, dry, and greasy, and adhere to the surface of the scalp. The skin is not inflamed and appears normal, but the hair is dry and has lost its luster. From time to time there are exacerba- tions (especially in hot dry climates). The skin becomes irritable, with small erythematous areas and excoriations, due to scratching. Extension occurs on the forehead where the skin becomes red and irritable (corona seborrhoeica). The sebaceous glands become involved, infected with staphylococci, giving rise to small vesicles which, if irritated by friction of the hat, may result in boils. Occasionally this may extend to the eyebrows and eyelashes. The sebaceous glands which are not pilosebaceous are rarely involved by Malassezia, but are infected by staphylococci, producing styes, exfoliative cheilitis of the lips, etc. From 30 to 45 years of age, the Malassezia which has been largely restricted to the superficial layer of the scalp now invades the hair follicle and its seba- ceous gland, after a few years destroying them and producing seborrheic alopecia or baldness. The area may start at the crown and spread or at the MALASSEIZIA 365 forehead and grow backward, until, when baldness is complete, only a narrow fringe is left at the back and sides of the head. At first the number of hairs is not diminished, but they become finer in texture and, as the follicles are destroyed, the space between the hairs increases, until all the follicles are destroyed and the scalp is thin, shiny, and bald. During and after middle age, extensions to the body are more common than in adolescence, when it extends to the forehead as corona seborrhoeica, rarely to the whole face. The skin becomes dry and harsh, thickened and indurated and from irritation of the corium, the melanoblasts and basal layers are stimulated with deepening of pigmentation. When the nape of the neck becomes involved, with the added irritation of collar and other clothing, the skin is thickened, indurated, with deep furrows where the normal lines of skin were (lichenification). More rarely pustules form and the scars become keloid (dermatitis papillaris capil- litii. In elderly people Malassezia may also attack the skin at the flexures of the elbows and knees, producing lichenification or even acute exfoliative der- matitis. Among the poorer classes of England, circinate lesions with a slightly in- flamed base and greasy scales appear on the body, a condition known as flan- nel rash. During middle age the acneiform type of lesions tend to disappear. In acne rosacea, there is a reflex erythema on the nose or the face produced by some irritation in the upper portion of the alimentary canal, causing the skin to become red and thickened and the sebaceous glands to become hypertro- phied. The latter are then invaded by Malassezia, with the production of papular lesions. As a result of the irritation of the mouths of the sebaceous glands these are blocked and the sebum cannot find an outlet; as age advances the glands be- come enlarged from the accumulation of sebum, and retention cysts (wens) are formed. More rarely in adult life the moist skin of the axilla and scrotum is attacked, being covered with a layer of greasy scales, where retention of the secretion of the sebaceous glands and staphylococcal infection occur. Sometimes the glands of the scrotum become blocked, producing a condition similar to Fordyce's disease of the lips, with thickening, induration, and intense irritation, an eczematous condition whose diagnosis is to be carefully distinguished from tinea cruris. In the tissues, Malassezia appears as a yeastiike budding form (the flask- shaped bacillus of Unna or the Morococcus of Sabouraud) . During its growth it interferes with the flattening and the proper keratinization of the super- ficial cells of the scalp, and they are shed as the delicate scales of dandruff. When the disease is spreading rapidly, the scales separate rapidly, and only the yeast cells are seen. At middle age, the mouth of the hair follicle is in- vaded, the homy layer fails to keratinize, and the mouth of the follicle is filled with large swollen cells. The yeast cells divide rapidly at the bottom of the follicle and invade the sebaceous gland, while the leucocytes show an inflam- matory reaction about the neck of the gland. The hair becomes distorted and 366 MEDICAL MYCOLOGY the shaft is invaded by staphylococci. Deeper doAvn, the root is surrounded by an inflammatory exudate and atrophies, the epidermis is thin and atrophied, and the corium consists of a dense white fibrous tissue, causing the glazed, scarred appearance of the skin of a bald head. The site of the follicle is oc- cupied by a thin fibrous scar that has destroyed the hair root, leaving perma- nent baldness. The sebaceous glands not connected with hair follicles are larger and more racemose. As Malassezia ovalis invades the mouth of the gland, it causes the cells of the mouth to disintegrate more rapidly than usual, because at this site there is very little normal transformation of homy cells to sebaceous material. The cells destroyed by the fungus are more resistant and block the mouth of the gland by a fatty plug; the external part is impregnated with dirt and forms the blackhead, filling the enlarged mouth of the sebaceous gland. The flow of sebum is blocked and collects behind the plug, distending the gland (comedo). This sebaceous plug, which on expression comes out as a little coiled "worm," has been called by Sabouraud the seborrheic cocoon. These plugs are completely soluble in ether. Often these plugs are filled with a fine felted mass of the acne bacillus, which has nothing to do with the disease but finds suitable conditions (fat and anaerobiosis) for growth as a secondary invader. As a complication, the comedones may be infected by Staphylococcus alhus or ;8'. aureus, producing the acne pustules. In some individuals, part of the pustules become cicatrized superficially and give rise to keloid scars; in others, the cicatrization is deeper. The gland may become atrophic and its wall reduced to a single row of cells as a result of the excessive sebum fonnation. The inflammation may spread from the gland into the corium, usually without suppuration, giving rise to induration of the corium. The melanoblasts are irritated so that they secrete more pigment, producing a darkening of the skin (chloasma). In the scales of relatively quiescent stages, the fungus cells are large, spherical, 8-12/a in diameter, showing occasionally smaller cells and some bud- ding. In the active stage, the fungus cells are smaller, budding forms are numerous, 2-3/x in diameter in the resting stage. The cell and its adherent bud gave the picture described by Unna as his flask bacillus. When the cells are resting, they give the picture described by Sabouraud as morococcus. In the dandruff, some irregular hyphae are seen, 2-3/x in diameter and 15-20/1, long; often slightly bent. As staphylococci in a given field increase, the number of Malassezia cells decrease. Attempts to cultivate Malassezia on the usual laboratory media have been unsuccessful. Panja obtained his first successful cultures on Petroff's glycerol medium, an Q^g medium colored with 0.0004/^. gentian violet, to inhibit the staphylococci. The scales should be washed in sterile normal saline to free them from as many extraneous foreign organisms as possible. The washed scales are placed on the upper part of Petroff's medium where the slope has dried MALASSEZIA 367 up. The primary colonies appear small, dry, and chalky, visible on the third day. Secondary cultures will grow on the thick part of Petroff's medium, which is still moist, on glucose agar, ordinary agar, and even on glucose broth. Macleod & Dowling- (1928) and Benedek (1930) have also claimed the culture of the bottle bacillus of Unna, but in no case have they established the pathogenicity of the organism cultivated. Benedek describes his organism as follows : Cells mostly spherical, hyaline, 7.8-10.4/x, solitary or rarely in small groups, forming neither hyphae nor spores. On Sabouraud's agar (pH 4.7-4.9) and on 8% glucose agar (pH 6.9-7.1), under strictly aerobic conditions, colony smooth, rounded and brown. He also reports growth on malt extract, glucose, and maltose broth, and acid formation with glucose, fructose, sucrose, and mal- tose solutions and on milk. No fermentation, no liquefaction of gelatin. Ota & Huang (1933) report further details of cultures, using a strain isolated by Acton & Panja, Crypfccoccus graciloides Castellani and a strain isolated by Huang which they consider conspecific. Cells typically ovoid, al- though spherical and bottle-shaped forms may be seen, 2-2. 5/a often united in pairs. Very rarely suggestions of arthrospores were observed, but probably no true arthrospores are produced. On egg medium at the end of several months, senescent cells are 3-5/x, thick-walled, usually solitary spherical, ovoid, or more or less angular, with a large central oil drop, sometimes occurring in short chains with a thin gelified sheath. No mycelium seen. Gram stain colors the protoplasm completely in young cells, less so in adults ; Giemsa stain variable. Petragni's medium found to be the most favorable, colonies becoming 1 mm. in diameter, becoming confluent, especially in moister portions, forming a crustose membrane, granular, moist, ivory white, finally brownish. Ten per cent glucose agar to which a little (about 10%) butter has been added is also favorable, slightly more so than Petroff's medium. On Sabouraud glu- cose, colonies very fine, suggesting those of Neisseria. On malt extract, a slight deposit, but no ring or pellicle. No ascospores observed No fer- mentation. Ota & Huang were unable to prove the pathogenicity of their strains (intravenous and intraperitoneal injections!). Moore (1934) succeeded in isolating Malassezia on wort agar, proving its pathogenicity. Growth is ver,y slow at first but increases after repeated subcul- ture on this medium. He describes his cultures as follows : On Sabouraud agar, colony flat, light ochraceous salmon, dull Avith slight ridges at the margin, cells spherical, 3-lOju mostly 4-5;Ui in diameter. On wort agar, colony pulvinate with radial ridges, surface rough, light ochraceous salmon to pinkish buff, dull cells 3-15/a mostly 4-5/^, bottle-shaped cells com- mon, the larger cells with a thick capsule. On malt extract agar, colony pink- ish cinnamon with a circular flat plateau in the center Avith fine radial ridges. Cells spherical or ovoid, 2-7/x mostly 4-5/a. On Raulin's agar and Richard's agar, growth similar but very poor. On corn meal agar colony ochraceous buff, 368 MEDICAL MYCOLOGY slightly glistening, ridges faint, margin irregular. Long cells occasional. On potato glucose agar colony pulvinate, waxy, pinkish buff, cells spherical to ovoid, mostly bottle-shaped, 2-6/t mostly 4/x. On ManevaFs modification of '°°^^(35a68 °°oo 6 6 88 O0G(3 O0S>Q ooOOObh °^^§ 6 0 6^ SoodS?f 0 n^^S88©aA L5^.S^f!^M u^ 00 % Pig. 70. — Malassezia ovalis. 1, development of ovoid cells ; 2, development of spherical cells ; S, both types of cells on serum agar ; 1,, on Richards' agar ; 5, on Malt extract agar ; 6, on blood agar, bipolar sprouting- ; 7, thick-walled form on yeast-glucose agar ; 8, on Sabouraud agar; 9, on corn meal agar; 10, bottle forms on potato glucose agar; 11, on Maneval modifica- tion of Gorodkova agar ; n, on wort agar ; IS, on glycerol agar ; H, on Raulin's agar ; 15, in lactose broth ; 16, in Sabouraud broth. MALASSEZIA 369 Gorodkova agar, colony similar, light ochraceous salmon, cells similar but some long cells present. On lactose broth agar and nutrient agar, colony flat, or slightly elevated, cinnamon buif, cells spherical, 2-5/a. On lactose broth agar, colony pulvinate, light ochraceous buff, cells 4.5/^, spherical or ovoid. On glycerol agar, colony light ochraceous salmon, smooth, glistening, radial ridges, pulvinate. Cells spherical or ovoid, 2-7/a with long cells. On yeast glucose, colonies similar, cells spherical, 2-5/*, sometimes clinging together in short chains of 3 cells with few long cells. On serum agar, growth poor, colony dull and pasty, ochraceous buff. On blood agar, colony glistening or waxy, pinkish buff, appearance of that on malt extract agar. On peptone broth, no pellicle or ring, sediment shows cells spherical to ovoid, 3-9/* mostly 5fi in diameter, forming chains 3-5 x 12-15/i with many thick-walled cells. On lactose broth, similar, but spherical cells up to 12/x. in diameter with long cells suggesting catenulate conidia, ovoid cells about 6/t in long axis, larger cells thick-walled (Fig. 70). Therapeusis. — As with other fungus infections, there is no simple treat- ment and the standard texts should be consulted. Since the dandruff scales of the scalp form a reservoir of both Malassezia ovalis and staphylococcus, any treatment should be aimed at their complete removal to prevent reinfection. When this has been accomplished, the mouths of the sebaceous glands may be softened by an appropriate lotion and the comedones expressed. Staphylo- coccus infections may be cleared up by vaccines, although vaccines alone with- out an attempt to get rid of the dandruff scales as a reservoir of reinfection are of very uncertain value. Malassezia Macfadyemi Castellani, 1908. Microsporon Macfadyeni Castellani, Brit. Med. Jour. 1905, 2: 1272, 1 fig., 1905; Trans. Internat. Derm. Congr. 6: 661, 1908; Jour. Trop. Med. Hyg. 10: 65-66, 1907. Trichophyton Macfadyeni Castellani, Jour. Cut. Dis. 26: 396-397, 1908. Trichophyton Macfadyeni Castellani, Jour. Trop. Med. Hyg. 11: 261, 1908. Atrichophyton Macfadyeni Castellani & Chalmers, Man. Trop. Med. ed. 3, 1009, 1919. Found present in cases of pityriasis alba. Hyphae slender, short threads of regular outline; spores small 3-3.5/x in diameter, ovoid, sometimes appearing in large clusters. Cultivated twice for short period on Sabouraud maltose agar, gave colo- nies of very slow growth, deeply pitted yellowish mass, very firmly and deeply rooted in medium. Transfers always failed. Malassezia tropica (Castellani) Schmitter, Jour. Trop. Med. Hyg. 26: 194, 1923. Microsponmi tropicum Castellani, Brit. Med. Jour. 1905, 2: 1271-1272, 1905; Trans. Internat. Derm. Congr. 6: 661, PL 44, 1908; Jour. Trop. Med. Hyg. 10: 65-66, 1907; Jour. Cut. Dis. 26: 396, 1908. Appears in tinea flava or pityriasis flava. Not cultivated by Castellani. 370 MEDICAL MYCOLOGY Spores spherical or ovoid, 3.5-4.5/*, thick-walled. No white spots or craters on surface of glycerol agar cultures. Indol not produced. Litmus in broth strongly decolorized. No acid formation in lac- tose broth. — Schmitter. Malassezia pachydermatis (Weidman) Dodge, n. comb. Pityrosporum imchydermatis Weidman in Fox, Kept. Lab. Mus. Comp. Path. Zool. Soc. Philadelphia 36, 1925. Pityrosporum rhinoserosum Sabouraud in Lodder, Anaskosporogenen Hefen 1 : 189, 1934. Isolated from scaling of the skin of Rhinoceros unicornis in the Zoological Garden in Philadelphia. In scales, cells small, sprouting, polar. In malt extract, cells ovoid or flask-shaped, 1.5-3 x 2.5-5/x, sprouting over a broad base, polar. Growth very slow, elevated, center pointed, dark yellowish, thick, surface smooth, regular, dull. On malt agar, colony brown, margin lighter, dull, some- what irregular surface, margin smooth. On malt gelatin, colony gray, dull, smooth, margin irregular. On malt extract, a few islets, a thin ring, and granular sediment. On alcohol, a few thin islets. No fermentation; gelatin liquefied after 12 weeks. Doubtful Species Mycoderma Dombrayi Vuillemin, C. R. Acad. 8ci. 189: 405-406, 1929. Isolated from white pityriasis of the skin of penis and scrotum of a der- matologist. The scales show filaments varying from 3-6.75/a in diameter with chlamy- dospores ; same appearance on liquid media. Chlamydospores reach a diam- eter of 14/x on germination. BIBLIOGRAPHY The following articles relating to Hemispora infections were overlooked when this genus was transferred from the Toriileae at a late stage in the prepa- ration of this work : Falchi 1925, see p. 689 ; Fonseca & Area Leao 1927, see p. 689 ; Porcelli 1922, see p. 692 ; Sartory, Meyer & Meyer 1930, see p. 692, and Vuillemin 1906, see p. 693. Aars, C. G. 1930. Piedra, Arch. Derm. SyphiJol. 22: 401-409, 9 figs. 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A variation of gemmation of Blastomyces dermatitidis in the tissue lesion, Jo^ir. Infect. Dis. 18: 618-629, PI. 22,23. Wade, H. Windsor & George S. Bel. 1916. A critical consideration of systemic blastomycosis with notes on certain special features and report of five cases. Arch. Int. Med. 18: 103-130. Wallace, G. I. & Fred W. Tanner. 1927. An etiologic agent in bronchomycosis, Amer. Bev. Tuberculosis 15: 373-379. Watanabe, K. 1927-1928. Die chromolytische Studie der Hefe. I. Der chemische Aufbau der Hefe, Jap. Jour. Derm. Urol. 27: 373-385. [27-33] 1927. II. t)ber freie Fette und Fettarten, Ibid. 27: 735-747. [57-59] III. tJber die Hefealbumose und Grobuline, Ibid. 27: 945-959. [68-72] IV. Die Zuckerarten der Hefe, Ibid. 28: 521-532 [29-33], 1928. *Watanabe, N. 1919. On blastomycetic meningitis. Mitt. Med. FaTc. Viviv. Kyushu, Fukiioka 5: 1-16. Watts, James W. 1932. Torula infection, a review and report of two cases, Amer. Jour. Path. 8: 167-191, Pis. 29-33. Weidman, Fred D. 1922. Resemblance of yeasts in cutaneous scrapings to Hyphomycetes, Arch. Derm. Syphilol. 5: 325-328. — . 1930. The effects of gentian violet and crystal violet on certain pathogenic yeasts, Urol. Cutaneous Bev. 34: 215-225. — . 1933. Cutaneous torulosis, the identification of yeast cells in general in histologic sec- tion. Southern Med. Jour. 26: 851-863. Weis, Joseph D. 1902. Four pathogenic Torulae, Jour. Med. Bes. 7: 280-311, Pis. 21-24. [n. s. 2.] Weiss, Charles. 1928. A review of the recent literature on tropical sprue. Arch. Path. Lab. Med. 6: 885-900. MALASSEZIA 423 Weiss, Charles & Francis Laudron. 1928-1929. Immunological investigations of tropical sprue in Porto Rico. I. Monilicidal activity of the blood in sprue, Amer. Jour. Irop. Med. 9: 83-86, 1929. II. Skin reactions with toxins of Monilia, Ibid. 9: 87-91, 1929. III. Attempts at transmission of sprue to monkeys and human volunteers, Hid. 9: 92-95, 1929. IV. Biology of Monilia psilosis in relation to sprue, Jour. Infect. Dis. 43: 557-564, 1928. Weller, C. V. & A. D. Riker. 1930. Rhinosporidium Seeberi; pathological histology and re- port of a third case from the United States, Amer. Jour. Path. 6: 721-732. Wells, H. Gideon. 1898. Preliminary report of a case of blastomycetic dermatitis. New York Jour. Med. 67: 427-430. Wernicke, Robert. 1892. Ueber einen Protozoenbefund bei Mycosis fungoides? Centralbl. BaU. 12: 859-861, PI. 6. Wettstein, Richard von. 1885, Untersuchungen iiber einen neuen pfianzlichen parasiten des menschlichen Korpers, Sitcungsher. K. Alad. TViss. U'ien. 1, 91: 33-58. White, Charles J. 1927. Fungus diseases of the skin, clinical aspects and treatment, Arch. Derm. Syphilol. 15: 387-404. A\niite, Charles J. & J. H. Swartz. 1928. Cryptococcosis epidermica, Arch. Derm. Syphilol. 18: 692-705, 5 figs. White, Cleveland. 1927. Studies in mycotic dermatitis. I. Superficial yeast iiLfections of the glabrous skin, New York State Jour. Med. 27: 1116-1120 [19]. [Same title, Arch. Derm. Syphilol. 18: 429-438, 1928.] Whitfield, Arthur. 1908. Microscopical specimen and pure culture of a yeast derived from a case of intertriginous dermatitis of the cruroscrotal region, Proc. B. Soc. Med. 1: Derm. Sect.: 190. [Reprinted, Brit. Jour. Derm. Syphilis 20: 274, 1908.] — . 1909. A microscopical specimen of a mould fungus obtained in a scraping from a case of chronic dermatitis of the glans penis, Proc. E. Soc. Med. 2: Derm. Sect.: 90. Wliitnian, Ross C. 1913. A contribution to the botany of the organism of blastomycosis. Jour. Infect. Dis. 13: 85-94. Wieben, Magdalene. 1927. Die Infektion, die Myzel iiberwinterung und die Kopulation bei Exoascaceen, Forsch. Gehiete Pflansenkranlch. Immunitdt im Pflanzenreich 3: 139-176. Wilhemi, Armand. 1899. Beitrage zur Kenntnis des Saccharomyces guttulatus (Buscalioni), Centralbl. Bakt. II, 4: 305-309, 353-361, 412-417, 713. Wilhelmj, C. M. 1925. Primary meningeal form of systemic blastomycosis, Amer. Jour. Med. Sci. 169: 712-721. Will, H. 1903-1916. Beitrage zur Kenntniss der Sprosspilze ohne Sporenbildung welche in Brauereibetreibung und in deren Umgebung vorkommen. I, Centralbl. Bakt. II, 10: 689-700, 1903; II, Ibid. 17: 1-3, 1907. HI, Ibid. 17: 3-7, 75-90, 137-146, 331-334, 428-445, 604-614, 693-712, 3 pis., 1907. IV, (nach Untersuchungen von J. Dachs), Ibid. 21: 386-392, 459-469, 1908. V, (nach Untersuchungen von J. Schenckenbach), Ibid. 34: 1-35, 1912. VI, Ibid. 46: 226-280, 1 pi, 1916. — . 1910. Beitrage zur Kenntnis der Gattung Mycoderma nach Untersuchungen von Hans Leberle, Centralbl. Bakt. II, 28: 1-37, 15 figs. — . 1915. Vergleichende morphologische und physiologische Untersuchungen an vier Kul- turen der Gattung Pseudosaccharomyces Klocker (Saccharomyces apiculatus Reess), Centralbl. Bakt. II, 44: 225-290, 1 pi, 6 figs. Williams, J. R. 1922. Systemic blastomycosis, Med. Jour. Australia 2: 186, 187, 4 figs. Winkel, Ludwig. 1920. Ueber die Lymphangitis epizootica des Pferdes mit besonderer Beriicksichtigung der Therapie, Monatsh. Prakt. Tierheilk. 30: 385-433, 1 fig. *Winkler, Jos. 1898. Lungenentziindung bei Rindern, Woch. Thierheilk. 111. Wlaeff. 1900. Levures pures dans un sarcome de 1 'uterus chez une fomme, C. E. Soc. Biol. 52: 759, 760. Wold, Michael G. 1923. Fungous diseases of man in the state of Nebraska. Sporotrichosia blastomycosis, actinomycosis, Trans. Sect. Path. Physiol. Am. Med. Ass. 74: 26-45, 10 figs. [Condensed, Jour. Amer. Med. Ass. 81: 647-643, 10 figs.] 424 MEDICAL MYCOLOGY Wolbach, Simeon Burt. 1904. The life cycle of the organism of dermatitis coccidioides, Jow. Med. Res. 13: 53-60, Pis. 1-3. — . 1915. Eecovery from coccidioidal granuloma, Boston Med. Surg. Jour. 172: 9-4-96. Wolfram, St. & Franz Zach. 1933. tJber einige durch niedere Pilze verursachte Nagerler krankungen beim Menschen, Arch. Derm. Syphilis 169: 95-104, 5 figs. Wood, Edward Jenner. 1915. The occurrence of sprue in the United States, Amer. Jaur. Med. Sci. 150: 692-699. — . 1919. Eecognition of tropical sprue in the United States, Jour. Amer. Med. Ass. 73: 165-168. — . 1925. Pernicious anemia in its relationship to sprue, Amer. Jour. Med. Sci. 169: 28-39. Wortis, S. Bernard & Henry B. Wightman. 1928. Case report of Torula meningitis, Biill. N. Y. Acad. Med. 4: 531. Wright, Jonathan. 1907. A nasal sporozoon (Rhinosporidium Kinealyi), N. ¥. Med. Jour. 86: 1149-1152. Wright, R. E. 1922. Rhinosporidium Kinealyi of the conjunctiva, Indian Med. Gaz. 57: 6, 7, 82, 83, S figs. — . 1922. Rhinosporidium Kinealyi of the conjunctiva cured by tartrated antimony (tartar emetic) and notes on a case in vrhich the lacrymal sac wis affected by this sporozoon, Indian Med. Gaz. bl: 81, 82, S figs. Wright, R. E. & T. S. Tirumurti [Trimurthi]. 1922. Rhinosporidium Kinealyi, Indi< Ota & Lanp-eron, Ann. Para- sitol. Hum. Comp. 1 : 329, 1923. Closteroaleurosporia Qirivckeana Grigorakis, Ann. Sci. Nat. Bot. X, 7: 412, 1925. Microsporum (Closteroaleurosporia) Quinckeanum, Guiart & Grigorakis, Lyon Med. 141: 377, 1928. This species presents many problems. It is very difficult in the early eases to be sure whether the authors were dealing with accidental inocula- tions of mice by A. Schoenleini or by this organism. Even the descriptions of the early cultures leave us in doubt, but since the lesions on mice are more severe and more common, reaching epizootic proportions, when caused by this species, it seems quite probable that the early workers, such as Bennett (1842), Draper (1854), Friedrich (1857), Gluge & d'Ukedem (1857), Pieschel & Voigtlander (1857), Zander (1858), and Schrader (1858) were dealing with this species. The statements regarding morphology are extremely contradic- tory. Quincke, Busquet, Sabouraud, and Grigorakis figure closterospores, Bodin does not mention them, while Langeron & Milochevitch deny that clos- terospores are produced on any medium. Producing severe favie lesions on mice, usually attacking the head and largely destroying the skin and the cornea, causing death by starvation, fol- lowing blindness. Found also on a hedgehog (herisson) which had been em- ployed to rid an infested house of mice. Easily inoculable into man, in whom the lesions are usually confined to the glabrous skin, usually with few favie crusts, so that the human lesion might be diagnosed as herpes circinatus. Micro- scopically indistinguishable from Achorion Schoenleini in the tissues. Inocu- lable into guinea pig, producing either favie scutula or dry scaly lesions which heal spontaneously. Common in Germany, occasional in England, Austria, and Hungary, rare in France. In hair, hyphae 1.5-2/x in diameter, arthrospores 3-6/x in diameter. In cul- tures closterospores septate, walls thick, 40-70/*, 4-7-celled (3-6 septate) ; chlamy- dospores 7-10/*, rarely up to 15/i ; aleurospores 4-5/1, borne on compound thyrses. Langeron & Milochevitch report spirals on straw and dung. Colony a pure white disc resembling pleomorphic colonies even in the pri- mary colony, with a short marginal fringe, becoming rough, with more or less concentric furrows, with some irregular hills and channels around the margin; reverse yellowish white at room temperature, deep violet at 35° C. On malt agar (3% maltose and 3% carbohydrates, expressed as glucose), colony more 1 TRICHOPHYTONEAE 557 sharply folded, approachiny cerebriiorin. Ou glycerol peptone agar, inter- mediate between that on malt agar and on Sabonraud glucose agar. On beef broth, forming white islets, light yellow below with some grayish fiocci on the liquid. Gelatin liquefied; milk casein clotted then liquefied, colony white, floating. On potato, a fine short, white velvet, with short irregular folds and furrow; growth mediocre. The resemblance of primary cultures to pleomorphic cultures of A. Schoen- leini at fii-st led Sabonraud to think that they might be the same species, with the characters observed due to host reaction, as had been elaborately suggested by Busquet (1890, 1891, 1892), without experimental evidence, but this hypoth- esis was abandoned by Sabonraud before his classic work in 1910. Grigorakis (1925) has recently revived and extended this idea to include all the derma- tophytes. Bodin (1902) after a study of the nutrition of this organism claimed that it was more closely related to the species of Microsponim and Trichophyton than to Achorion Schoenlemi. Ota & Langeron (1923) also placed it in their genus SahoiD-audites along with Microsponim and Ectotrichophyton. Achorion cupressiforme Aoki, Festschr. Keizo Dohi, 517-574, Pis. 40-42, 1917. Achorion sp. Aoki, Derm. Woch. 59: 863-872, 1914. Producing favus turriformis and favus confertus on glabrous skin with lesions also on scalp. In one case there was swelling of lymph nodes with cheesy pus formation. Easily inoculable to mouse and rabbit, producing whitish scutula and healing spontaneously. Mycelium septate with swollen cells becoming chlamydospores, a few closterospores present; arthrospores in scutula 3-8 x 2.5-5/^,. Growth rapid in medium, slow on surface, rapid in depths suggesting cypress leaves, white becoming dirty white and finally yellowish white, sur- face smooth and waxy. Center elevated, surrounded by a flat waxy zone with periphery below surface of the medium. Reverse bright yellow. On broth, floating floccose white colony becoming yellowish white, chlamydospores abun- dant. On milk, growth slow, coagulated and digested after several weeks. On lactose agar, center elevated, irregular then crimped, brownish yellow. On potato, irregular brown to black brown, elevation surrounded by a white flat surface, growth slow. On beef broth peptone agar (slightly alkaline), colony as on Sabonraud, but moist shining and waxy, elevated in the center, sur- rounded by a yellow white zone, marginal portion radiating, submersed. Gelatin slowly softened (partial liquefaction), surface white then yelloAvish white, reverse yellow. Achorion f ormoseum Hasegawa, 1927 ; Bruhns & Alexander, in Jadassohn, Handbuch d. Haut- u. Geschlectskrank. 11 : 233, 1928. Lesions size of lentil, grayish white with favic scutula. Infected hair lustreless and easily epilated. Fonnosa. Inoculation of guinea pig easy, with typical scutula ; not inoculable to mouse, ape, or hen. Spores 3.5-4/x in the scutulum. along with septate, wavy mycelium; hair showing spores, but not the air bubbles so characteristic of A. Schoenleini. 558 MEDICAL MYCOLOGY In culture, mycelium septate, wavy, 3-4/a in diameter, spore chains, chlamy- dospores, favic candelabra and nodular organs present. On glucose agar, bright yellow and moist, then brownish. Surface of ir- regular folds; in the medium radial branching fibrils. Achorion Schoenleini (Lebert) Remak, Diagnostische und pathogenetische Untersuchungen 193, Figs. 5, 6, 1845.* Oidium Schoenlemi Lebert, Physiologic pathologique 2: 490, 1845.1 Oidium sp. Miiller & Retzius, Arch. Anat. Physiol. AViss. Med. 1842: 198- 212, 1842. Oidium porrigi7iis, Montague apud Berk. & Broome, Ann. Mag. Nat. Hist. II, 7: 177, 1851. Favuspilz /3, Quincke, Arch. Exp. Path. Pharm. 22: 62, 1886. Oospora porriginis, Saccardo, Sylloge Fungorum 4: 15, 1886. Oidium Schoenleinii Zopf , Die Pilze 481, 1890. Schoenleinium achorion Johan-Olsen [Sopp], Centralbl. Bakt. II, 3: 276- 284, 1897. Gruhyella Schoenleini Ota & Langeron, Ann. Parasitol. Hum. Comp. 1: 330, 1923. Arthrosporia Schoenleinii Grigorakis, Ann. Sci. Nat. Bot. X, 7: 414, 1925. Achorion (Gruhyella) Schoenleini Guiart & Grigorakis, Lyon Med. 141: 377, 1928. Probably also the folloAving names are synonyms : Achorion kerotophagus Ercolani, Mem. Accad. Sci. 1st. Bologna III, 6: 363- 381, 1 pi., 1875; Archivio med. vet. 1: 5-21, 1876. Oospora porriginis var. ceratophagus Saccardo, Sylloge Fungorum 4: 15, 1886. Achorion atacton Unna. Achorion dichroon Unna. Achorion radians Neebe & Unna, Monatsh. Prakt. Derm. 16: 17-31, 57-72, 1893. Achorion akromegalicum Neebe & Unna, Monatsh. Prakt. Derm. 16: 17-31, 57-72, 1893. Achorion demergens Neebe & Unna, Monatsh. Prakt. Derm. 16: 17-31, 57- 72, 1893. Achorion cysticum Neebe & Unna, Monatsh. Prakt. Derm. 16: 17-31, 57-72, 1893. Achorion moniliforme Neebe & Unna, Monatsh. Prakt. Derm. 16: 17-31, 57-72, 1893. Achorion tarsiferon Neebe & Unna, Monatsh. Prakt. Derm. 16: 17-31. 57-72, 1893. Producing scutula and kerions in man, inoculable to guinea pig, rat, mouse, cat, and rabbit, but very rarely occurring spontaneously on these ani- *This work appeared in the latter part of 1845 according- to Hinrichs, J. C. Verzeichnisse der Bticher Landkarten u.s.w. welche von Juli bis Dezember 1845 neu erscheinen slnd. 194, 1845. tThis work appeared not later than July 25, 1845, according to Bibliographie de France 399, 1845. No. 3818 for July 26, 1845. TRICHOPHYTONEAE 559 mals. In 1890, this species was reported common in Scotland, Italy, Spain, Austria, Poland, Volga and Caucasus governments of Russia, China, Central Asia, and Abyssinia, occasional in Holland, Scandinavia, and France, rare in England, Switzerland, America, and Japan. At present it is reported com- mon in Southern Japan, Algeria, Holland, Rhine Valley, and Eastern Germany, especially in Schlesien, Bosnia, Hungary, Poland, and the Russian border states, Transcaucasia, Bessarabia, Italy, and Scotland. Occasional in France, rare in Formosa, Northern Japan, Manchuria, United States, Portugal, Argen- tina, and Sao Paulo in Brazil. Ordinarily only arthrospores present, but aleurospores abundant on grains of barley and carrot, much less on potato and wheat, none o(n other media so far reported. Aleurospores 6.5-7 x 5-5. 5/x. Cultures on Sabouraud agar yellowish white similar to fresh beeswax; colony cerebriform, heaped, not spreading. Form constant on the usual media, varying in the fineness of convolutions, size, and rapidity of development. Suc- cessive subcultures grow more rapidlj^, colonies never velvety until pleomor- phism sets in when they become cottony, white. Before pleomorphism, best growth on high nitrogen and low carbohydrate media, while after pleomor- phism, the proportions are reversed. Pleomorphic cultures of A. Schoenleini are suggestive of primary cultures of A. muris. On potato and carrot, colonies dirty white, elevated, irregular, not velvety, not producing pigment on media. On gelatin and coagulated serum small irregular colony, whitish, not velvety; liquefaction begins in about a week but is very slow. On milk, casein com- pletely digested in 3 weeks. At 37° C. growth rapid, colony disciform. On wheat, colonies small, irregular, grayish yellow masses with wrinkled surface. After 7 months they become powdery, white. Grigorakis (1933) has renamed a pleomorphic strain of this species Arth- rosporia Gougeroti. Chen. Kurotchkin and Hu (1931) described a buff variety common in Peiping, China. Colony 2.5-5 mm. in 7 days, small round, smooth surface, slightly powdery and buff. Submerged growth profuse 3-4 cm. in 10 days, composed of deep radial rays. After 2 weeks, secondary growths buff and glabrous or white with short velvet. Morphology and lesions of A. Schoenleini. Guinea pig inoculation resulted in about 50% infection. When scutula were used, only superficial scaly lesions developed, while inoculation from cultures produced small typical scutula. Var. mongolica (Hashimoto & Ota) Dodge, n. comb. GruhyeUa Schoe^ileini var. mongolica Hashimoto & Ota, Jap. Jour. Derm. Urol. 27: 386-409 [33-35], 1927. Producing favus in Mongolia, mouse very susceptible, with a large deep scutulum. On guinea pig and rabbit slight lesion, spontaneously healing. On glucose, maltose, or conservation agar, mycelium straight, frequently branched, rarely undulate, septa distant. Intercalary chlamydospores, pecti- nate organs, fa vie candelabra, nodular organs (?) (resembling "cocon d'uue chrysalide entre des brindelles"). 560 MEDICAL MYCOLOGY Disc 4 cm., chestnut maroon with irregular radial furrows, margin fringed ; daughter colonies may appear on mother colony or at its margin, elevated white chalky or yellowish brown. Subcultures on conservation agar increasingly elevated and folded, becoming yellowish gray. Subcultures on Sabouraud sugar media begin to resemble T. flavum or T. plicatile, 5-8 cm. in diameter, crumpled, center brown, cracks in gray powdery surface, margin of fine rays. Achorion passerinum Fischer, Derm. Woch. 87: 1359-1361, 2 figs., 1928. Producing large scutula on canary, a herpetic lesion on man, not patho- genic for white mouse. Only intercalary or, rarely, terminal chlamj^dospores present ; clostero- spores and aleurospores absent. Colony very slow growth, center with a thin white velvet, with radial folds, fracturing in the center. Probably close to Achorion gallinae but white. Achorion niveum Greco, Origine des Tumeurs 67-83, Figs. 15-28, 1916. Producing small nodule like a lentil on the " cou-de pied," covered with a crust, followed two months later by swelling of leg and redness. Followed by similar nodules on other parts of the body. Pathology and case history de scribed in detail. Filaments flexuous, l-2yu in diameter, granulous, intercalary chlamydo- spores 4-6/A, round or fusiform, occasionally up to 8-lOja, sometimes terminal. Endoconidia sometimes formed. Growth does not show for about 15 days on ordinary agars. On simple agar, small cerebriform elevations resembling A. Schoenleini. On Sabouraud conservation agar, potato, and carrot, elevations dull gray, with a fine pure white nap of fascicles of hyphae. Similar structures below the surface brownish gray. Colony finally crateriform, on Sabouraud agar. Achorion caninum (Costantin & Sabrazes) Dodge, n. comb. Oospora canina Costantin & Sabrazes, Arch. Med. Exp. 5: 354-358, 1893. Bodinia canina Brumpt, Precis Parasitol. ed. 4, 1297, 1298. 1927. Producing favus in dog, inoculable to man, mouse, and dog. Mycelium septate without favic candelabra; chlamydospores similar to those of A. Schoerdeini, rare; in the scutula, long chains of ovoid arthrospores 5-6/a and fascicles of hyphae staining less deeply. Colony with short, velvet, thick pure white, here and there powdeiy, red pigment diffusing as in A. gallinae but darker. On milk, colony bright red. On malt, pellicle adherent to walls of tube, wavy, snow white, reverse dark red and medium becomes deep red. On potato, brown acuminate peaks, surrounded by an extensive white velvet. Only chlamydospores and arthrospores known, the former rare. Cytology not studied and its positions somewhat uncertain. In lesions agreeing so closely that it seems to belong here. Achorion africana Dodge, n. sp. Achorion sp. Mitchell & Robertson, South African Med. Record 1915 ; Med. Jour. S. Africa 20 : 122-125, 4 fi,gs., 1924. TRICHOPHYTONEAE 561 Producing witkop or dikwakwadi in syphilitic natives in Bechuanaland ; for description of lesions see p. 443. Inoculable to mice where lesion causes pru- ritus, minute papules, the exudate causes several hairs to adhere and after a time a grayish white powderj^ dust occurs about the edge of the lesion. The scratching caused by the pruritus prevents the typical white scutulum from developing. The fungus was reisolated from the experimental lesions. Arthrospores isodiametric, cylindric, invading hair follicle, while medium is moist and colony young, suggesting colliform colonies etched with fine wavy lines. After 10-12 days fine snow white filaments, coarse and white as if dusted with dry lime. Slight growth downward into the medium, making the colony very adherent to the medium. Mitchell & Robertson argue that it is not syphilitic in origin since arseni- cal and mercuric antisj^philitics have no effect, while it clears up in 2-3 weeks on treatment with dilute nitrate of mercury ointment, although 2-3 months are necessary to eradicate it completely. The incidence of the disease is from in- fancy to late puberty. It tends to disappear or become less after puberty. Occasionally new hair grows Avhere the crust or scutulum is removed, but more often scar formation results in alopecia, in which case the condition is called ''kaalkop." One boy showed no evidence of present syphilis and gave a negative Wassei-mann reaction. None of the 4 cases intensively studied showed any other lesions suggestive of syphilis. Achorion annulosum Cazalbou, Rev. Path. Comp. 14: 131-144, 3 figs., 1914. Bodinia annulosa Ota & Langeron, Ann. Parasitol. Hum. Comp. 1: 329, 1923. Colony brown gray greenish, glabrous with concentric rings, mycelium hyaline with refringent granules, spores 3-6/a. Grubyella ang-olensis Brumpt, Precis Parasitol. ed. 4, 1304, 1927 ; Froilano de ]\rello & Paes, Rev. Med. Angola, 505-513, 1923 ; Cong. Med. Trop. Afrique Occidentale, Loanda, 1923. Lesions similar to Microsporum in many details, crusts lighter and less ag- glomerated than in Trichophyton: on horee, France. Not inoculable into guinea pig. Hair grayish. Invading mycelium 2-4/a(-5/a) in diameter, slightly branched, straight or slightly sinuous, rarely septate. Spores variable in size in spore sheath, up to 7/x, sometimes irregular. Fringe of Adamson in root of hair. Chlamydospores up to 20/a, hyphae 3-4yLi, of variable diameter. Aleurospores present. Colony in 20 days at 25° C. is 4 cm. in diameter, little elevated above me- dium, surface of dark brown, smooth, concentric zones between fine, velvety, greenish gray zones of equal width, center slight velvety elevation, periphery of fine radiations, outer half of slight radial folds. The velvet gradually dis- appears and by the thirty-fifth day colony is glabrous, moist, finely verrucose, at first deep brown, then color of gingerbread. Immersed margin deep orange; reverse concolorous. On Sabouraud glucose, less growth, glabrous zones deep orange, velvety zones grayish, the glabrous zones giving way to 562 MEDICAL MYCOLOGY short velvet of the same grayish shade. On peptone glucose broth, large, im- mersed yellowish gray plaques, soon filling the space occupied by the medium. Odor of urine. BIBLIOGRAPHY Abramowitz, E. WiUiam. 1931. Local medication in diseases of the skin. Indications, ac- tions and uses of the various active ingredients and vehicles, Arch. Derm. Syphilol. 23: 644-680. Acton, Hugh W. & C. McGuire. 1927. Tinea cruris: its manifestations, diagnosis and treatment, Indian Med. Craz. 62: 419-428, 8 pis. Agostini, Angela. 1930. Una nuova specie di "Bodinia" causa di tigna umana nell 'Eritrea, Atti 1st. Bot. E. Univ. Pavia. IV, 2: 117-125. *Akagi, Shozo. 1926. A new dermatosis caused by a fungus, Jap. Jour. Derm. Urol. 26: 55. * — . 1927. Parasitic skin disease (Unusual previously undescribed skin lesion), Jap. Jour. Derm. Urol. 27: 9. Alexander, Arthur. 1912. Beitrage zur Kenntnis des Eczema marginatum, Arch. Derm. SyphiHs. 113: 11-38, Tl. 1. — . 1922. Statistik der Pilzflora. 1919-1920. (a) Trichophyton persicolor; (b) Interdigital Soormycose; (c) Lychen chronicus Vidal, Arch. Derm. SypMlis 138: 410-415. — . 1922. Die Trichophytie der Hande und Fusse, Med. Klin. 18: 2: 1550-1553. — . 1925. Zur Klinik seltener Soorformen. (a) Dyshidrosis sicca lamellosa oidiomycetica. (b) Paronychie mit sekundarem Soor der Nagel, Derm. Zeitschr. 53: 5-12. — . 1927. tJber die durch den Kaufmann-Wolfschen Pilz hervorgerufenden Hauterkrankungen der Hande und Fiisse mit besonderer BeriicksichtigTing der Dyshidrosis sicca lameUosa, Med. Klin. 23: 275-278. Alfonso, Jose. 1929. Estudio de los hongos en algunas enfermedades parasitarias Bol. Sac. Cubana Derm. Sifilogr. 1: 217-224. Allison, J. Richard. 1927. Fungus disease of the skin. Southern Med. Jowr. 20: 604-606. Amberg, Samuel. 1910. The cutaneous trichophytin reaction, Joihr. Exp. Med. 12: 435-459, PI. 39. Ambrosoli, Gian Angelo. 1921. Coltura di Trichophyton gypseum dal sangue circolante in tricofizia profonda con lichen trichophyticus, Giom. Ital. Mai. Yen. Pelle. 62: 233-240, 1 pi. — . 1923. Favide, reazione cutanea generalizzata alia tigna favosa, Giom. Ital. Mai. Yen. Pelle. 64: 147-157, 1 pi. — . 1925. Lichen trichophyticus in tre fratelli affetti da kerion Celsi (da Trichophyton rosaceum), Giom. Ital. Derm. Sifilol. 66: 628-640, 1 pi. * — . 1926-1927. Coltura di Achorion Schoenleinii dal sangue circolante in un caso di tigna favosa, Giom. Ital. Derm. Sifilol. 67: 1552, 1926; 68: 820, 1927. [Same title, PoUclinico, Ses. Prat. 34: 487, 488, 1927.] — . 1927. Epidemia di tigna da Trichophyton glabrum osservata in Italia, Giom. Ital. Derm. Sifilol. 68: 1011-1020. Americo da Veiga. 1929. Algumas especies novas de cogumelos causadores de tinhas, Brasil Med. 43: 830-838, 4 figs. AndrevfS, George C. 1927. Tinea capitis in an adult, Aroh. Dei'm. Syphilol. 16: 477. Aoki, T. 1910. "Ober die Mikrosporie besonders auf der unbehaarten Haut (in Japan Ha- take genannt), Monatsh. Prakt. Derm. 50: 390-396. — . 1912. Methode der Ausstrichkulturen beim Studium von Epidermispilzen, Jap. Zeitschr. Derm. Urol. 12: 838-872, 11 figs. [66-68]. — . 1914. tiber den Favus der unbehaarten Haut in Japan mit besonderer Beriicksichtigung der bakteriologischen Untersuchung, Derm. Woch. 59: 863-872. — . 1917. Untersuchungen iiber den Favus in Japan, Festschrift, Keiso Dohi 517-574, Pis. 40-42. [Erganzungsband zur Jap. Zeitschr. Derm. Urol.l TRICHOPHYTONEAE 563 Arijewitsch, A. 1930. Zur Frage der Pilzerkrankungen der Augenlider und Wimpern, Derm. Woch. 90: 683-685, 1 iig. Arnold, Walter. 1921. Die intradermale Trichophytinreaktion beim Kind, Arch. Derm. Syphilis. 136: 125-134. Aroeira Neves. 1923. ContribuiQao ao estudo das dermatomycoses em Bello Horizonte. Observa§6es sobre casos piovocados pelo Microsporon felineum C. Fox e F. Blaxal 1896, Brasil Med. 36: 1: 144-149. * — . 1923. ContribuiQao ao estudo das dermatomycoses em Bello Horizonte. Trichopliycia do couro cabelludo pelo Tricophyton cerebriforme Sabouraud, 1893 (Tricophyton flavum Bodin 1902), Arch. Mineiros. de Dermat. SypMUffr. 5: Nos. 17-20. — . 1923. Sobre um caso de Favus em um gallo causado pelo Achorion gallinae (Megnin 1881), Brasil Med. 37: 2: 198. Aronstam, Noah E. 1922. Epidermophyton inguinale with unusual distribution, with report of 4 cases [in one family], Urol. Cut. Bev. 26: 280. Artom, Mario. 1933. Imponenti formazioni tumorali da Trichophyton in tricofizia universale, Giom. Ital. Derm. Sifilol. 74: 563-582, 10 pis. Arzt, Leopold & Herbert Fuhs. 1921. tjber mykotische AUgemeininfektionen bei Trichophytie und Mikrosporie ("Trichophytosen und Mikrosporosen"), Arch. Derm. Syphilis 136: 333-344. — . 1921. tJber Sycosis parasitaria mit besonderer Beriicksichtigung der speziellen Therapie, Derm. Zeitschr. 32: 91-113. — . 1922. Weitere Beitrage zur Klinik und Therapie der Audouinischen Mikrosporie. (Herpes tonsurans microsporicus und Mikrosporid), Acta Dermato-venereol. 3: 1-20, PI. 1. — . 1923. tJber Allgemeinerkrankungen bei Audouinischer Mikrosporie, Acta Dermato- venereol. 4: 59-94, Pl. 3. — . 1923. Zur Entstehung der AUgemeinexantheme bei Mikrosporie, Arch. Derm. Syphilis 143: 52-56, 1 fig. — . 1923. tJber durch Trichophyton violaceum hervorgerufene Pilzerkrankungen. Ein Beitrag zur Pilzflora in Wien, Derm. Woch. 76: 409-414. — . 1923. tJber diagnostische Impfergebnisse mit Mikrosporin (Hochst) bei Mikrosporie und artverwandten Pilzerkrankungen, Derm. Woch. 77: 997-999. — . 1924. Zur Kenntnis der durch das Epidermophyton inguinale (Sabouraud) hervor- gerufenen Hauterkrankungen, Derm. Zeitschr. 41: 97-103. — . 1925. tJber eine seltenere Form der atypischen Audouinischen Mikrosporie: kerion Mikrosporicum, Arch. Derm. Syphilis 149: 9-12. — . 1928. Die Mikrosporie, Handbuch der Haut- und Geschleohts-Tcrankheiten. [Jadassohn.] 11: 607-654, 30 figs. Ashford, Bailey K., Earl B. McKinley & George B. Dowling. 1930. Experimental inocula- tion of monkeys (Silenus rhesus) and guinea pigs with two dermatophytes and one Blastomycoides. Porto Bico Jour. Publ. Health Trop. Med. 5: 452-457, 1 pl. Ayres, Samuel Jr. & Nelson Paul Anderson. 1934. Inhibition of fungi in cultures by blood serum from patients with phytid eruptions. Arch. Derm. Syphilol. 29: 537-547. Balbi, Edoardo. 1933. Anticutine e procutine nelle dermatomicosi, G^oni. Ital. Derm. Sifilol. 74: 889-910. •Balina, P. E. Aubrun, & Pablo Negroni. 1930. Epidermomicosis por Epidermophyton rubrum de Castellani, Asoc. Argeoitina Derm. Sif. 4 abril. Ballagi, Stefan. 1926. Die in Ungarn einheimischen Mikrosporon Trichophyton Epidermophy- ton und Achorionpilze, Derm. Wodi. 83: 1155-1169. Ballagi, Stefan & Stefan Laubal. 1933. tJber Meerschweinchenimpfungen mit Sehimmel- pilzen und Dermatophytenarten, Arch. Derm. Syphilis 167: 394-400, 5 figs. Ballarini, M. 1925. Casi di tricofizie a sede insolita. Arch. Ital. Derm. Sif. Ven. 1: 76-80 5 figs. 564 MEDICAL MYCOLOGY Balzer, Felix. 1883. Notes sur 1 "histologie des dermatophytes, Aroh. Physiol. Ill, 2: 466- 474. — . 1881. Recherches histologiques sur le favus et la trichophytie, Arch. Gen. Med. 148: 385-410. Bang, Henrik. 1910. Sur une trichophytie cutanee a grands cercles causee par un derma- tophyte nouveau. Trichophyton purpureum Bang, Ann. Derm. Syphiligr. V, 1: 225- 238. Barbaglia, Vittorio. 1914. I Tricofiti della provincia di Sassari, Giorn. Ital. Mai. Ven. Pelle. 55: 911-929. — . 1931. Su un case di favide. Contribute all' etiologia ed alia i^atogenesi del favide, Arch. Ital. Derm. Sif. Ven. 6: 337-354, 5 figs. Barbalian. 1932. Epidemie de trichophytie cutanee dans une maison de couture, Bioll. Soc. Frang. Derm. Syphiligr. [43:] 543-546. Bargum. 1909. Kerion bei Mikrosporie, Monatsh. Pralct. Derm. 39: 84-87. *Bassewitz, F. von. 1926. Frequencia das dermatomycoses no Rio Grande do Sul, Con.gr. Sudwmericano de Derm. 1: 21. [Ann. Derm. Syphiligr. VI, 7: 365, 1926]. *Baudet, E. A. R. F. 1921. Herpes bij het paard veroorzakt dorr Trichophyton grauulosum, Tijdschr. Diergeneesk. 48: 379-390. — . 1930. Sur une teigne trichophytique du dromedaire produite par une espece nouvelle de Grubyella, G. Langeroni n. sp., Ann. Parasitol Hum. Comp. 8: 411-418, ^ pis., 6 figs. — ■. 1930. Sur un cas de teigne humaine produite j)ar un dermatophyte megaspore, Aim. Parasitol. Eim. Comp. 8: 512-519, 1 pi., 6 figs. — . 1930. Particularites d 'une teigne megasporee d 'un veau, Ann. Parasitol. Hum. Comp. 8: 520-522. — . 1931. Les Trichophyton a cultures faviformes sur milieux a base de polysaccharides d'Langeron et Milochevitch, Ann. Parasitol. Hum. Comp. 9: 546-551, Pis. 12, 13. — . 1932. Recherches experimentales sur les Trichophyton animaux a cultures faviformes, Ann. Parasitol. Hum. Comp. 10: 520-541, Pis. 24-30. *Bazin, [Antoine Pierre] Ernest. 1854. Considerations sur la mentagre et les teignes de la face, Paris. * — . 1855. Recherches sur la nature et le traitement des teignes, Paris, 152 pp. — . 1858. LeQons theoriques et cliniques sur les affections cutanees parasitaires (redig^es par A. Porquet) Paris, 236 pp., 5 pis. ed. 2, Paris; 292 pp., 5 pis., 1862. Beckett, Clinton G. 1930. Ringworm or tinea infection of the toes, JJ. S. Veterans Hospital Bidl. 6: 793-798. Beclerc. 1894. Les teignes tondantes a I'ecole des teigneux de I'hopital Saint-Louis (£:cole Lailler), Ann. Derm. Syphiligr. Ill, 5: 685-693. Beeson, B. Barker. 1915. Ringworm of the scalp in Chicago, a bacteriological study of one hundred cases. Jour. Cut. Dis. 33: 731-737, Pis. 42, 43. Behdjet, Houloussi. 1927. Die Trichophytie der Augenbrauen und der Wimpern und die seltenen Formen der Trichophytie, Derm. Woch. 85: 1028, 1029. — . 1930. Epidermophytie Castellani oder Trichophyton purpureum Bang, Derm. Woch. 91: 1623, 1624. Beigel, Hermann. 1865. tJber Auftreten und Bersten der Haare, eine eigenthiimliche Erkrankung des Haarschaftes, Sitsmigsher. Math. Natww. Kl. E. Alcad. Wiss. Wien. 17: 612-617, 1 pi. Beintema, K. 1933. Klinische und kulturelle Beobachtungen bei Achorion Quinckeanum, Derm. Zeitschr. 68: 21-27, S figs. — . 1934. Tiber einem neuen Pilz der Endothrixgruppe: Trichophyton florifornie n. sp., Aroh. Derm. Syphilis 169: 577-581. Bellini, A. 1899. Studio sulla profilassi e la cura delle tigne, Giorn. Ital. Mai. Ven. PeUe. 40: 70-104. TRICHOPHYTONEAE 565 Bennett, John Hughes. 1842. Ou the vegetable nature of tinea favosa (Poirigo lupinosa of Bateman) its symptoms, causes, pathology and treatment, London 4~ Edinburgh Monthly Journ. Med. Sci. 2: 50-1-519, Pl. 5. Berde, Kd.roly von. 192(5. tiber die Widerstandsfahigkeit des Mikrosporon Audouini, Arch. Derm. Syphilis 150: 225-235. — . 192fi. tJber die pathogene Fadenpilzflora in Siidungarn, Areh. Derm. Syphilis 152: 126- 131. — . 1927. Delmagyarorsz4g Korokozo fonalgombaflorajol, Folia Cryptogamica [Szeged] 1: col. 279-282. [Tr. tJber die pathogene Fadenpilzflora in Siidungarn. col. 283-286.] — . 1927. Favus capitis unter dem Bilde einer Folliculitis sclcrotisans. Derm. Woch. 84: 669-676, 5 figs. [*Borgyogysucrose>glucose>lactose. On lactose, filaments have a chocolate brown color. Raulin's neutral gelatin becomes clear, colorless liquid in 6 days. Broth gelatin is similar, with aspergilloid forms common in growth. Egg albumen not liquefied. Thom and Church describe this organism as follows : "Colonies on Czapek's solution agar with cane sugar spreading broadly, with vegetative hyphae mostly submerged, with fruiting areas at first color- less, then passing through orange yellow shades to brown in old colonies (variously Isabella color, light brownish olive, buff}' citrine, medal bronze or raw umber. Ridgway, column 19, on Plates XXX, XVI, IV and column 17, Plate III), not showing true green; reverse uncolored or occasionally pinkish ; stalks arising from submerged hyphae, up to 1 to 2 mm. in length, becoming several millimeters in length upon com or other concentrated media, 10-20/^ in diameter, increasing in diameter towards the apex and passing rather abruptlj^ into vesicles with walls rather thick, 1 to 2/a, becoming abruptly thinner at the base of the vesicle, pitted more prominently in upper than in lower half (often appearing as rough or echinulate with low magnifications) and frequently showing irregular thickenings witliin ; vesicles 25-50;u, in diam- eter, with fairly thin walls which frequently crush in mounts; heads varying greatly in size in the same fruiting area, from more or less columnar to nearly but not completely spherical and up to 350/i, in diameter, with radiating chains and columns of conidia ; phialides, one series in small heads, two series in large heads, primary commonly 7 to 10 b}' 3 to 4/x, becoming 20 to 35//. long in gigantic heads upon corn, secondary 7 to 10 by 3/a; conidia more or less pyriform toward globose, tuberculate especially at the distal end in the chain, 5, 6, occasionally up to 8/x in diameter, rough from prominent masses and ASPERGILLACEAE 627 bars of orange yellow coloring matter deposited under the loose outer wall upon the firm inner wall. ISclerotia occasionally produced, usually purple or reddish ])uri)le, globose to pyriform with apex white." Aspergillus Gratioti Sartory, C. R. Acad. Sci. 170: 523, 524, 1920. Found on the nails of a Chinese workman in France. Pieces of the nail, on being treated with 40% KOH, show sometimes sep- tate mycelium, sometimes unsegmented. Chlamydospores 20-40;u, in diameter, sometimes terminal, frequently intercalary. Conidia free, 3.5-4^ in diameter. In culture, hyphae grayish, then brown and black, 0.6-1. 5/i, in diameter, much branched. Fertile hyphae short with delicate walls about 4-5ju, in diameter, vesicle spherical, 8-20/a in greatest diameter. Phialides elliptic, 6/a long, very close-packed. Conidia globose, 3-3. 5/x in diameter, brown. Sclerotia and peri- thecia not seen. On Raulin's maltose agar, colony is moist and grayish, becoming visible on the fifth day, black by the tenth. Colony elevated, irregularly rugose in the center, less so at the margin. Conidiophores in concentric circles. Seen from below, culture appears brown with the substrate also colored. No marked odor. Growth on glucose agar the same. Growth the same, but less pronounced on other media; e.g., potato, potato glycerol, carrot, Sabouraud's nutrient gel- atin, a variety of Raulin's sugar solutions (sucrose, glucose, lactose, maltose), and milk. No growth on coagulated beef serum, egg albumen, or acid potato. Milk coagulated on the twelfth day, curd digested. Gelatin liquefied on the fifth day. Aspergillus subfuscus Johan-Olsen [Sopp] & Gade, Nord. ^led. Ark. 18: 9:14-16, 1886; Johan-Olsen [Sopp], Bot. Jahresber. 14: 476, 1886; Meddel. Naturhist. Foren. Kristiania, 1885. Sterigmatocystis suhfu.^ca, Saccardo, Syll. Fung. 10: 526. 1892; Sartory & Jourdre, C. R. Soc. Biol. 64: 928, 1908. Pathogenic when injected into rabbits. Mycelium snow-white, coarsely articulate, sparingly branched. Curious raylike involution forms present, conidiophore 18-20/x from base to vesicle, vesicle 30-90/a, head, including chains of conidia, 90-200/a in diameter. Phialides 20/i long, conidia 3-3.5/* in diameter, olive brown, spherical, smooth. At opti- mum temperature of 36-38° C, conidia appear in 30-36 hours, more slowly at lower temperatures. No perithecia or sclerotia reported. Organism grows best on bread medium (10 c.c. finely crumbed bread. 125 c.c. H2O, and 2 drops lemon juice), also groAvs on agars and gelatin. Giant cultures, grown at room temperature, are olive brown in color. At 37° C, color deeper, then greener. Mycelium white, quite smooth then raised above the substrate. Colony lightest at the edges, deepest in color at center. Aspergillus cinnamominus (Weiss) Dodge, n. comb. Sterigmatucystis cinnamuminus Weiss, Ann. Parasitol. Hum. Comp. 8: 189- 193, 5 iigs., 1930. 628 MEDICAL MYCOLOGY Isolated from a case of pityriasis versicolor fiava of brown spots with discrete depigmentation and a few small vesicles at the borders. Reinocula- tion into human skin gave typical lesion. On rabbit, scarification produced epidermic lesions, while subcutaneous inoculation gave rise to subcutaneous gummata, with the picture of generalized pseudotuberculosis revealed at autopsy. Hyphae septate and branched ; conidiophores simple, 5/a in diameter, vesicle 12 x 9/a. Primary phialides cylindric, 4/a long, bearing 2-3, sometimes 4 secondary phialides, 5-6/t long. Conidia spherical, 1-2/a in diameter, brown. Other spores, borne laterally on short branches (phialides?), 3-4)u,. Chlamydo- spores occasional. Under some conditions, only monstrous phialides are formed, with a "pleomorphic" mycelium resulting. Aspergillus Hortai (Langeron) Dodge, n. comb. Sterigmatocystis Hortai Langeron, Bull. Soc. Path. Exot. 15: 383, 384, Fig. 1, 1922. Found in a case of otomycosis in Brazil. Mackinnon (1932) refers this species to A. terreus Thom. Conidiophore appears empty of protoplasm while neighboring cells ap- pear full. Vesicle 5-10/i, in diameter, primary and secondary phialides 5 x 1.5/i,, scarcely covering upper half of vesicle. Primary phialides give rise to two, occasionally three, secondary phialides. Conidia smooth, 2.5fi in diameter, in long chains. Aspergillus Phoenicis (Corda) Thom & Church, Aspergilli 175, 1926. Ustilago PJioenicis Corda, Icon. Fung. 4: 9, PL 3, Fig. 26, 1840. Sterigmatocystis Phoenicis Patouillard & Delacroix, Bull. Soc. Myc. France 7: 119, PI. 9, 1891. Sterigmatocystis antacustica Cramer, Vierteljahrschr. Naturf. Ges Ziirich 4: 325-337, PI. 2, 1859. Saprophyte, originally isolated from dates. 8. antacustica Cramer isolated from external ear of man. Aspergillus niger Risel (1906) non al. from lungs apparently belongs here. Vesicles 45/t in diameter ; primary phialides 20-50/x long, several secondary' phialides to a primary phialide ; conidia 2.5-3. 5/i,. This species has often been confused with Aspergillus niger Tieghem by subsequent writers, but differs in the much longer primary phialides. It is probable that Aspergillus nigricans Wreden, C. R. Acad. Sci. Paris 65: 368-370, 1867, belongs here, but it was too briefly described to place defi- nitely. Apparently it was not cultivated. Aspergillus niger Tieghem, Ann. Sci. Nat. Bot. V, 8: 240, 1867. Sterigmatocystis niger Tieghem, Bull. Soc. Bot. France 24: 102, 103, 1877. This species is predominantly saprophytic but occasionally parasitic, es- pecially in the human ear. Unless pathogenicity is clearly proved, reference ASPERGILLACEAE 629 to this organism should be regarded witli suspicion, as it is widespread and common. Thom characterizes the species group as follows: "Colonies rapidly growing, with abundant submerged mycelium, hyaline, aerial hyphae usually scanty; conidiophores rising directly from the sub- stratum, hyaline or yellow to brown near the vesicle, smooth, wall thick, with- out pits, but frequently uneven on the inner surface, splitting lengthwise into strips when broken, not septate or with occasional thin septa, varying greatly in length on different media ; conidial heads deep brown to black, spherical, radiate ; vesicles spherical or subspheric, thick-walled, commonly 20-50/x in diameter, hyaline or yellow brown ; phialides usually in two series, the primary covering the vesicle, 20-30/* in length ; secondary phialides 6-10 x 2-3/i, brown to black ; conidia spherical, thin-walled, smooth with diffuse brown or fuscous color then rough or spinulose from coloring matter deposited as tubercles or bars between the outer and inner walls, 2.5-4/i, in diameter." Aspergillus giganteus (Mattlet) Dodge, n. comb. Stcrigmatocystis gigantens Mattlet, Ann. Soc. Beige Med. Trop. 6: 31, 32, 1926. Sterigmaiocystis cas V ]\Iattlet, Ann. Soc. Beige Med. Trop. 4: 167-176, 3 figs., 1924. Along with this species a Parasaccharomyces irritans was isolated from a case of severe bronchomycosis in the Belgian Congo. Animal inoculations not reported. Colony yellow saffron at first, then becoming black from the conidia. My- celium variable, 1.5-6/a in diameter, with dark granules ; conidiophore hyaline 12-13/x in diameter, 800-1,000/1 long; vesicle 50/i in diameter, about 50 phialides per great circle, covering the whole vesicle ; primary phialides about 30/i long, rarely 2-celled, bearing 3-4 secondary phialides about lOfx long, heavily pig- mented; conidia in chains, 3/a in diameter, echinulate, black, thick-walled, finally 4-5/* in diameter. Optimum temperature 37° C, growth slow at 21°- 23° C. There is little to distinguish this species from the preceding. Aspergillus Macfiei Dodge, n. sp. Sterigmaiocystis sp. Maefie, Ann. Trop. Med. Parasitol. 15: 279-281, 1921. Isolated from a case of otomycosis in the ear of a European. Caused irritation but no serious damage or deafness. Nonpathogenic in scarified ear of pouched rat (Cricetomys gambianus) , sheep, or monkej^ (Cercopithecus patas). No effect on intraperitoneal injection in rat. Hyphae hyaline, conidiophores about 1 mm. long, 14/t in diameter, white, then darkening. Vesicle slightly broader than long, 40 x 37/i-, almost com- pletely covered by phialides. Primary phialides 12/x, long (average of ten), secondary phialides 8/t long (average of ten). Conidia dark brown, spherical, 4-5.2/A in diameter, average 4.5/i. On Sabouraud maltose agar, growth is rapid and spreading, yellowish white and feltlike, then fluffy as conidiophores are produced. These are brown to black. On glucose agar at 28° C, there is a white, puckered, and wrinkled 630 MEDICAL MYCOLOGY surface growth in 24 hours. Then yellow, wrinkled, feltlike, darker with conidiophores beginning. Growth more rapid at 37° C. Growth on potato rapid and abundant. On gelatin stab, there is no deep growth. After a while, cerebriform whitish masses appear just below surface growth. In peptone water, growth is mainly surface with some "puffy balls." Glu- cose peptone water is acidified but not fermented. Litmus milk turns acid and clots. Surface growth is yellowish. Coagulated blood serum shows a very slow white growth with rare conidiophores. It is eventually liquefied. Gelatin is not liquefied. Aspergfillus Menderi Sartory & Flamant Apud Sartory, Champ. Paras. Homme Anim. 578, 1922. Aspergillus sp. Sartory & Flamant, C. R. Soe. Biol. 93: 1114, 1115, 1920. Isolated from sputum of one suspected of having tuberculosis. Found three times. Not pathogenic to laboratory animals. Mycelium branched. Conidiophores erect, 220-740/x high, 14-16/* in di- ameter. Vesicle 30-50/a in diameter, green at first, becoming brown. Phialides on upper half of head only, irregular but usually twice as long as wide. Conidia irregular, 9-12/t in diameter. Disjunctors present. Perithecia canary yelloAv on carrot or potato, 200-220/* in diameter, with a bright yellow mem- brane. Asci spherical, 15-20/* in diameter, 8-spored. Spores lenticular with groove and crest along greatest diameter, 10 x 4.7/t. Mycelium sprouts on carrot, potato, and licpiid media. Growth good on the usual media. Glucose only fermented. No action on starch, cooked rice, eg^ albumen, coagulated serum. Milk coagulated and curd dissolved. Gelatin liquefied. Aspergillus Amstelodami (Mangin) Thom & Church, Aspergilli 113, 1926. Euroiium Amstelodami Mangin, Ann. Sci. Nat. Bot. IX, 10: 360, 361, 1909. This species is a common saprophyte that from time to time has been re- ported pathogenic. Weil, Gregoire, Chevallier & Flandrin (1928) claim to have isolated this species from splenomegaly [probably as a contaminant]. Duche describes a var. alophote similar to the species, but lacking the crest on the ascospore. Fonseca (1930) reports this species from a case of mycetoma of the foot with yellowish grains, his organism being determined by Mangin. Conidia green, spherical, finely echinulate, 2.8-4.7/* or somewhat ellipsoid (Fig. 100, A), larger at lower temperatures (5.6 x 7.5/i at 10° C). Perithecia numerous, yellow, deeply surrounded by floccose mycelium; ascospores with furrow only, hyaline, smooth, 4.7 x 3.7/i (Fig. 100, B, C). Colony floccose, tardily green, glaucous or olive green. Aspergillus ruber (Spieckermann & Bremen) Thom & Church, Aspergilli 112, 1926. Eurotium ruhrum Spieckermann & Bremer, Landw. Jahrb. 31: 81-128, 1902. Case by Agostini, Atti 1st. Bot. R. Univ. Pavia IV, 2: 65-77, 1930 [1931]. ASPEKCILLACEAE 631 Isolated from small subepidermal abscesses in the hairy areas of head which open to the surface by small yellow orifices, covered by a purulent crust. Pathogenic for white rats, producing ulcers. Conidiophores arise either from submerged or aerial hypliae, 120/a long and more, when forking from tlie latter, up to 30()-500/x or even 800/x in length and 8-16(0, in diameter. Vesicle clavate to subspheric, 24-30/z in diameter, phialides radiating, 4-12(a, mostly 6-10/x by 2-4/^. Conidia ovoid to spherical. 5-8/x in long axis, rough, pale green (blue green in mass). Perithecia globose, yellow then rusty brown, 80-120/a in diameter; asci 10-12^ in diameter; asco- spores 6 x 4-5/a Avitli shallow, smooth furrow, no frill or ridges. Colonies more or less floccose, at first white, then bluish green with de- veloping conidia, then through yellow to rusty red throughout and intense red in the substratum. Aspergillus montevidensis Talice & Mackinnon, Soc. Biol. Montevido 61 Sesion. 4 pp. 1931; Reunion Hoc. Argent. Patol. Reg. del Norte, Tucuman 7: 278-284, 1932. Fig. 100. — Aspergillus Amstelodami. A, conidia; B, ascospores ; C, asci containing ascospores. (After Fonseca 1930.) Isolated from a ease of otomycosis in ^Montevideo. Tympanic membrane was covered with dark, slightly adherent spots in which the Aspergillus was demonstrated. Fertile mycelium at first glaucous (glaucus 38 Saccardo, Chromotaxia), then yellowish (flavo-virens 33 Saccardo), and finally yellowish green (oliva- ceus 39 Saccardo). Sterile mycelium Avhite or grayish (cremeus 27 Saccardo). Conidiophores, 55-234^i long (mean 170/i) by 5.1-8.5/i, in diameter (mean 23/u.), not visible on casual inspection, smooth, terminating in a spherical vesicle, 13.5-25.5/x in diameter (mean 18.7/a), of greenish or yellowish green color. Phialides elongate, inserted over almost the entire surface of the vesicle, not crowded, 6.8-13.6 x 2.5-3.4/i (mean 10.8 x 3/^). No secondar}* phialides. Conidia usually ovoid, sometimes ellipsoid or spherical, 3.4-o.6;u in diameter (mean 4.9/x), generally smooth and forming short chains which are easily broken up in the making of preparations. Fertile perithecia abun- 632 MEDICAL MYCOLOGY dant, greenish yellow, each having a variable number of asci with 8 ascospores per ascus. Spores grooved and crested, the latter visible but not very distinct. Perithecia 34-102ju in diameter (mean 68/x) ; asci 7.6-11.4/x, in maximum diameter (mean IO.6/1.) ; ascospores 5.45 x 3.75/t. Optimum temperature for growth 25°-30° C. On glucose peptone agar a diffusible yellow pigment is formed. Gelatin is not liquefied within 10 days. Aspergillus Fontoynonti Gueguen, C. R. Soc. Biol. 66: 1052, 1053, 1909; 67: 10-12, 1909; Arch, de Parasitol. 14: 177-192, 1911. Isolated from secondary abscesses in the cervical region of a European resident in Madagascar, not from "nodosites juxta-articulaires" as first re- ported by an epistolary error. Abscess was on indurated base in the hypo- derma, nearly painless, about the size of a large nut. Full case history given. Medication with 2 gm. KI per diem for several months successful. Intravenous or intraperitoneal injection into rabbits, guinea pigs, or pigeons gave negative results. Conidiophores about 150-200/x high, excipuliform, subcontinuous, 14-18/* in diameter. Phialides of flask shape, 8-12 x 2/a, giving rise to chains of 10 to 30 glaucous conidia which are spherical to ovoid, 4-6 x 3-5/i,, more or less covered with extremely fine warts. Conidia in short flammiform plumes, rarely cylindric. Early subcultures with heads proliferating and very irregu- lar. Later subcultures more regular, optimum temperature 22°-25° C, no growth at 37° C. On gelatin, there is visible growth by the fourth day with variable glau- cescence. On agar, growth with less glaucescence, culture becoming greenish gray. On potato, punctiform colonies appear on the fifth day and become confluent on the ninth, show few conidiophores and remain dirty white. On glycerol potato, growth less abundant and white. On carrot, growth visible the fourth day, pale glaucescence the eighth, with finally a large elevated cushion. On Jerusalem artichoke, culture turns green on the eighth day and color is somewhat deeper in 3 weeks. No growth on Raulin's normal medium. On Raulin's neutral medium, punctiform colonies appear on fourteenth day with grayish, circular covering the third week. On Raulin's normal gelatin, very small star-shaped colonies appear on the tenth to twelfth days, fruiting in 3 weeks, greenish white. On Raulin's neutral gelatin, growth better, no liquefaction after 40 days. In peptone broth, fructifications in 12 days, glau- cous color third week. Coagulated albumen shows white punctiform colonies on the eighteenth day and thereafter is stationary for one month. Organism grows on a variety of sugars but best on maltose. It hydrates urea with feeble formation of NH3. Milk is coagulated in 12 days, with curd subsequently digested into a citron yellow liquid with light chalky sediment. Plain gelatin liquefied after a month. Aspergillus malignus (Lindt) Thom & Church, Aspergilli 132, 1926. Eurotium. malignum Lindt, Arch. Exp. Path. Pharm. 25: 256-271, Figs. 1-11, 1889. ASPERGILLACEAE 633 Obtained from human ear. Pathogenic to puppies, with production of extensive lesions. Conidiophores up to 1,000;^ long ; vesicles 22-24/a in diameter, approxi- mately spherical instead of cylindric, phialides hyaline, 10 x 4-4.5/x, more or less divergent, covering upper two-thirds of the vesicle [while in A. fumigatus the upper half only is covered, and the phialides measure 8 x 3-4/t] . Conidia 3-4/* in diameter. Perithecia are common, being particularly abundant on bread. They are white, of woven hyphae, 40-60/x in diameter, surrounded by a pseudoparenchyma. Asci 8-spored, 14-18/* in diameter, spherical, scattered throughout the sclerotioid perithecium. Ascospores produced on bread in 7 days. Spores lenticular or biconvex, 6-8/i in long axis, colorless, having a thick exospore with two ridges and a furrow and some markings on surface of the valves. Colonies on agar gray blue. Judging from the figure, the antheridial hypha degenerates, unless Fig. 6 shows a much younger stage than interpreted by the author. Aspergillus fumigatoides Bainier & Sartory, C. R. Soc. Biol. 66: 22, 23, 1909; Bull. Soc. Myc. France 25: 111-118, PI. 5, 1909. A saprophyte found to be pathogenic to rabbits on inoculation. Foot tortuous, without partition, lightly and progressively curved from bottom upward. Conidiophore short, 50-310/i high, 5-6/*, in diameter at the base. Vesicle 30-35/1. broad, phialides 8-14/i long, wholly covering the vesicle, colorless. Conidia dark olive, ovoid, small, 2-3/* in diameter. Perithecia appear on solid media, 65-92/i in diameter, collected in masses. Asci spherical to ovoid, 20-26 X 12-18/1 with usually 8 ascospores per ascus, rarely 4, spherical, 3-3. 5/c in di- ameter, rough. Organism not very heat resistant, dying at 48°-49° C. Color of cultures on carrot slightly green, not ashy gray fuliginous and blackish as in A. fumigatus. Aspergillus bronchialis Blumentritt, Ber. Deutsch. Bot. Ges. 19: 442-446, PI. 22, Figs. 1-6, 1901; 23: 419-427, PI. 19, Figs. 1-3, 6-8, 19, 23, 1905. Found by Chiari in the bronchial tubes of a diabetic patient. Studied by Blumentritt. Not pathogenic to dogs and guinea pigs, pathogenic to hens and pigeons (Luksch 1902). Aerial hyphae 5-8/t, submerged hyphae 2-4.2/t, conidiophores erect, simple, seldom septate, hyaline, 280-300/i tall; vesicle 12-19/t, conidia gray, greenish gray, and brownish, depending on medium, mostly gray green. On agar colony, at first white, then greenish, becoming dark green or brown. On broth, white pellicle becomes green, tube finally filled with myce- lium; obligate aerobe. On gelatin, colonies as on agar, gelatin liquefied after a few weeks. Colony floccose in contrast to velvety in A. fumigatus. Aspergillus virido-griseus Costantin & Lucet, Ann. Sci. Nat. Bot. IX, 2: 119-171, 1905. Pathogenic to rabbits, not to fowls. Mori (1916) isolated it from the pleuropulmonary exudate of goat. 634 MEDICAL MYCOLOGY Colonies very floccose, areas partly white, partly g'ray, cream gray to dark gray green or green; eonidiopliores thin-Avalled, septate, undulate, slightly colored or tardily colored near the vesicle, 400-620/x, long, vesicle 25-36/a in diameter, phialides 5/x in length; conidia 2.8/>i in diameter. Doubtfully distinct from A. fumigatus. Aspergillus fumigatus Fresenius, Beitr. z. Myk. 81, PL 10, Figs. 1-11, 1850. Aspergillus glaucoides Spring, Bull. Acad. Sci. Belg. 19: 560-572, 1852. Aspergillus nigrescens Robin, Hist. Nat. Veg. Paras. 518, PI. 5, Fig. 2, 1853. Aspergillus pulmonum-hominis Welcker in Kiichenmeister, Die in und auf des lebenden Menschen vorkommenden Parasiten. II. Planzliche Parasiten 144, 1855: Dusch, Arch. Path. Anat. Phys. [Virchow] 11: 561-566, 1857. Aspergillus ramosus Hallier, Zeitschr. Parasitenk. 2: 266-269, PI. 6, Figs. 1-6, 1870. Aspergillus aviarius Peck, N. Y. State Museum Rept. 44: 120, PI. 4, Figs. 9-12, 1890 [1892]. Aspergillus fumigatus var. alpha Sion & Alexandrescu, C. R. Soc. Biol. 64: 288, 289, 1908. This is the commonest species isolated from cases clinically resembling tuberculosis of the lungs in which Mycohacterium tuberculosis has not been found. Apparently causing- severe epizootics in birds, less fatal in man, and not reaching epidemic proportions. Pathogenic for laboratory animals. Conidiophores short, usually densely crowded, as long as 300/a (rarely 500/a) by 2-8ju in diameter, more or less green in color, especially above, arising either from submerged or aerial hyphae, septate or unseptate, enlarging to- ward the top with flask-shaped vesicles, 20-30/a in diameter, usually fertile only on the upper half. Primary phialides only, 5-10 (mostly 6-8) x 2-3/li, crowded, with axis approximately parallel to axis of the conidiophore. To- gether with the chains of conidia, the growth is occasionally about 400/* high, though usually shorter, and measures 50/jb across at the top. Conidia appear dark green, spherical, 2-3.5 (mostly 2.5-3 )/x in diameter. Growth good at 37° C. Colonies on Czapek's solution agar velvety to felted floccose, green to dark green to almost black in age, spreading. Reverse and medium colorless to yellow, occasionally^ reddish in age. On the basis of ascospores reported by Sartory, C. R. Acad. Sci. 183: 1360-1362, 1926, Vuillemin erected Sartorya fumigata (Fresenius) Vuillemin. C. R. Acad. Sci. 184: 136, 137, 1927. Perithecia in sclerotia, walls of several cell layers, spherical; asci ovoid or spherical, thin-walled, disappearing at maturity. Ascospores 8, ellipsoid, thick-walled, black at tips, lighter in centei ; perithecia 70-110/1, in diameter; asci 15-22. 5/a; ascospores 4-6.25/j, long, perithecial filaments 3-5. 5/x in diameter, walls 0.8-1.2/. thick. The perithecia were reported following treatment with x-rays. This work has not been confirmed. Previous reports of perithecia in this species have all been shown later to be erroneous. ASPERGILLACEAE 635 Var. minimus Sartory, Bull. Acad. Med. Paris, III, 82: 304, 305, 1919. Isolated from the sputum of a tuberculous suspect. Pathogenic to rabbit and guinea pig. Mycelial hyphae 2-3/x in diameter, forming a close felt. Conidiophores erect, slightly tortuous, 100-150/* long by 4-5/1, in diameter, near base (gradu- ally enlarging upward), fuliginous gray at the base, deepening in color to- ward the head, vesicle spherical, 20-25/i in diameter, covered in upper half or two-thirds by phialides 5-8/i long. Conidia spherical, rarely ellipsoid, 2-3/x in diameter, bronzed. No perithecia found on any media tried. Growth good at 37° and ceases at 42° C. Organism has been grown on starch paste agar, Raulin's agar, gelatin, Raulin's sucrose, glucose, maltose, and lactose media. Sucrose is inverted. Neither glucose, maltose, nor lactose is fermented. Starch liquefied with reducing sugars present by the fifth day. Egg albumen unchanged. Milk coagulated on tenth day, transformed to an opalescent liquid. Gelatin lique- fied in 8 days (color 373, Code des Couleurs). Asperg^lus tunetaneus (Langeron) Dodge, n. comb. Sterigmatocystis tunetana Langeron, Bull. Soc. Path. Exot. 17: 345-347, 1 fig., 1924; Blanc & Caillou [case history only], Bull. Soc. Path. Exot. 17: 343-345, 1924. Isolated from tumors between the third and fourth metacarpals on the left hand of a native of Tunis. Tumors removed surgically. Animal inocula- tions negative. Conidiophores simple, 200/x high by 2.5-4/a in diameter, not septate except in old cultures. Vesicle hemispheric above, obconic tapering below, varying with age from 5-10/x in diameter, the average being 7-8/x. Primary phialides short, oblong to inverted conical, 6-8 x 2.5-3. 5/i. Secondary phialides ampul- liform with more or less elongated beak, in groups of two to three, measuring 7-8.5 X 3-3.5/1. Conidia spherical, minutely verrucose, 3-3.5/1 in diameter, in long chains, greenish under the lens, deep blue green in mass (390-395, Code des Couleurs) and remaining so even after the mycelium is red. Optimum temperature 25°-30°, growth weak at 37° C. Colonies white, then green blue (368, Code, later 390-395), later orange (121, Code, ochraceus 29 of Chromotaxia) finally becoming orange (113, Code, fulvus 32 of Chromotaxia) . Asperg^illus Brodeni (Mattlet) Dodge, n. comb. Sterigmatocystis Brodeni Mattlet, Ann. Soc. Beige Med. Trop. 6: 31. 1926. Sterigmatocystis cas I, II, IV Mattlet, Ann. Soc. Beige Med. Trop. 4: 3: 167-176, 3 figs., 1924. Isolated from bronchomycoses with abundant sputum containing spores and later some filaments of Aspergillus, no trace of Mycohacterium tuberculosis either in smears or guinea pig inoculation. First case fatal (refused medica- tion), the second, a prisoner, partially recovered and disappeared, the third case apparently cured by KI. 636 MEDICAL MYCOLOGY Colonies greenish from conidia, sterile mycelium white. Mycelium vari- able, 1-3/A in diameter, cylindric, ovoid, or spherical ehlamydospores, 6-7/x in diameter. Conidiophores lOO-350/i, high, vesicle ovoid, 16 x 20fx, whole surface covered with phialides, 18-22 per great circle, 4/a long. Three to four sec- ondary phialides per primary phialide, 1-2 x 5-6ju,. Conidia in chains, 3/x in diameter, finely echinulate and bright green. Var. Vancampenhouti (Mattlet) Dodge, n. comb. Sterigfnatocystis Y ancampenliouti Mattlet, Ann. Soc. Beige Med. Trop. 6: 31, 1926. Sterigmatocystis cas III, Mattlet, Ann. Soc. Beige Med. Trop. 4: 167-176, 3 figs., 1924. Isolated from bronchomycosis, cases similar to those reported in A. Brodeni. Recovery followed Kl-emetine treatment. Close to A. Brodeni but less velvety, deeper green, with radial furrows in the colonies. Conidiophores very similar as to shape and size. Conidia bottle green, spherical, 3-4^ in diameter, or ovoid, 3 x 5/a. Aspergillus cyaneus (Mattlet) Dodge, n. comb. Sterigmatocystis cyaneus Mattlet, Ann. Soc. Beige Med. Trop. 6: 32, 1926. Isolated from a case of chronic bronchitis Avith a little emphysema. Au- thor unable to follow up case (Tamahunga). ]\Iycelium white, hyphae variable, 1-5/x, in diameter. Conidiophores 3-4|U in diameter by 150-300/a high. Primary phialides 3 x 8-lOyu,, secondary phialides about 1.5-2 X 4-6/x; conidia blue gray, smooth or echinulate, 3/x in diameter. Colony appears grayish blue, interrupted by straw yellow areas formed of many cespitose sclerotia (suggesting raspberry). Sclerotia [perithecia?] vary from 120-350/x in diameter and are composed of polyhedral cells, 5-18/a in di- ameter with very thick walls, 4-6/* in larger cells. Sclerotia hollow and allow a large volume of fatty droplets [ascospores?] to escape. Optimum temper- ture 30°-37° C. Aspergillus nidulans (Eidam) Lindt, Arch. Exp. Path. Pharm. 21: 284, 1886. Wtomyces purpureus Wreden, Arch. Augen-Ohr-Heilk. 3: 56-90, PI. C, 1874. Sterigmatocystis nidulans Eidam, Beitr. Biol. Pflanzen [Cohn] 3: 392- 411, 1883. Diplostephanus nidulans Neveu-Lemaire, Precis Parasitol. 101, 1921. Emericella nidulans Vuillemin, Champ. Parasit. 58, 1931. Examination of the Berkeley type in the Curtis herbarium shows that Emericella is unre- lated to A. nidulans. Found by Siebenmann in 2 cases of otomycosis. Conidiophores cinnamon brown in color, more or less sinuous, septate or not, 50-100/A, rarely 200 jx long by 3-5/* in diameter at base, increasing gradu- ally to obconic vesicle, glaucescent, becoming brown, 12 x 10/x,. Primary phialides approximately 5-8 x 2-3/x, secondary usually also present, 7-10 x 2-2.5/1,. Conidia spherical, 3/// in diameter, in chains. Perithecia in a nest of ASPERGILLACEAE 637 hypliae, spherical, 200-300/a in diameter. Asci 10.5-11/a in longest axis, 8-spored, filling the perithecia. Aseospores purple brown, slightly ovoid, 4 x 5/x, with a frill, giving the appearance of a biconvex lens, separating into two valves on germination. Colonies on Czapek's solution agar, white to yellowish green or fairly deep green, velvety to more or less floccose in purely conidial areas, becoming defi- nitely floccose when perithecia are forming. In aseosporic strains, colonies floccose with conidium production much reduced and green color absent or nearly so. Reverse and medium usually reddish to dark red or red brown. Perfect stage of A. fumigatus described by Grijns, 1903, belongs here (see Pinoy & Masson, 1915). Var. NicoUei Pinoy, apud Nicolle & Pinoy, C. R. Acad. Sci. 144: 396, 397, 1907. Sterigmafocysfis niduJans var. NicoUei Pinoy apud Nicolle & Pinoy, Arch, de Parasitol. 10: 457, 458, 1 pi, 1906. Diplosfej)hanus nidulans Neveu-Lemaire var. NicoUei Neveu-Lemaire, Precis Parasitol. Hum. 101, 1921. Isolated from a case of Madura foot in Tunis. Tissues of the foot ex- tensively invaded by the parasite. Puestow (1929) reports this organism from a case of maduromycosis of the forearm and mycetoma back of the right ear. (Organism determined by Thom as A. nidulans.) Not pathogenic to guinea pigs or rabbits. Young mycelium colorless. Conidiophores erect, simple, continuous or rarely septate, glaucescent or sometimes brownish, SOO/a long by 4/* in di- ameter, head obconic, 12 x lO/x. Primary phialides 8 x 3/x with two, rarely four, secondary phialides to each. These are 4 x 2.5/a, each giving rise to a chain of conidia, spherical, 2-3/x in diameter, smooth or slightly roughened, somewhat green. Chlamydospores terminal, spherical, 8-16/* in diameter, brownish. Sclerotia black brown, 50-300/* in diameter, embedded in nests of hyphae. Optimum temperature 36°-38° C. Var. Cesarii Pinoy & Masson, Bull. Soc. Path. Exot. 8: 11, 12, 1915. Caused mycetoma in the lung of an ass. Phialides often simple and inserted near the base of the vesicle. Conidia 3/* in diameter. Perithecia fertile, aseospores 4.1 x 4/*. Membrane turns brown rapidly. Aspergillus unguis (Weill & Gaudin) Dodge, n. comb. Sterigmafocystis unguis Weill & Gaudin, Arch. Med. Exp. Anat. Path. 28: 463-465, PI. 12, 1919. Found on rather superficial lesions of nail of great toe in deeply penetrat- ing brownish crevices of the nail, the remainder of which is deep yellow with a marbled appearance resulting. Mycelium 2-5/* in diameter (mean 3/*). Conidiophores simple, nonseptate, of uniform width, 250-300 x 5/*. Vesicle spherical, 12/* in diameter, or pyri- form, 14-16 x 8/*. Primary phialides cylindric, 5 x 3/*. Secondary phialides flask-shaped, 6/* long to the base of the neck and 3/* in diameter, four per 638 MEDICAL MYCOLOGY primary phialide. Conidia spherical, 3/x in diameter, becoming slightly ver- rucose in age, brownish in original lesions. Optimum temperature 30° C. Colonies at first white, then greenish or chrome green in center, then dirty, finally becoming chocolate brown. Almost always brown at 37°C. Giant colony after 10 days is 2 cm. in diameter, white, velvety, with center slightly depressed. GroAvth good on the usual laboratory media ; e.g., Sabour- aud maltose, carrot, Raulin's medium. Aspergillus Jeanselmei Ota, Ann. Parasitol. Hum. Comp. 1 : 137-146, 1923. Found on the nails of an old man in Paris. Nails had disappeared from all but four fingers, where they Avere deformed, thickened, grayish yellow. On other fingers there were thick crusts in place of the nails. Epidermis not affected. Mycelium branched, sparingly septate, 2-5 fx in diameter, sometimes 7/a in young cultures. Conidiophores 100-400/i, long, rarely 80/*, expanding gradu- Fig. 101. — Aspergillus Jeanselmei. 1, normal conidiophore ; 2, branched conidiophore, showing abnormal phialides ; S, conidia {1 and 2 X300 ; S X400). (After Ota 1923.) ally from 4-5/x at the base to nearly 8/a in diameter at the top. Heads includ- ing phialides and conidia measure 16-29/x across, occasionally up to 50/i. when both primary and secondary phialides are present. Phialides mostly simple, cylindric or fiask-shaped, 4-10 x 3-5/*, or when in two rows, the primary cylindric or slightly clavate, bearing two secondary flask-shaped phialides, 10-15 X 5/A, to each (Fig. 101). Conidia spherical, sometimes ovoid, 3.5-6. 5/x in diameter (mean 5/*), slightly green, becoming reddish brown in 7-month- old cultures, sometimes slightly verrucose, in long chains. On malt agar, glucose agar, carrot, or potato at 25°, mycelium white, greening on the third day, remaining yellowish green for months. On malt agar at 35°, growth is more rapid with colony turning green the second day. Colony on glucose agar becomes chocolate brown, with a greenish lustre in ASPERGILLACEAE GoO 7 months. On egg albumen, growth is feeble, graniilai-, greenisli brown. Conidia not produced on peptone agar without sugar. Mackinnon (1932) refers this species to A. flavus Link. Doubtful Species There is little to distinguish Sterigmatocystis tropicalis Matta, Bol. Inst. Brasil. Sci. 3: 51-54, 2 figs., 1927, from Aspergillus tunetanus, except its ver- milion color. It was isolated from ulcerous lesions on the back of the foot, removed surgically. PENICILLIUM Penicillium Link, Mag. Ges. Naturf. Freunde Berlin 3: 17, 1809. The type species is Penicillium expansum Link. Vegetative mycelium abundant, entirely submerged or more or less effused, monopodially branching, septate, commonly showing vegetative anastomoses, colorless or secondarily colored by products of metabolism wdiicli frequently also discolor the substratum, never with hyphal walls brown or black; colonies greenish or less commonly colorless, hazel, yellow, reddish, purplish or other shades ; conidiophores arising- as branches from the vegetative mj^celium, fre- quently perpendicular to the vegetative hyphae, but not showing differen- tiated foot cells as in Aspergillus, with walls in some species smooth, in others more or less conspicuously pitted or roughened from secondary thickening ; conidial apparatus forming a brush or broom, the penicillus, ranging from a single terminal verticil of conidiiferous cells or phialides or a terminal verticil of equal branches or metulae, bearing phialides in verticils ; phialides bearing single unbranched chains of conidia, each cut off as a cylindric segment from an apical tube ; conidia cylindric to ovoid, ellipsoid or commonly finally spherical, smooth or roughened, hyaline or variously colored, especially in mass; sclerotium or peritliecium formation known only in a few species. While Penicillium is frequently mentioned in medical literature, there are cnl^' two references where the organism has been sufficiently described to be sure that it belongs in this genus. With the exception of P. mycetomato- gerium and P. Bertai, none of the descriptions are adequate to permit identifica- tion of a modern culture as probably belonging to them. In general, it seems that the imperfectly described organism was picked up as a contaminant, and there is a probability of pathogenicity in only one of the cases cited. Penicillium Bertai Talice & Mackinnon, Ann. Parasitol. Hum. Comp. 7: 97-106, 1929. Isolated from case of chronic pseudotuberculosis on three successive occa- sions. Not pathogenic for rabbit and guinea pig. ]\Iycelium rampant, hyaline at first becoming yellow, about 2.5/x in di- ameter (2-4), with a tendency to become funiculose. Conidiophore erect, 20-90// X 2/^, tip enlarged to 3.2 ( 2-4.5 )/i terminated by up to 5 phialides, about 9 X 2.2|u, (6-16 X 1.5-3.5), which produce chains of conidia up to 20/*. Conidia smooth, spherical, 1.7-3.5/x in diameter (Fig. 102). 640 MEDICAL MYCOLOGY Grown on neutral Raulin's (Diercks) medium, beef broth gelatin, malt, agar, bread, potato, and Sabonraud agar (prepared according to the direc- tions of Biourge). Stab colony white with sulphur colored center, then green (glaucus of Saccardo Chromotaxia) finally dark green (atrovirens), reverse sulphur. Streak practically same color, changes finally, folded and fuliginous (11 Saccardo Chromotaxia), reverse luteous, the orange finally badious in the central zone with clear orange (137, Code des Couleurs). In beef gelatin, the reverse finally becomes ferruginous. In Sabouraud's conservation agar, colonies remain pinkish white (53a, Code des Couleurs). This organism belongs in the Aspergilloid section, close to P. platense Spegazzini. Penicillimn mycetomag'enum Mantelli & Negri, Giorn. R. Accad. Med. Torino 78: 165, 166, 1915 [as P. mycetogenum] ; Negri Atti R. Accad. Sci. Torino 66: 67-78, 1921. Fig. 102. — Penicillium Bertai. (After Talice & Mackinnon 1929.) Isolated from the foot of a man of thirty in Turin, Italy. The foot had been stepped on by an ox and gradually developed lesions which did not respond to iodine or KI treatment and finally, after 3 years, had to be ampu- tated. In section, the foot showed a fibrous periphery and a soft interior filled with black grains. Organism pathogenic to pigeon but not to guinea pig. In grains, hyphae septate, crowded into pseudosclerotia, subspheric or lobate, carbonaceous, 200-500ju, in diameter, solitary or in groups of two or three. Outer hyphae of the granules larger, 2.2/x in diameter, shorter, twisted and with a clavate apex, bound together into a crust by a black cement. Central hyphae are hyaline, 1.5/i in diameter, with conidial cells larger, some terminal, ovoid or pyrif orm, 3-4/a in diameter, with some larger intercalary ones 6-10.5/t (Carter's capsules). In cultures chlamydospores 4-5/a in diameter, yellowish gray, blackening the substrate, reproducing abundantly at 20°-25° C, but growing slowly and normally between 3° and 37° C, hyaline, septate. Conidiophores not in coremium, septate, 2-3 furcate with the next rank verti- ASPERGILLACEAE 641 cillate, umbellate, 50-60/x beyond bifurcation. Primary phialides 7.5-1.2/x long, secondary phialides splierico-depressed, 3.7/1. in diameter. Conidia spherical, 2.2-3. 7/A, smooth, hyaline, aeruginous greenish, never long catenulate. Growth on glycerol or Sabouraud agar from isolation, colonies round, slightly umbilicate, grayish white or gray, velvety, hyphae radiating from the middle, finally giving a hemispheric effect. On potato, bread, sugar car- rot, glycerol carrot, or orange, colonies ragged, floccose, moderately rugose or undulate, completely covering the substrate, at first white then, on appear- ance of fructifications, aeruginous green and mouse color. Reverse cream colored, with edge blackening in age. No odor. On pear and apple, an arachnoid colony is formed. Colony on gelatin is round, plane, slightly ochraceous. In bean decoction, with or without sugar or glycerol, growth is at first immersed, with the formation of a basal veil of hyphae, which are hyaline, septate, 1.5-2/x in diameter, with intercalary chlamydospores, the lat- ter ovoid, 8-12/A in diameter. Later there appears a supernatant pellicle with normal conidiophores and hyphae. Litmus agar is turned slightly alkaline. Gelatin not liquefied. Penicillium Giordano! Vuillemin, Champ. Parasit. 62, 1931. P. glaucum Giordano, Annali. Med. Nav. Colon. 24: 567, 1918, non aliorum. Producing fatal pseudotuberculosis. I have been unable to see a copy of this work. Penicillium minimum Siebenmann, Die Schimmelmykosen der mensch- lichen Ohres. 82, 83, 1889. Penicilliumdhnlich Bezold, Arch. Ohrheilk. 25: 1885. Isolated from a case of catarrh of the middle ear following scarlet fever. Three years later the organism was found abundantly fruiting in epidermis and a large, crouplike membrane. Mycelium 2/t in diameter. Conidiophores up to 20/a long, septa 6/t apart. Conidia 2.5-3/a in diameter and about the color of those in Aspergillus niger. The case cited by Buscino & Cardia appears somewhat doubtful. The fig- ures suggest an Aspergillus, although the organism was determined by Pollacci as P. minimum. Penicillium crustaceum [Linne] Fries, Syst. Myc. 3 : 407, 1832. Mucor crustaceus Linne, Species Plantarum, 1186, 1753. Monilia digitata Persoon, Syn. Fung. 693, 1801. Penicilliiim glaucum Link, Mag. Ges. Naturf. Freunde Berlin 3: 17, 1809. Greco, Origine des Tumeurs . . . 67-122, 1916, reports this species as the etiologic agent in 3 cases. He gives much detail, but absolutely no evidence that the organism cultivated was the etiologic agent or that the Penicillium isolated was this species. The cases were undoubted mycoses and showed sep- tate, occasionally binucleate mycelium but he was consistently unable to re- produce in animals with his cultivated organism either the lesions produced by 'inoculations with uncontaminated pus from the human lesions or the lesions as they appear in human subjects. 642 MEDICAL MYCOLOGY Penicillium Montoyai Castellani & Chalmers, Man. Trop. Med. ed. 1, G14, 1910. Penicillimn picior Neveu-Lemaire, Precis I'arasilol. 1908. Isolated by Montoya y Flores (1898) from a grayish violet pinta. Hyphae branch dichotomously. Spore chains with more or less cylindric sporophore. Penicillus short and has only a few branches. Conidia spherical or ovoid, smooth, 3-4.5/x in diameter. Organism grows well on ordinary agar and maltose agar. On glycerol agar, colony is woolly, short, kinky, white at first, becoming glaucous and finally violaceous, old cultures presenting rose-colored mammillae, with re- verse rose. Glycerol broth shows a pellicle with smooth white tubercles, broth becoming yellow as picric acid. Penicillium pruriosnm Salisbury, 1873, Marchand, Bot. Crypt, Ice. 1880. Isolated from mucous membrane of vulva and vesica urinaria. Penicillium quadrifidum Salisbury Zeitschr. Parasitenk. [Hallier] 4: 3, PI. 6, Fig. 11, 1875. Isolated from the blood under fairly sterile conditions; this organism was seen in abundance in the freshly removed blood. The patient recovered after medication with 2 gm. of quinine and 20 drops of tincture Peni-chloride in a glass of water every four hours. In addition, the swollen surfaces were painted with tincture of iron every 3-4 hours. Kesten et al. (1932) report Penicillium spicuUsponim Lehman from open ulcers on the toes, but were unable to reproduce the lesions on experimental animals. PAECILOMYCES Paecilomyces Bainier. Mycotheque de I'Escole de Pharmacie 11: Bull. Soc. Myc. France 23: 26, PL 7, 1907. Type species is P. varioti Bainier. Phialides short, tubular, or more or less enlarged, tapering into long conidiiferous tubes, curved or bent slightly away from the long axes of the phialides; variously arranged, partly in verticils and branching systems sug- gesting Penicillium, partly irregularly arranged upon short branchlets, partly arising singly along the fertile hyphae ; conidia in chains, never green ; macro- spores variouslj^ borne, usually solitary and terminal on branchlets either sub- merged or close to the substratum (Fig. 103). It is possible that some species described as Spicaria should be placed here. The nomenclature of Spicaria is involved, having been used in various senses by various authors, so that it seems wiser to retain Paecilomyces than to treat it as a synonym as was done by Oilman & Abbott (1927). Paecilomyces Burci (Pollacci) Thom, Penicillia 548, 1930. Penicillium Burci Pollacci, Atti 1st. Bot. R. Univ. Pavia 18: 128, 129, PI. 31, 1921; Berti, Policlinico Sez. Chiiiirg. 29: 484-486, Figs. 1-5, 1922; Campa- telli, Pensiero Med. 12: 217-219, 1923. ASPERGILLACEAE 643 This organism was isolated by Dorti from a small tumor along with Acre- moniella Berti in Florence, and was also used by Campatelli to produce an ex- perimental granuloma. Specialized (•onidio])horps scarcely found; penicilli consisting of single phialides or small, short-stalked clusters or verticils of phialides, each with its conidial chain, scattered along the separate hyphae or ropes of hyphae in large numbers; phialides 10-12/i with long tubes set at angles to their axes and divaricate Avhen in verticils ; conidia up to 6, 7 or even S/j. x 2.5-4/x, almost hyaline (Thom). Colonies in Czapek's solution agar broadly spreading, forming a loose, bristly network of hyphae, making a funiculose felt 200-400/* deep, white with more or less yellow in areas, and with yellowish drops or their residue, reverse in yellow orange shades. Fig. 103. — Paecilomyces Va7-ioti. C, conidia; CCr, germinating conidia; P, penicilli; T, section of a trailing hypha bearing phialides and various grades of penicilliferous branchlets. (After Thom 1910.) Colonies on glucose agar, white floccose, then grayish chestnut. Sterile hyphae branched, hyaline, septate, creeping or ascending, 6-7/i in diameter. Conidiophores erect, simple, septate, hyaline. 50-110/* long, sparsely branched above, ending in chains of conidia. Conidia spherical, rarely ellipsoid, smooth, pale, fuliginous, 4-5/* in diameter. On broth, colonies floating, similar to those on agar, SCOPULARIOPSIS Scopulariopsis Bainier emend. Thom, Penicillia 511, 1930; Bainier, Bull. Soc. Myc. France 23: 99-103, PL 9, Figs. 1-6, 1907. AcauUum Sopp, Vidensk. Selsk. Skrift. I. Mat. Naturv. Kl. 1912: 11:33, 42-46, 1912. The type species is Penicillium hrevicaule Saceardo. 644 MEDICAL MYCOLOGY Colonies never green, with aerial hyphae partly at least in trailing and anastomosing ropes or fascicles (funiculose) ; conidiophores very short or al- most wanting, commonly borne along the funiculose hyphae; conidial ap- paratus as in PenicilUum, consisting of varying aggregations of branches and phialides, at times reduced to single phialides scattered along aerial hyphae ; phialides tapering gradually from a basal tubular section or even the base itself toward a conidiiferous apex, or narrowly tubular without tapering, ab- jointing conidia from the apex ; conidia more or less pointed at the apex and truncate at the base with more or less thickened basal ring surrounding the basal germinal pore, with walls usually thickened and often variously marked or roughened (Fig. 104). Sopp describes perithecia as showing small but definite ostioles. The species appear as agents of decomposition after the usual green Penicillia have ceased to be active ; i.e., in the later stages of decay. Fig-. 104. — Scopulariopsis brevicaulis (Sacc.) Bainier. (After Thorn 1910.) While there is little real evidence that PenicilUum is ever pathogenic, there is plenty of evidence for several species of Scopulariopsis, at least as invaders of the nails and secondary invaders, if not the primary cause of gummata and other lesions. Key to Pathogenic Species Perithecia present after 3 weeks, black, 250/;i in diameter, asci evanescent, 8-spored, 10-12 x 8fi; ascospores brown planoconvex, 6-7 x 3-3.5fi. S. cinereus. Perithecia perhaps developing in time, but only rudiments observed, conidia smooth, color- less, 3-4 X 1.5-2M. S. Blochi. Perithecia usually absent (not yet observed). Conidia smooth. Conidia yellow to tawny, 5-7/i. 8. rufulus. Conidia hazel (avellaneous) 6-8,^. S. Koningi. Conidia brown (2-5/i). Phialides in groups. S. sehnsuohta. Phialides single. S. Bertaoini. ASPERGILLACEAE 645 Conidia white. Conidia 3-4 x 1.5-2ja. S. Blochi. Conidia 1.25-1.5/t. S. mmirrms. Conidia rough. Conidia chestnut, 6/t. S. ivorensis. Conidia white. Conidia ellipsoid, 5.2 x G.o/j., eehinulate. S. sputicola. Conidia subcitriform, 7-9^. S. Castellanii. Conidia some shade of yellow. Conidia canary yellow, 3-5/*. S. Mencieri. Conidia golden yellow, 3-4.5At. S. a/wreus. Conidia pale yellow, 7 x 6^. S. hrevicauUs var. hominis. Conidia yellowish gray chestnut, 5-8 x 4/i. S. venerei. Scopulariopsis cinereus Weill & Gauclin, Arch. Med. Exp. 28: 458-460, 1919. Found on infected toenails, which crumble into white layers with some yellower parts. Mycelium and conidia brown in age. Conidia have a truncate base and pointed disjunctor, which is slightly longer than broad, 2.5-3 x 4-5/a. Coremia sometimes present. Chlamydospores up to lOO/t, mostly terminal, sometimes intercalary. Perithecia develop in about 3 weeks at 25° C. They are spheri- cal, surrounded by radiating mycelium, taking a full month longer to mature. Asci ovoid, 10-12 x 8fi, 8-spored, early diffluent. Ascogonium soon surrounded by other hyphae which make up the pseudoparenchymatous wall of the peri- thecium. Ascospores brown, planoconvex, 3-3.5 x 6-7/a. Optimum tempera- ture 25° C. Grows on usual Penicillium media and on sterilized toenails. Colonies at first white, then ashy, mouse-gray, becoming brown green in age. On Sabouraud maltose, in Erlenmeyer flask, after 16 days, colony attains 3 cm. in diameter, is white, radiating, granular with a large central prominence. Green circle near the periphery and outside it a white circle which remains white. Scopulariopsis Koningi (Oudemans) Vuillemin, Bull. Soc. Myc. France 27: 143, 1911. Monilia Koningi Oudemans, Arch. Neerl. Sci. Exact. Nat. II, 7: 287, PI. 21, 1902. Scopulariopsis rufulus Bainier, Bull. Soc. Myc. France 23 : 105, PL 12, 1907. Vuillemin (1911) thinks this a synonym of Penicillium hrevicaule Saccardo as does also Biourge (1923), excluding Scopulariopsis rufulus Bainier, also Weill & Gaudin, 1919. Jannin (1912) isolated this from a gumma on the hand. Raymond & Parisot, 1916, isolated this organism from a number of cases of trench foot and gave a brief account of the lesions following experimental inoculations. Liegard, 1925, found it in ulcerated human eye. Originally obtained by Koning from humus or leaf mold in Holland. Koning describes his organism as follows : Colonies on soil extract gelatin orbicular, subzonate, rosy avellaneous; hyphae all hyaline, 4-5/a in diameter, septate. Creeping hyphae dichotomously branched, ascending hyphae race- 646 MEDICAL MYCOLOGY mosely branched with phialides lageniform, 30-40/a long, bearing simple chains of conidia with as many as twenty in a chain. Conidia snbspheric, apiculate at the apex, smooth, 6-8/i, in diameter, delicately rosy avellaneous. Scopulariopsis sehnsuchta Froilano de Mello, Bull. Soc. Patli. Exot. 25: 295, 296, 1932. A saprophyte isolated from lesions produced by Ectotrichophyton men- tagrophytes (Trichophytori asteroides). Hyphae hyaline, branched; 2.5-3/^, conidiophores septate, bearing- 2-8 phialides suggesting those of Paeciloniyces, 20-150 x 2.15-3;u,; metulae 5.5-7 x 1.5-2fi; phialides 3.5-11 x 1.5-3yu,; conidia smooth, brownish, slightly ovoid in chains of 10-40 cells, 2-5/a in diameter or 4 x 2.5/x. Sclerotia 56-88 x 68-108/*; coremia also present. On Czapek agar, colony velvety white, becoming chamois or golden yellow, more or less zonate, amber guttulate, reverse yellowish; later the colony be- comes chestnut to clear chocolate; reverse almost black. On potato, bread, and liquid media, similar colors, sometimes with a slight heliotrope shade. Scopulaxiopsis Bertacini Redaelli in Bertacini, Giorn. Ital. Derm. Sifilol. 75: 783-828, PZs. 1-9, 1934. Isolated from a fatal case of blastomycosis originating in Brazil, showing cutaneous ulcers mostly about mouth and neck with lesions in the lung, etc. Pathogenic for laboratory animals. In tissues, cells 5-22/* mostly 10-12/a; primary mycelium 3.5-4/*; secondary mycelium 1.5-3/*; chlamydospores intercalary 20-25/*; phialides erect, short, clavate, tapering toward distal end, 4 x 6-10/*; conidia smooth spherical or slightly ovoid, slightly brownish, 3.5-4/* in diameter in chains of 5-6 spores, often only a single spore remaining attached. Some phialides are branched from the middle of the club. On Sabouraud agar, colonies of compact yellowish white felt, glabrous, margin grayish white, in old cultures becoming mammillate, color varying from smoky gray to brownish, depending on substrate. On Pollacci agar, colony round, glabrous, moist, shining Avith small verrucose center which be- comes cerebriform, margin white. On liquid media, colony forms dense yellow brown ring, grayish above, finally becoming a thick pellicle. The relationship of this organism is not altogether clear. The colony on Pollacci agar suggests the imperfect Eremascaceae. Scopulariopsis Blochi (Matruchot) Vuillemin, Bull. Soc. Myc. Prance 27: 144-146, 1911. Mastigocladium Blochi Matnichot, C. R. Acad. Sci. 152: 325-327, 1911. Named for Bruno Bloch, who first isolated it [Arch. Derm. Syphilis 108: ^n-512, Pis. 19-21,1911]. Isolated from a case of gummatous lymphangitis, clinically resembling sporotrichosis. Mycelium 0.5-1.5/* in diameter, septate, little branched, tending to aggre- gate. Conidiophores erect, simple, with broad base but with slight constric- tion, 20-30/*, tapering as a long cone to a fine point, bearing, in a centripetal ASPERGILLACEAE 647 manner, indefinite heads of conidia whieh adhere in chains, are ovoid with unequal ends, 3-4 x 1.5-2/x, indefinite in number. In okl cultures there are Avhite, creamy formations which may be undeveloped perithecia. Spores brownish and terminated with a filiform papilla. On Sabouraud maltose, growth attains a diameter of 1-1.5 cm. in 8 days, is round, flat, with indentations on the periphery. Central portion pure white, periphery gray like gray rubber. Finally, center becomes cratoriform and the periphery develops radial folds. On glycerol agar slant, growth is more rapid, brighter, ivory in color, and with broader convolutions. On glycerol and potato, groAvth is good and about the color of the substrate with a surface of furry papillae, becoming snow white in the drier parts of the culture. Scopulariopsis minimus Sartory, Hufschmitt & Meyer, Bull. Acad. Med. Ill, 103: 604-606, 1930. Isolated from a case of onychomycosis. Mycelium slender, 1.25-2.5|a in diameter, hyaline, branched. Conidiophores 10 X l-2fi, often grouped in bundles. Phialides 3.75-4.5 x 1.25-1.5;u,, sometimes in groups of two to three. Conidia spherical, apiculate, 1.25-1. 5/^ in diameter. Perithecia not found. Optimum temperature 28°-32° C. Colony crateriform, small, white, after 21 days pale green, then deep green, then ashy with the margin remaining white. Scopulariopsis ivorensis Boucher, Bull. Soc. Path. Exot. 11: 308-321, 1918. Several lesions over bony areas on each of 16 cases in Africa. Cured by medication with KI. Lesions reproduced on pigeon and guinea pig from each ease encountered. Hyphae septate with cells of metacarpal form, 1/a in diameter. Conidio- phores short, irregular with long phialides. Conidia rounded or truncate, G/j. in long axis, brownish, ecliinulate, thick-walled. Live 5 months in culture. Gram-negative. All cultures reported for 25° C. in diffuse light. On poor Sabouraud agar, growth powdery on the third day, chestnut on the fifth, deep brown on the seventh. On simple glucose agar on the third, colonies filamentous and rounded with central point refringent, browning on the fourth day, becoming powdery on the fifth and chestnut colored. Giant colony is 5 mm. in diameter on the second day, 1 cm. on the third day, central zone white and filamentous, outer hyaline and radiate, 5 cm. in diameter on seventh day, on eighth day whole colony becomes chestnut colored, on tenth dry and powdery, chestnut by reflected light, black filamentous by transmitted light; growth stops the twelfth day. If several inoculations have been made, colonies are. not eon- fluent. On potato, second day white, filamentous colonies, third day dry, light brown membrane with white border which spreads. Sixth day, culture wholly chestnut. On potato glycerol, second day culture white and dry, third day light brown, fourth day folded, fifth day powdery, medium covered on the eleventh day. Growth is slower in the dark. In glucose bouillon colony shows white clots on the surface the second day, on the fourth day a floating chest- nut mat, elevated, with simple white border. 648 MEDICAL MYCOLOGY Scopulariopsis sputicola (Galippe) Dodge. Monilia sputicola Galippe, Jour, de Anat. 21 : 538-553, PI. 27, 1885. Isolated from human sputum, perhaps as a contaminant in the author's attempt to secure sterile sputum by filtration. Mycelium variable in diameter, branched, septate; phialides isolated or attached to a principal hypha. Conidia ellipsoid, in chains up to twenty-five, becoming- long, echinulate, scabrous. Spores catenulate on side branches, end ones oldest, 3.7-7.5/x in short diameter, the mean measurements being 5.26 x 6.36/A. Grows in white tufts of mycelium and white spores. Scopulariopsis Castellanii Ota & Komaya, Derm. Woch. 78: 163-165, 1924. Isolated, probably as a contaminant, in a case of tinea flava. Mice killed in 14-16 days after intraperitoneal injection, and organism recovered from peritoneum, but pathogenicity mild or doubtful. Hyphae about 5//. in diameter, with relatively thick septa, much branched. Fertile hyphae 3-4/a, unbranched or dichotomous. Conidiophore 2-40;u, (sic), tips thickened; conidia spherical or ovoid, or roughly ovoid, 7-9/t, also sub- citriform, echinulate. Scopulariopsis Mencieri Dodge, n. sp. Scopulariopsis sp. Sartory, Bull. Acad. Med. 81: 783, 784, 1919. Found in pus from a war wound. Pathogenic to rabbit and guinea pig. Mycelium hyaline, white, septate, sometimes narrowed at the septa, 2.5- 3.6/x in diameter. Fertile hyphae in fascicles, sporophores 10-60/a, disposed regularly along a hypha, not isolated at first by a wall. Conidia in chains with the insertion characteristic of Scopulariopsis, 3-5/x in diameter, canary yellow, echinulate or nearly smooth. Optimum temperature 28°-30° C, maxi- mum 40°-41° C. Growth good on potato, carrot, Jerusalem artichoke, banana, agar, gelatin, ordinary Kaulin and sugar broths. Only glucose and maltose are attacked, the latter hydrolyzed. Milk is coagulated the tenth day, and the coagulum peptonized. Gelatin liquefied in 5 days. Scopulariopsis aureus Sartory, Champ. Parasit. Homme Anim. 680, 681, 1922. Scopulariopsis sp. Sartory, C. R. Acad, Sci. 167: 703, 704, 1919. Found as a parasite on human nails. On nails appears as irregular, septate hyphae, 2.5-9 or 10/x with terminal or intercalary chlamydospores, 20-35/a in diameter, occasional conidia. In cul- ture mycelium white, then golden yellow. Hyphae 0.5-1.4/i in diameter, very much branched and having a tendency to aggregate conidiophores. Phialides erect, sometimes differentiated, tapering at the tip with spherical conidia, '^ aureus," usually ornate, 3-4. 5/a in diameter. On gelatin at 22°, conidia are sometimes smooth and hyaline, especially in old cultures. Chlamydospores sometimes intercalary and on maltose agar reach diameter of 130/*. Optimum temperature 29°-30° C, growth ceasing at 39° C. ASPERGILLACEAE 649 Organism grows on Raulin maltose agar, potato, acid potato, glycerol potato, carrot, Sabouraud nutrient gelatin, Raulin 's solution with sugars. No growth on egg albumen or coagulated beef serum, hence the latter not lique- fied. Milk not coagulated in 40 days. Gelatin liquefied in 6 days. Scopulariopsis brevicaulis (Saccardo) Bainier, var. hominis Brumpt & Langeron apud Brumpt, Precis Parasitol. ed. 2, 902-905, 1913. Penicillium hrevicaule Saccardo, var. hominis Brumpt & Langeron apud Brumpt, Precis Parasitol. ed. 1, 838-840, 1910. Isolated from onychomycosis. In nails, mycelium septate, 2-10/a in diam- eter; chlamydospores both terminal and intercalary, lO-30/i,. Weil & Gaudin (1919) report conidia in the nails. The parasitized portions of the nails are a dull yellow brown. Hyphae on the surface of medium in tufts, aggregated, also in ropes ; conidia cafe-au-lait, ornate, spherical to citriform. Colonies on potato glycerol, sweet potato, carrot, Sabouraud 's glucose and maltose agars, velvety. Scopulariopsis venerei Greco, Origine des Tumeurs. 709-717, PL 21, Figs. 437-440, 1916. Isolated from cases of venereal granuloma. Mycelium abundant, septate, 2-5/x in diameter; chlamydospores ( ?) present; conidiophores short, 10-20/* long, arranged on main hyphae to give a verticillate appearance ; phialides slightly conical or blunt ; conidia 5-8 x 4/i with thick, outer wall spherical or citriform, with the surface showing yellowish refractive drops that give a rugose appearance. Colonies slightly yellowish grayish chestnut to chestnut, with yellowish radiate spots, spreading, with whitish gray ropes at the center and in striae, margin light gray. Scopulariopsis lingualis Neto & Martins, C. R. Soc. Biol. 106: 1179-1181, 1931. Isolated from lesions on human tongue, case not described here. Mycelium slender, unbranched, septate, sometimes branching dichoto- mously at the tips. Conidiophores either simple, often reduced to a single conidium, or branched, each branch producing a single conidium. Conidia 1-2/x in diameter, spherical or ovoid. On serum glucose, growth appears on third day, 1 cm. in diameter, on the eighth, irregular, velvety, rugose, slightly brownish. Same on carrot and maltose serum. On potato, growth slow and not characteristic. On Gorod- kova agar, small colony, 3-4 mm. in diameter, yellowish, irregular, surface rugose apiculate, no tendency to spread over the surface. On gelatin, growth very slow, slight whitish membrane on twentieth day, no liquefaction. In Raulin 's liquid, clear flocculent precipitate. Scopulaiiopsis d'Agatae (Saccardo) Dodge, n. comb. Oospora d'Agatae Saccardo apud d'Agata, Policlinico Sez. Chirurg. 25: 80-87, 1 pi., 1918. 650 MEDICAL MYCOLOGY Sporendonema epizoum Ciferri & Redaelli, Jour. Trop. Med. Hyg. 37: 167- 170, 1934; Redaelli & Ciferri, Atti 1st. Bot. R. Univ. Pavia IV, 5: 145-198, 1934. Isolated from small, indolent nodules which become ulcers with blackish crust, 1.5 cm. in diameter. Nonpathogenic to guinea pigs, pathogenic to white rats; lesions similar to those produced by Sporotrichum. It is possible that some of the Italian cases referred to Hemispora stellata should be referred here. Cerri (1932) identifies this organism with Torula Sacchari Corda. Hyphae septate, branched, 2.5-3/i. in diameter, forming a mat 90-100/i thick ; conidiophores erect 50-60/x high, densely fasciculate, flexuous, sparingly branched, 2.5-3.5/a in diameter, 50-60/i, high at first, becoming 150-160/a, hyaline or subhyaline, minutely guttulate, sparingly septate, chains of conidia, 6-8 cells, developing slowly in the phialides and hanging together for a long time ; conidia spherical 2.5-3.5/* smooth, uniguttulate, ochraceous at first becoming olivaceous; chlamydospores intercalary, hyaline, 8-lO/t. On Sabouraud glucose agar at 22° C, colonies grayish at first, becoming mammillate, grayish brown with brownish or chocolate powder, finally cere- briform and confluent. On glucose broth, chocolate brown, powdery, super- ficial, brown sediment but no true pellicle. Doubtful Position The following species are either too imperfectly described to place ade- quately or I have been unable to locate the original descriptions. Scopulariopsis Vignolo-Lutatii (Matruchot) Dodge, n. comb. Acaulium Yignoli-Lutatii Matruchot apud Vignolo-Lutati, Arch. Derm. Syphilis 118: 681-698, PI. 21, 1 fig., 1913. Isolated from lesions on skin following wound with acacia thorn, Novara Province, Italy. Lesions clinically suggest sporotrichosis or tuberculosis. Pathogenic for guinea pigs. Mycelium of branched, hyaline, septate hyphae 2-3fx in diameter, not fasciculate. Phialides simple or in groups of two or three, 10-15 x 4/x ; conidia in chains of 8-10, blackish brown. Colony on Sabouraud glucose or maltose agar more or less cerebriform, center becoming very dark brown, outer zone yellow. Scopulariopsis americanus Ota, Jap. Jour. Derm. Urol. 28: 4: [7], 1928, nom. nud. [figured and perhaps described in the Japanese text] ; Ota & Kawatsure, Arch. Derm. Syphilis 169: 160-163, Figs. 4-6, 1933 [citing Ota, Jap. Jour. Derm. Urol. 26: 1:111, 751, 1926, as place of original publication, but I have been unable to see this publication]. American oidiomycosis. Cultures of Weidman 1135, 1136, 1233, and 1234 (Ota collection 538, 539, 540, 541). Kawatsure (1933) reports pathogenicity for mice, white rats, guinea pigs, and rabbits. Mycelium produces irregular arthrospores [suggesting those of Madur- ella] • conidia in chains of 3-4 spores on short side branches or more often ASPERGILLACEAE 651 singly, but never in typical phialides, smooth and hyaline at first, then cream yellow, verrucose, 5-6/a in diameter. On peptone and peptone glucose agar, colony circular moist, cream gray, with radial folds, then producing numerous cervine coremia, becoming dull, dry with concentric rings of velvet, and finally powdery. From the foregoing brief description it is evident that the species is not closely related to Scopidariopsis but is probably closely related to some of the species of Zymovema, although Kawatsure's Fig. 6 Bh with its verrucose eonidia is suggestive of Hemispora. As the description is based on four strains, which show some differences in pathogenicity and no reference is given to the case histories, it is not possible to place this organism definitely. Scopulariopsis Fincki A. & R. Sartory, 1925 ; Vuillemin, Champ. Parasit. 65, 1931. Scopulariopsis Mottai Vuillemin, Champ. Parasit. 62, 1931. ? PenciUium epigaeum Motta, Atti Clin. Ota-Rino-Laring. 25: 305-322, 1927, non Berkeley & Curtis. PHAEOSCOPULARIOPSIS Phaeoscopulariopsis Ota, Jap. Jour. Derm. Urol. 28: 4: [7], 1928. Characters of Scopulariopsis, but spores brown to black, producing chains of spores on phialides. Phaeoscopulariopsis Paisi (Pollacci) Ota, Jap. Jour. Derm. Urol. 28: 4: [7], 1928. Torula Pais Pollacci, Atti 1st. Bot. R. Univ. Pavia 18: 129, 130, PI. 31, 1921. Found in axillary hair in Sassari. Colony on glucose agar black, verrucose, with lanuginous margin. Hyphae sterile, sparsely septate, hyaline, and slender, with pale ochraceous and thicker tips, 4-6/i, in diameter. Conidiophores short, sparingly septate, olivaceous, 4-12/x long. Conidia spherical, smooth, eatenulate or glomerate, brown, 4-5//, in di- ameter. Phaeoscopulariopsis Bestae (Pollacci) Ota, Jap. Jour. Derm. Urol. 28: 4:[7], 1928. Torula Bestae Pollacci, Riv. Biol. 4: 313-318, 1 fig., 1922. Isolated from subcutaneous abscesses in a patient suffering also from high fever, gastrointestinal disturbances, and bronchopulmonary trouble. On lanc- ing, the lesions produced bloody, purulent matter. Similar smaller foci of in- fection developed. Trichosporium Mantegazzae also isolated from the same lesions. Sterile hyphae repent, branched, filiform, pale at first then rosy or pale avellaneous, 5-6.5/x in diameter. Conidiophores short, septate, scarcely distinct from sterile hyphae, 6-20/x, long, apex rounded. Mature conidia spherical, smooth, rose colored at first, then avellaneous, 8-IO/1, in diameter, shortly eatenulate. 652 MEDICAL MYCOLOGY On glucose agar, 15°-20° C, colony circular, pale at first, floccose, then covering the medium, rose with more or less white margin, ragged, then avel- laneous ; not liquefying gelatin. ALLESOHERIA Allescheria Saccardo & Sydow, Syll. Fung. 14: 464, 1899. Eurotiopsis Costantin, Ann. Inst. Pasteur 11: 1-43, 1897, non Karsten, 1889. Eurotiella Lindau in Engler & Prantl, Die Naturl. Pflan^enfam. I, 1 : ** : 383, 1900. The type species is Eurotiopsis Gayoni Costantin. Perithecia spherical, minute, thin-walled ; asci spherical, ascospores ovoid, pointed at the ends. Conidia in chains at the tips of sporophores but compressed as in Cephalosporium. So far known only from a single case of mycetoma, with white grains. Allescheria Boydii Shear apud Boyd & Crutchfield, Amer. Jour. Trop. Med. 1: 258-268, 1921 ; Mycologia 14: 242, Figs. 1-3, 1922. Isolated from the foot of a male negro farm laborer in Texas. Thorn in foot near metatarsophalangeal articulation twelve years before. Thorn re- moved and wound healed. Three months later patient felt a stinging pain in the ankle, followed by swelling. At intervals soft openings appeared in the side of the ankle which discharged pus and healed in a few weeks. Process has not extended, remaining localized for many years and showing white grains. Ankle swells when patient is on his feet long. Perithecia numerous, crowded, covering the surface of the medium, usu- ally erumpent or subsuperficial, spherical, thin-walled, dark brown, astoma- tous, 100-200/i in diameter; asci spherical or subspheric, thin-walled, evanes- cent at maturity, 10-20/x in diameter ; no paraphyses ; ascospores 8, spherical to subspheric or somewhat ovoid, continuous, smooth, pale yellowish brown when mature, spherical form about 7/* in diameter, other mostly 5.5-7 x 4-4.5/a. Pycnidia wanting or unknown. Cephalosporium Boydii, byssoid, thin, floccose, white at first, soon gray, margin radiate-fimbriate, later changing to pale greenish ochraceous as sporu- lation begins, fertile hyphae much branched, spreading, conidia adhering in small or large subspheric masses, continuous, subspheric to oblong ellipsoid, very variable in size and shape, hyaline at first, becoming pale, yellowish brown when old, smooth, 8-15 x 4-7.5//,, mostly 10-12 x 5-6/jl. Coremia (Dendrostilhella Boydii) with dark brown synnema very variable in height and thickness, 200-300/* or more high, head subspheric ; conidiophore alternately branching, ultimate branches once or twice the length of the conidia; conidia practically the same size, shape, and color as in the byssoid condition and adhering in a globular mass after abstriction. Organism was cultivated from granules on plain, Sabouraud, Huntoon, and glucose agar. No bacteria found. Huntoon 's agar is best. In 14 days, ASPERGILLACEAE 653 colony 5 cm. across with four zones, center 3 mm. with alternate light gray and darker brownish gray bands with smaller, dark coremia. On Saboiiraud agar, colonies smaller and zoning less noticeable, pale grayish green with white border. No fructifications seen. Perithecia formed on Sabouraud agar and potato, 200/x in diameter. Perithecia cespitose on potato glycerol. Organism is strongly aerobic, showing very little growth in liquid media, but a good pellicle is formed. Carbohydrates not fermented. Milk and Loeffler's serum are peptonized. Blood not hemolyzed but in agar, gradually becomes green. Gelatin liquefied. ONYGENACEAE The members of this family are saprophytic but confined to animal sub- stances, such as hoofs, horns, claws, feathers, etc. They are included here, since it is possible that eventually some members may be found parasitic. When the family is studied more fully, it is possible that it may show resem- blances to the dermatophytes, AcJiorion, or to some genera of Aspergillaceae which seem to show some specialization on nails, e.g., Scopulariopsis. Onygena equina is the only species studied carefully. The fructifi- cations are up to 1 cm. in height. They consist of solid homogeneous coremia which abjoint, on their surfaces, so many chlamydospores that they seem to be covered with a brown powder. Later each is differentiated into a solid stipe, composed of parallel hyphae and a somewhat looser head, consisting of radiating hyphae. On the outer surface of the head, the hyphae intertwine to form a firm, pseudoparenchymatous peridium. The interior hyphae de- velop into capillitium and aid in the dissemination of spores. Nothing is known of the cytology of this species. Within the head in various places, tAvo short, septate branches coil about each other into a solid knot. At maturity, the cavity of the head is filled by a dark spore mass through which run the capillitium threads, generally starting at the base. The peridium is ruptured irregularly, or around the base of the head, and the spores are scattered. They germinate after a resting period, which may be shortened by placing the spores in a mixture of HCl and pepsin, similar to that in the gastric juice. Immature ascospores and chlamydospores germinate without this stimulation (Ward 1899; Brierley 1917). 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Dissertation on aspergillosis. The Victoria Univ. Manchester, England, 66 pp. — . 1901. Experimental aspergillosis, Jour. Path. Baot. 7: 34-52, PI. 1. Sabouraud, Raimond. 1898. Les carates, Ann. Derm. Syphiligr. 9: 673, 674. Salisbury, J. H. 1875. Vegetations found in the blood of patients suffering with erysipelas, Zeitschr. Parasitenlc. 4: 1-5, PI. 1. Sartory, Antoine. 1916. Mycose a Scopulariopsis Koningi, Progres. Med. 31: 107, 108. — . 1919. Un champignon pathogene nouveau du genre Scopulariopsis isole d'un pus de plaie de guerre. Bull Acad. Med. [Paris] III, 81: 783, 784. ■ — . 1919. Onychogryphoses et onychomycoses, Bull. Acad. Med. [Paris] III, 82: 162, 163. — . 1919. £tude d'un Aspergillus du groupe fumigatus, Aspergillus fumigatus var. mini- mus. Bull. Acad. Med. [Paris] III, 82: 304, 305. — . 1919. Sur un nouveau champignon du genre Scopulariopsis isole d'un cas d 'onycho- mycose, C. B. Acad. Sci. 169: 703, 704. — . 1920. 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ASPERGILLACEAE 663 Schmidt, Albert. 1897. tJber die Bediiigungen der Conidien-Gemmen und Schlauchfruclit- production bei Sterigmatocystis nidulans, Inaug. Diss. Greifswald, 40 pp., 2 fls. Schonlierr, August. 1932. Ueber die Schimmelpilzerkrankung der menschlichen Lunge, Inaug. Diss. Med. Fak. Univ. Heidelberg, 39 pp. Schubert, Paul. 1884. Zur Casuiatik der Aspergillusmykosen, Devtsch. Arch. Klin. Med. 36: 162-179. — . 1889. Fadenpilze in der Nase, Berlin. Klin. Woch. 26: 856, 857. Sc.hiitz. 1884. Tiber das Flindringen von Pilzsporen in der Athmungswege und die dadurch bedingten Erkrankungen der Lungen und liber den Pilz des Hiihnergrindes, Mittheil. K. Gesundheitsamte 2: 208-227. Schwartz, Georg. 1904. Ein operatic behandelter Fall von Pneumomykosis aspergillina, Zeitschr. Klin. Med. 56: 120-127. Schweizer, A. 1927. tJber aegyptische Splenomegalie, Schweiz. Med. Woch. 57: 1017-1027, 7 figs. Scott, H. H. 1928. Reports on deaths occurring in the society's gardens during the year 1927, Proc. Zool. Soc. London 1928: 81-119. Segal, J. 1923. Notes on a fungus isolated from guinea pigs inoculated with the virus of typhus fever. Jour. Path. Bact. 26: 156-163, PI. 2S. Shear, Cornelius Lott. 1922. Life history of an undescribed Ascomycete isolated from a granular mycetoma of man, Mycologia 14: 239-243. Siebenmann, F. 1883. Die Fadenpilze Aspergillus flavus, niger, fumigatus; Eurotium repens (und Aspergillus glaucus) und ihre Beziehung zur Otomycosis Aspergillina, Zeitschr. Ohrenheilk. 12: 124-161. — . 1889. Ein zweiter Fall von Schimmelmykose des Eachendaches, Monatsher. Ohrenheilk. 23: 73-76. — . 1889. Die Scliimmelmykosen des menschlichen Ohres. Wiesbaden ed. 2, 118 pp., 4 pis. Silva, Flaviano. 1926. Consideragoes en torno de um caso de "puru purii, " Brasil Med. 40: 2: 113-119, 3 figs. Simon. 1922. Aspergillose saprophyte des brouches. Anevrysme aortique, P,ev. Med. de I'Est. 50: 339-342. Sion, V. & N. Alexandrescu. 1908. Sur la toxicite d'un type d 'Aspergillus fumigatus isole du mais avarie. C. B. Soc. Biol. 64: 288, 289. Solmersitz, F. 1906. Beitrag zur Aspergillus Mykosis der menschlichen Lunge, Deutsch. Med. Woch. 32: 1490-1492. Spegazzini, Carlos. 1925. Un nuevo Aspergillus patogeuo, Physis Bev. Soc. Argentina dent. Nat. 8: 115-117. Steele, Albert E. 1926. A case of infection with Aspergillus versicolor, Boston Med. Surg. Jour. 195: 536-538. Steudener, F. 1870. Zwei neue Ohrpilze nebst Bemerkungen iiber die Myringomycosis, Arch. Ohrenheilk. 5: 163-168, PI. 1. Stokes, W. R. & S. McCleary. 1928. A case of pulmonary mycosis, Boston Med. Surg. Jour. 197: 1350-1353, 3 figs. Talice, Rodolfo V. & Juan E. Mackinnon. 1929. Penicillium Bertai n. sp. agent d'une mycose broncho-pulmonaire de I'homme, Ann. Parasitol. Rum. Comp. 7: 97-106. — . 1931. Aspergillus (Eurotion) montevidensis n. sp. aislado de una otomicosis de hombre. Sociedad de Biol, de Montevideo, 61 sesion, 4 pp. — . 1932. Tres casos de otomicosis por Aspergillus observados en Montevideo, Beunidn Soc. Argent. Patol. Beg. del Norte, Tuoiomdn 7: 278-284. Tourmanoff, K. 1928. Au sujet de I'aspergillomycose des abeilles, C. B. Acad. Sci. 187: 391-393. Ureta, Eduardo. 1924. Pinta or carate with special reference to treatment, Internet. Conf. Health Prob. Trop. Amer. United Fruit Co. 524-532. Vignolo-Lutati, Carlo. 1913. Ueber eine neue Mykosis (Aeauliosis), [tr. J. Ullmann], Arch. Derm. Syphilis 118: 681-698, PL 21. 664 MEDICAL MYCOLOGY Vuillemin, Paul. 1904. L 'Aspergillus fumigatus est-il connu a I'etat ascospore? Arch, de Parasitol. 8: 540-542, — . 1911. Difference fondamentale entre le genre Monilia et les genres Scopulariopsis, Aerosporium, et Catenularia, Bull. Soc. Mye. France 27: 137-152. — . 1920. Fructifications de champignons decouverte dans I'ongle par Louis Jannin, C. B. Acad. Sci. 170: 788-790. — . 1927. Sartorya, nouveau genre de Plectascinees angiocarpes, C. B. Acad. Sci. 184: 136, 137. Watjen, J. 1928. Zur Kenntnis der Gewebsreaktionen bei SchimmeLmykosen, Arch. Path. Anat. Physiol. [Virchow] 268: 665-685, 3 figs. Wehmer, Carl. 1901. Die Pilzgattung Aspergillus, Mem. Soc. Phys. d'Hist. Nat Geneve 33: 2d. part. No. 4, 160 pp., S pis. — . 1903. Das Aspergillus des Tokelau, Centralbl. BaTct. I, 35: 140-146, 8 figs. — . 1913. Uebergang alterer Vegetationen von Aspergillus fumigatus in Riesenzellen unter Wirkung angehaufter Saure, Ber. Deutsch. Bat. Ges. 31: 257-268, 7 figs. Weichselbaum, A. 1878. Eine Beobachtung von Pneumonomykosis aspergillina, Wien. Med. Woch. 28: 1289-1292. Weidman, Fred. D. 1915. A Penicillium (species?) superimposed on a tuberculous peri- tonitis and pleurisy, Proc. Path. Soc. Philadelphia 17: 62. — . 1920. Penicillium brevicaule var. hominis (Saccardo 1877) Brumpt & Langeron 1910 in an American case of ringworm of the toes, Arch. Derm. Syphilol. 2: 703-715. Weill, P. £mile, Paul Chevalier & P. Flandrin. 1928. Contribution a 1 'etude des pseudo- sarcomas spleniques, Le Sang 1: 361-373, 5 figs. Weill, P. £mile, Paul Chevalier, Raymond Gregoire, & P. Flandrin. 1928. La splenomegalie primitive aspergillaire, Le Sang 1: 509-609, 1 pi., 6 figs. Weill, P. flmile & L. Gaudin. 1919. Recherches sur les onychomycoses, C. B. Soc. Biol. 82: 121, 122. — . 1919. Contribution a 1 'etude des onychomycoses a Penicillium, k Scopulariopsis, k Sterigmatocystis, a Spicaria, Arch. Med. Exp. 28: 452-467, PI. 12. Weill, P. £mile, Raymond Gregoire & P. Flandrin. 1927. Splenomegalie mycosique, Bull. Mem. Soc. Med. Hop. Paris III, 51: 713-717. — . 1927. Anatomic pathologique de la splenomegalie mycosique, Ann. Anat. Path. Anat. Norm. Med. Chirurg. 4: 587-593, 1 pi. Weiss, Pedro. 1930. Sur un Sterigmatocystis agent d'une dermatose semblable au Pityriasis versicolor flava. S. cinnamominus n. sp., Ann. Parasitol. Eum. Comp. 8: 189-193, 5 figs. Wheaton, S. W. 1890. Case primarily of tubercle in which a fungus (Aspergillus) grew in the bronchi and lung, simulating actinomycosis, Trans. Path. Soc. London 41: 34-37, PI. 2, Fig. 1. Winfield, James McF, 1897. A favus-like eruption of the oral mucous membrane caused by Aspergillus nigrescens. Jour. CtU. Genito-Urin. Dis. 15: 13-17, 3 figs. Wreden, Robert. 1867. Recherches sur deux especes de vegetaux parasites (Aspergillus flavescens et Aspergillus nigricans) de I'homme, C. B. Acad. Sci. 65: 368-371. — . 1874. Die Myringomycosis Aspergillina in den Jahren 1869-1873 nach eigenen und fremden Beobachtungen besprochen, Arch. Augen-Ohrenheilk. 3: 56-90, PI. C, Figs. 1-4. Xalabarder, X. 1927. Micosis pulmonar producida por una especie de Penicillium, Bev. Med. Barcelona 8: 574-577. Zarnirko, C. 1891. Aspergillusmykose der Kieferhohle, Deutsch. Med. Woch. 17: 1222. Ziegenhom, Otto. 1886. Versuche iiber Abschwachung pathogenen Schimmelpilze, Arch. Exp. Path. PJharm. 21: 249-268. Ziirn, A. 1876. Beitrage zur Lehre von den durch Pilze hervorgerufenen Krankheiten der Hausthiere, Arch. Wiss. Prakt. Thierheilk. 2: 110-114, PI. 2. CHAPTER XVIII FUNGI IMPERFECTI The many fungi whose life cycle is little known are usually grouped as Fungi Imperfecti. From its origin, it is obviously a large artificial group of more or less unrelated species. In this work I have removed from the Fungi Imperfecti all those genera where there seemed reason to believe that they were closely related to other genera whose whole life cycle is known; e.g., the group usually called Monilia by medical men has been removed to the Eremascaceae Imperfectae, although it is quite possible that when the life cycles of these species are better known, some may be found to belong else- where. I have placed the dermatophytes in an appendix to the Gymnoascaceae, although only one species has yet been found to produce asci. Similarly the species of Aspergillus and Scopulariopsis, in which no perithecia have yet been demonstrated, have been placed in the Aspergillaceae. Even after all these groups have been removed, there remain several important genera whose rela- tionships with the Ascomycetes are still obscure. It is possible that many of these species represent degenerated lines in which asci never will be produced, but in the last 50 years so many species have yielded to improved cultural technic and media that it seems probable that eventually many more may be placed. Hence, the Fungi Imperfecti are conceived as a temporary dumping ground until the gaps in our knowledge have been filled. The older authors attempted to divide the group on the basis of color and then septation of spores. They divided the Fungi Imperfecti into several groups on the basis of the elaborateness of the fructification in which the conidia were borne. Of these, only those groups in which the spores are borne in coremia or on more or less differentiated conidiophores without protection from sterile tissues need be considered here. For our purposes, even the presence of coremia has little importance. The Hyphomycetes, in which we are interested, were next divided upon the basis of color of spores into the Mucedineae with light colored or hyaline spores and the Dematieae with black spores. While this differentiation is often useful, it is very artificial and difficult to apply in certain well-defined groups. Further division was based on the conidiophore and septation of the spore. This system culminated in the complete and elaborate works of Saccardo, and of Lindau and his German coworkers. Even with this system, the Mucedineae with unicellular spores and undifferentiated conidiophores were not well studied, yet it is in these groups that the genera of most interest to the medical man are to be found. In France, largely due to the work and influence of Vuillemin, the dif- ferences between blastospore, arthrospore, and conidium in the narrower sense have been emphasized. To him we also owe the concept of phialide as a highly 665 666 MEDICAL MYCOLOGY specialized conidiophore. The various spores of Vuillemiu undoubtedly rep- resent different structures, although his distinctions are somewhat difficult to apply. Groups built up on such characters have much more unity than those based on the lumping of the older Saccardian system. Unfortunately Vuillemin was not primarily a systematist, and there has been no systematist among his followers, so that he failed to give us a carefully worked out clas- sification. In his last publications he was very reactionary and tried to undo the good work he proposed earlier. In this book I have already considered the genera with blastospores, arthrospores, and light colored mycelium in connection with various families of the Endomycetales and Plectascales (Gymnoascaceae) ; all of the conidial genera with spores borne in chains from phialides in the Plectascales (Asper- gillaceae). This leaves the genera with blastospores and arthrospores having black mycelium (the Toruleae) and those not bearing conidia in chains from Fig. 105. — Alternaria sp. (After Hopkins, Benham & Kesten 1930.) phialides to be treated here. In the following key I have attempted to include all groups in which no ascospores (perfect stage) are known, with page refer- ences to where each group is discussed further. It is felt that, in general, each of these groups is relatively uniform. The tabulation is somewhat arbitrary, but an attempt has been made to arrange the remaining groups in the order of increasing complexity of conidiophore, following tradition in placing groups with multicellular spores at the end of the series. Key to Larger Grroups of Fungi Imperfecti — Hyphomycetes No differentiation of sporophores, multiplication by sprouting or arthrospores (oidia). Cells mostly isodiametric or nearly so, occasionally adhering in chains, but never forming a true filamentous mycelium. Saccharomycetaceae Imperfect ae (p. 325). Cells mostly much longer in one diameter, forming a definite mycelium on most media. Colonies not brown or black, producing arthrospores, often multiplying by sprouting. Eremascaceae Imperfectae (p. 186). FUNGI IMPERFECTI 667 Colonies brown or black, arthrospores and chlamydospores present, sprouting not re- ported or rare. Toruleae (sensu Persoon et Saccardo) (p. 669). Sporiferous hypliae usually highly differentiated, at least other spore forms besides arthro- spores and chlamydospores present. Mycelium very slender, mostly less than Ifi, spores in chains, often borne in elaborate helices,* etc., colony usually chalky, powdery and very slow growing. Actinomycetale.s (p. 6!t4). Mycelium much coarser, spore chain.s, if present, not forming helices. Spores unicellular. Spores either sessile or on short sterigmata borne on undifferentiated vegetative hyphae. Deep cutaneous lesions. Sporotricheair (p. 786). Confined to the horny layer of the epidermis, hair and nails. Trichophytoneae (Gymnoascaceae Imperfecta^) (p. 433). Spores borne on well-differentiated fertile hyphae. Spores produced endogenously at the tips of pliialides. Spores typically in unbranched chains, not embedded in a gel. Pliialides hyaline. Aspergillaceae (p. 608). Phialides dark colored, conidia usually long cylindric, not spherical or isodiametric. Chalareac (p. 822)- Spores embedded in a gel at the tip of the phialide; chain structure rarely evident if present. Mycelium light colored. Ccphalosporieae (p. 823). Mycelium black, mouth of phialide dilated. PhialopJwreae (p. 833). Spores borne in groups on intercalary swollen cells along the vegetative hyphae. Gonatohotrytideae (p. 834). Spores borne singly, rarely in pairs, on variously branched conidio- phores, never from phialides. Spores hyaline or light colored. Mycelium usually lignt colojed, very rarely fuscous. Spores borne singly on tips of short branches. Botrytideae (p. 835). Spores borne in whorls. Trrticillieae (p. 841). Mycelium dark colored, brown to black. Spores in heads. Stachylideae. Spores borne singly. Chloridene. Spores colored, dark brown to black, mycelium often dark colored. Spores in chains, with youngest spore at the tip of the chain, not at its base as in the Aspergillaceae and Chalareae. IJaplographieae (p. 843). Spores in whorls. Arthrineae. Spores in heads. Perico'iiieae (p. 850). •Spore chains are extremely fragile so that a careless mount gives the appearance of a bacterial smear. For methods of making satisfactorj' mounts, see p. 702. 668 MEDICALi MYCOLOGY Spores borne singly on tips of conidiophores. Conidiophores unbranched. Monotosporieae. Conidiophores branched. Trichosporieae. Spores two-celled. Spores hyaline. Hyalodidymeae (p. 851). Spores Colored. Phaeodidymeae. Spores multicellular. Confined to the horny layer of the epidermis, hair, rarely on nails. Trichophytoneae ( Gymnoascaceae Imperfectae) (p. 433). Not isolated from such situations; several groups not known to be pathogenic although some, as Alternaria (Fig. 105), Stemphylium, Fusarium, etc., may be serious lab- oratory contaminants; in the case of the latter there are two or three cases where the fungus had multiplied on the host and caused irritation at least. Several species from these groups are suspected of producing allergy. (For re- view of this literature see Brown, 1932.) CHAPTER XIX TORULEAE Toruleae Saecardo, Syll. Fung. 4: 235, 247, 1886. The type genus is Torula Persoon non Turpin, Pasteur, Hansen, etc. Mycelium usually dark colored, often black ; spores usually dark colored ; no differentiation of sporophores; multiplication by arthrospores and chlamy- dospores, rarely sprouting. In this group I have combined the Coniosporieae and Toruleae of Sac- cardo. The Coniosporieae are a small group of species with more or less evanescent mycelium bearing single, terminal black spores of variable shape. The morphology is little known, but I assume that the spores are really arthrospores, or chlamydospores liberated by the disintegration of the hyphae which bear them, not by any specialized mechanism, as in the case of conidia. The Toruleae of Saccardo in a narrow sense include all the dark colored fungi with blastospores or arthrospores, hence the spores are usually in chains. To this group I have also added Madurella, a genus whose morphology is not well known. The cells are usually dark colored, in many ways suggesting a Hormiscium but apparently not normally producing arthrospores. In cul- tures, in my experience, it continues to produce sterile hyphae, forming hard, sclerotioid masses very adherent to media and hence very difficult to transfer, while Hormiscium produces moister and softer colonies. There are comparatively few pathogenic species in this group, which is predominantly saprophytic on decaying wood and other vegetable matter. It is quite possible that some species reported as pathogens were contaminants In the case of Cladosporium, it has been found frequently enough to justify its inclusion as having pathogenic species. Some would make it the type of another tribe, the Cladosporieae, since it regularly has 2-spored cells. So many confusing transitional states have been reported that it seems better for present purposes to leave it in the Toruleae. In Madurella, there are many species mostly isolated from black grained mycetomata, and in Indiella, those from light grained ones. In general, light grains are produced by species of Actinomyces, but occasionally Indiella is found instead. The problem of generic names is complicated somewhat as in the im- perfect yeasts, especially by the use of the name Torula by Turpin, Pasteur, Hansen, and even Guilliermond for asporogenous yeasts. Historically this usage is altogether incorrect. A discussion of the early application of the older generic names in this group follows. 669 670 MEDICAL MYCOLOGY DEMATIUM Vematium Pcrsoon, Neues Mag. f. Bot. 1: 121, 1794. Persoon first described this genus in 1794 reprinted with additional notes as Dispositio Meth. Fung. 41, 1797, as follows: " Filis subfasciculatis erectis pulverulentis. " He divides it into two sections: Rigida subfasciculata which includes D. mirev/m, and D. ai-ticulatum, on Allium, and Molliora cespitem. latum efformantia which includes D. ahietinum on Pinus and Abies (Picea excelsior) and D. virescens on decaying twigs. Hoffman, in Deutschl. Flora, PI. 13, 1795, used the same genus name without reference to Persoon for D. verticillatum.^ and D. antennaeforme, the former belonging in the genus Arthrinium, the latter transferred later by Persoon to Toriila and since treated there. In his Dispositio Meth. Fung, in 1797 Persoon, in supplementary material on p. ITi, gives a more complete description of D. aurewm with two varieties and adds D. Rippocaslani and D. herbarmn. It would seem best to consider D. articulatum the type of the genus since it is the only one figured, although the figure is not very helpful. Since it is reported on Allium, it is quite likely that Xenodochus Allii Harz (Torula Allii Sacc.) should be referred here. It is also likely that D. aiietinum on Picea excelsa {Pinus Abies L.) is the same as Tonola gnunulosa Lindau. In this case perhaps Groups 2 and 4 of Lindau 's Toi-ula belong in Dematium. In Persoon 's Syn. Meth. Fung. p. 694, 1801, Dematium is characterized as " Byssus forma indeterminata, erecta OAit depressa, subfasciculata OMt effusa. Fila laevia nee con- texta." Again lie divides into two groups "Rigida simplex aiU fasciculata" with D. articulatum, D. verticillatum Hoffm., D. ciliare (Hypoxylon ciliare Bull.), D. epiphyllum, D. strigosum (Byssus fulva Humb.), D. stuposum, D. bombycinu/m' (Byssus Bombycinus Roth); and "Cespitosa subintertexta nee panmmi s. pellem referentia" which included D. petraeum (Byssus petraeus Dillw.), D. olio/re, D. violaceum, D. einnabarinv/m, D. impressum, D. vvres- cens Pers., D. Hippocasfani., D. herbarum, D. brnssicae, D. fungorum, D. abietinum and D. salicinum,. It is obvious by this time that the idea of a dark color had dis- appeared, since D. abietinum is reported yellow cinnamon, D. impressum white, D. cinna- barinum, red, D. petraeum golden, D. bombycinum white, D. stuposum tawny ferruginous, D. strigosum tawny gray, leaving the rest varying shades of olivaceous to black. In his Mycologia Eur. 1822, Persoon again changes his concept very much. Several of his species of 1801 are grouped as Dematium vvlgare which also includes Cladosporium herbarum, etc. D. atrum and D. abietinum are recognized ; D. asserculorum and D. griseum are transferred here from Chloridiur,i ; D. grumosnvi, D. epiphyllum, and D. grnmirmmi are new; D. articulatum. and D. verticillatum are removed to the doubtful species. Link, in his revision of the fungi in Willdenow 's edition of Linne 's Species Plantarwm 6 : 131, 1824, uses Dematium in place of Racodiu/m of Persoon. Several of Persoon 's species of Dematium are included in Cladosporium, D. articulatum is referred to Coelosporium fruticu- losv/m, D. verticillatum to Spondylocladiii/m fumosum., D. abietinum and D. strigosu/m are said to be algae, and D. virescens is referred to Sporotrichum. Bonorden refers Dematium to Acladium in his Handbuch dcr Mylcologie, 1851. Saccardo, in his Sylloge Fung. 4: 308, 1886, uses Dem,atium in the sense of Sporodum Corda and does not include any of the species of older authors, which are referred to Cladosporium herbarum for most part. Lindau follows Saccardo. TORULA Torula Pers., Obs. Myc. 1: 24, 1796. Persoon first characterized the genus as follows: "Filis simplicibus articulatis inde- terminate efusis, mucidis." He included two species Torula monilis, from the rotting stem of an umbelliferous plant, characterized "late incrustans, atra, filorum. articulis globosis sub- contigwis," and T. fructigena., which is the common brown rot of plums, etc. He goes on to state that in this genus the spores are never in heads as in his Monilia nor branched in digiti- TORULEAE 671 form columns as in his Aspergillus (not that of Micheli) but atticulatc; the articles are deciduous, smooth, and simple in contrast to Dtnnatium. In the Syn. Meth. Fung. p. 693, 1801, lie reduces his genus to a subgenus of Monilia, changes tiie specific epithet monilis to herbaru/in to avoid reduplication, and transfers Dcmatimn onttnnaeforme Iloffm., DeiUschl. fl. cryptog. Fl. 13, Fig. 4, 1795, to this section. In his last treatment in the Myc. Eur. 1: 20-22, 1822, he again recognizes tlie genus and removes his T. fructigcna, adding T. tencra Link, Mag. Ges. Naturf. Freunde Berlin 3: 40, 3815 (Nees Syst. 2: 20, 1817); T. fuliginosa a pinophila (Antennaria pinophila Nees, Syst. 2: 72, 1817) and )3 ericophila (Antennariu ericophila, Link, Neues Jour. f. Bot. 3: 17, 1809). He also adds Hormi^cium expnnsam Kunze, Myc. Hefte 13, 1818, and H. alta Ehrenberg, keeping them distinct as a separate subgenus. He accepts the principle of Link and Nees' separation of Monilia and Tonila, although not the names for his other two subgenera, placing attenata in the former and the other species in the latter. Hence, we may conclude that Torula vionilis (T. herhar^im) is the type of Persoon's genus, since the other species first described along with it was later excluded, and that this species was retained in his central, largest section of the genus. Link, in his Ohservationes Mycologicae, Mag. Ges. Naturf. Freunde 3: 21, 1809, ac- cepts Persoon's 1796 treatment, but considers T. antennata Pcrsoon as type of his Monilia. In his revision of the fungi for Willdenow's edition of Linne's Species Plantariim 6: 128, 129, 1824, he limits it to black species with arthrospores, retaining only T. monilis (as T. herbarum) and T. tenera. He treats Hormiscium Kunze as Monilia, thereby using Momlia in a different sense from that of most other authors before or since. All the early authors used Torida for a species with dark colored moniliform hyphae creeping over bark and dead wood or plant stems, easily breaking up into arthrospores. In 1838, Turpin introduced the first serious complication by applying the name to an organism found in beer. Pasteur picked up the name for yeastlike organisms of beer which do not cause the usual fermentation of sugars. Hansen extencU-d this concept to include all asporogenous yeasts without regard to color or fermentation. Guilliermond continues the tradition of Hansen and, while admitting the possibility of black yeasts belonging here, gives the impression that these forms would probably be found to belong in Dematium when they are better known. CLADOSPORIUM Cladosporium Link, Mag. Ges. Naturf, Freunde Berlin 7: 37, 1816. Link originally characterized the genus: Thallus e floccis caespitosis erectis simplicibus aut subramosis, apicibus in sporldia secedentibus. A Sporothrico et Oidio differt Hoccis non intricatis, ab Acladio sporidiis apici primum innatis, dein delabentibus. He described four species: C. herbarum {Dematium herbarum Pers.), C. abietinum (Dematium abietinum. Pers.), C. aureum, and C. atrum as new. In Link's revision of the fungi, in Willdenow's edition of Liime's Species Plantarum 6: 39-42, 1824, he treats most of Persoon's species of Dematium, including here C. atrum. but not C. mireum. Therefore, we may eliminate C. avreum from consideration as the type of the genus. The choice thus narrows to the possibility of C. herbarum and G. atruim. Logically one would choose C. atrum, but apparently C. herbarum, has been most in the minds of later workers. C. atrum has unicellular spores while in C. herbarum these are 2-celled. Saccardo, in Michelia 2: 21, 1880, and Syll. Fung. 4: 235, 1882, kept the Persoon — ^Link tradition and has been followed by mycologists since. Lindau, in Eabenhorst, Kryptog. Fl. De^itschl. ed. 2, 8: 567, 1906, characterizes the genus as follows: Sterile hyphae absent or only a few, branched, septate, hyaline or dark colored. Conidio- phores either wholly lacking or only short side branches. Conidia either developing by the complete breaking up of the whole filament or in long chains on the tips of the short side branches which break up into single cells, which are black, brown, olive green, spherical or ellipsoid, or even fusiforni. 672 MEDICAL MYCOLOGY He divides the genus thus defined into four sections: (1) the whole mycelium breaking up into arthrospores ; (2) new spores being cut off continuously from the tips of hyphae eventually giving a basipetal chain; (3) the mycelium growing mostly by sprouting; (4) the mycelium comparatively light colored, with chains of spores developed on partially differenti- ated, short lateral conidiophores. Of the above sections, Torula should be retained for the first, 2 and 4 should be trans- ferred to Dermatium, and 3 to Pullularia or perhaps to Eormiscium. HORMISCIUM This genus was characterized by Hoffman, Myk. Hefte 1: 12, 1817, as follows: Fibrae aggregatae vel solitariae, simplices, stnctae, rigidiusculae, suppellucidae, articulatae ; articulis globosis continms. His Fig. 7 on PL 1 shows single chains of spores rising from the sub- stratum, with no traces of a conidiophore. He recognized only one species, K. expansum. Ehrenberg, Sylvae Myc. p. 22, 1818, added H. alta, from bark of Alnus glutinosus. Both these species were treated as a subgenus of Torula by Persoon, Myc. Eur. p. 22, 1822. Link, 1 '^ z Fig. 106. — 1, Horniiscium stilbosporum Corda ; 2, Hormiscium pinophilum Nees. (After Corda.) in his revision of the fungi for WUldenow's edition of Ldnnfi's Species Plantarum, places them in his Monilia which is apparently strictly a synonym of Rormiscium. He adds Dematium antennaeforme Hoffman, under name Monilia (Torula) antennata Pers., M. sparsa which may be the same as Dematviom articulatum Pers., and M. (Torula) Havimonis Ehrenberg in litt. Saccardo, Syll. Fung., and Lindau, in Rabenhorst, Kryptog. Ft. Deutschl. I, 9: 596- 604, recognize the characters as expressed by Link for his Monilia (Fig. 106). Lindau recog- nized the need for a revision of the species referred to Hormiscium and Torula but did not carry it out. Key to Genera Mycelium early evanescent, leaving single black spores varying from spherical to ellipsoid or lenticular, never fusiform. Coniosporium. Mycelium persistent, frequently dark colored (white in Indiella). Whole mycelium breaking up into arthrospores. Arthrospores short cylindric, chains not readily breaking up. Hormiscium. Arthrospores ellipsoid, chains readily breaking up. Torula. Arthrospores spherical to short ellipsoid, sprouting. Pullularia. TORULEAE 673 Whole mycelium not breaking up into arthrospores ; chains of arthrcspores borne on short lateral branches. Arthrospores ellipsoid to spherical. Spores rough. Jremispora (see p. 182). Spores smooth. D'lmatium. Arthrospores 2-celled. Cladofporium. Arthrospores not produced, mycelium tending to form sclerotia, chlamydospores abundant, strictly pathogenic, producing mycetomata. Mycelium dark gray to black. Madurella. Mycelium remaining white. Indiella. CONIOSPORIUM Coniosporium Link, Mag. Ges. Natiirf. Freunde Berlin 3: 8, 1809. The type species is Coniosporium olivaceum Link on Pimis maritima. Mycelium scanty, conidiophores hyaline, short, quickly evanescent ; co- nidia spherical to ovoid or lenticular, dark. Coniosporium onychophilum* Agostini, Boll. Sez. Ital. Soc. Internat. Mi- crobiol. 3: 214, 215, 1931. Isolated from lesions of the nails, Italy (case of Tarantelli). Not patho- genic for white mice. Mycelium dimorphic, some hyphae 2-2.5/x with refractive granules hyaline, little branched and irregularly septate, other hyphae 6-7.5/^ with oil droplets, a yellow brown pigment and regularly septate. Arthrospores lenticular, face view circular, 8-12fx in diameter; girdle view fusiform, 8-12 x 4-5/x; occasionally somewhat ellipsoid, 10-12 x 5-7/*. Epispore olivaceous or brown-olivaceous, contents granular, oily, hyaline. On treatment with acid, the wall splits into two halves. On Pollacci agar, carrot, and potato, colonies floccose, lanuginose, at first white then rose color at the surface and hazel below in the substrate. On Sabouraud agar, colonies less floccose, substrate browning. No growth on blood agar. On Raulin's solution, colonies floating, slightly floccose, rose at first, then hazel, and finally dark brown. In glucose, maltose, and galactose, colonies mucilaginous, becoming cartilaginous and dark brown. On leather, colonies small, floccose, yellow. On feathers, colonies small and whitish. No growth on hair. Ferments glucose, maltose and galactose within 6 days. Optimum temperature between 15° and 25° C; no germination at 35° C. PULLULARIA Pullularia Berkhout, De Schimmelgeschlachten Monilia, Oidium, Oospora en Tonila 55, 1923. Hormonema Lagerberg, Lundberg & Melin, Svenska Skogvardsforen. Tidskr. 1927: 219-233, Figs. 36-42, 1927. ♦Spelled onicophylum in original publication, spelling correctT^d in accordance with per- mission in International Rules of Nomenclature, Art. 70. In her more complete description, Agostini (1932) corrected the spelling. 674 MEDICAL MYCOLOGY The type species is PuUularia puUulans (Bary in Low) Berkhout. Hyphae dark colored, blastospores ovoid and lighter colored, occasionally absent. Hyphae composed of chains of dark, thick-walled large cells. Cul- tures at first yeastlike, then velvety, dark with lighter margin, sometimes remaining light colored for a long time. Sugars not fermented. It is extremely doubtful whether this genus is pathogenic, although P. puUulans is sometimes so reported. Pullularia puUulans (Bary in Low) Berkhout, de Schimmelgeschlachten Monilia, Oidium, Oospora en Torula 54, 55, 1923 ; Ashford, Porto Eico Jour. Publ. Health & Trop. Med. 5: 188-195, 1 pi., 1929. Deniatium puUulans Baiy in Low, Jahrb. Wiss. Bot. 6: 467-475, Pis. 29, 30, 1868. Isolated from extensive and deep-seated lesions, on the palms of the hands and the soles of the feet as well as on chest, arms, and legs. Lesions darkly pigmented and heavily crusted with deep infiltration of the skin. Eruption began as small vesicles on the toes, which broke and were covered later by crusts. Obstinate, finally cured by intravenous sodium iodide. No proof of pathogenicity. According to the authors, probably a saprophyte, but detailed to warn other investigators of this ubiquitous organism which vnsiy occur in cultures from various sources. In hanging drop at first sprouting from any part of parent cell. The daughter cell may separate in a few hours or it may grow to normal size while still attached, forming short, sometimes branched chains. Then almost simul- taneously all cells in the chain put forth sprouts laterally. These lateral cells usually remain smaller and are sometimes referred to as conidia. They may separate and either continue sprouting or develop true hyaline hyphae, espe- cially on solid media where the central axis of the colony is formed by the chain and its lateral sprouts, intersected by the netAvork of hyphae which the latter produce. These become so intricate that it is no longer easy to make out the structure of the colony. In liquid media, the sprouting cells often become uniseptate (the Cladosporium stage). Finally, some of the cells greatly thicken their walls and produce a black pigment. This process continues until the surface of the colony becomes a layer of these hypnospores. If con- ditions favorable for renewed growth are present, they germinate by fine filamentous hyphae. If conditions continue unfavorable, the walls of the hypnospores gelify and fuse into a blackish or dark yellow crust. They ordi- narily do not subsequently germinate but rather serve to protect the sprout cells beneath. Ciferri & Ashford report that the terminal cells of the hyphae are clavate to subspheric and sometimes sprout chains of cells, superficially suggesting Hormodendron. Young colonies white and similar to those of yeasts, older colonies with yellowish tan to brown spots, then black and shining, finally black, opaque (fumagoid type). In liquid cultures, colonies in tufts growing downward from surface, not forming sediment, white becoming grayish and greenish with a maroon surface. No fermentation has been noted in any sugars, al- TORULEAE 675 though growth is good in media containing glucose, fructose, maltose, dextrin, lactose, 2% methyl or ethyl alcohol, citric, malic, tartaric or acetic acid. Peptone, asparagin, glycocoll, ammonium sulphate or potassium nitrite or nitrate may be utilized as sources of nitrogen. Cryptococcus metaniger Castellani, Arch. Derm. Syphilol. 16: 402, 403, 1927; Amer. Jour. Trop. Med. 8: 393, 1928, probably also belongs here. It was isolated from a case of trichomycosis nigra and described as cells elongate, black, colonies black on all laboratory media. Too vaguely described for definite placing by any one at all familiar with this large group. Ciferri, Arch. Protistenk. 71: 445, 446, 1930, has also placed it here, while Ferrari (1932) places it in Cladosporium. Pullularia Jeanselmei (Langeron) Dodge, n. comb. Torula Jeanselmei Langeron, Ann. Parasitol. Hum. Comp. 6: 385-403, 12 figs., 1928. Fig. 107. — Pullularia Jeanselmei. (After Langeron 1928.) Isolated from black grain mycetoma in Paris in a mulatto woman from Martinique. Case history by Jeanselm, Huet & Lotte, 1928. Mycelium in lesions forms grains 500-1,000/x in diameter, soft, irregular, and appearing like hollow cylinders. Hyphae coiled, interstitial gel not well developed. Hyphae, 2.5-3/* in diameter, forming chains of arthrospores, 4.5-5/i, which soon disintegrate (Fig. 107). In cultures, the arthrospores form chains of spherical cells which laterally produce ovoid blastospores in the vicinity of the septa. Toward edges of the colony, fewer lateral chlamydospores are produced and terminal whorls like conidia are produced on short lateral branches. These chlamydospores are 2.5 X 5yu,, often with 3-8 per group. On sugar media contours are rounded and submoniliform. Cultures grow well at 22°-25° C. on the usual media (Sabour- aud glucose and malt agars, potato-glycerol, carrot and potato decoctions). 676 MEDICAL MYCOLOGY I have hesitated to place this species in PulluJaria but it apparently has blastospores as well as chlamydospores. From its soni'ce, one would expect it to be related rather with MadnreUa. DEMATIUM Dematium Persoon, Neues Mag. Bot. 1 : 121, 1794. Montoyella Castellani & Chalmers, Man. Trop. Med. ed. 2, 799, 1913. The type species is Dematium articulatum Persoon, isolated from Allium (Fig. 108). Hyphae hyaline, septate, branched, bearing chains of smooth, black arthro- spores on lateral branches (Fig. 109). The species are predominantly parasitic or saprophytic on plants. The pathogenic species referred here have not been very carefully described, and their reference here is somewhat doubtful. On the other hand, there is nothing in their descriptions to warrant placing them in Cladosporium which has two- Fig-. 108. — Demntiuni articulatum Pers. (Torula Allii Sacc.) type of the ffenus Dematium. (After Lindau 1906.) celled arthrospores. It is to be hoped that some one will undertake a careful study of the morphology of the whole group with black arthrospores. These species seem to produce relatively superficial lesions on the skin. Dematium Wernecki (Parreiras Horta) Dodge, n. comb. Carats noir Montoya y Florez, Recherches sur les carates de Colombie, These Med. Paris 25: 48, 49, 1898. Montoyella nigra Castellani & Chalmers, Man. Trop. Med. ed. 2, 799, 1913, non Dematium nigrum Link 1809. Cladosporium Wernecki Parreiras Horta, Rev. Med. Cirurg. do Brasil 29: 269-274, 1921. Cladosporium sp. Rietmann, Bull. Soc. Derm. Syphiligr. 37: 202-207, 1930; Sartory, Sartory, Rietmann & Meyer, C. R. Soc. Biol. 104: 878-881, 1930. The organism usually isolated from black Carate, pinta, or Kcara in South America. Pena Chavarria & Shipley (1925) report a species of Alternaria from such lesions [probably a contamination]. TORULEAE 677 We owe our first extensive study of this disease to Montoya y Florez who described it in Colombia as apparentlj^ confined to full-blooded negroes. Pro- ducing spots from 0.5-2 mm. in diameter on the forehead, chin, forearms, and legs; rarely generalized, smooth, not pruriginous or scaly. The skin is much vascularized and the epidermis very adherent. Sometimes it remains in this stage, at other times it results in achromia which may cover the whole body except the face, backs of the hands, and feet. The plantar surface is usually hyperkeratinized. The disease in Brazil was first described by Parreiras Horta from the case of Joao Ramos e Silva and Jose Torres in Werneck Machado's service. The lesion was a black spot on the left hypothenar region, slightly elevated with slight desquamation. Silva (1929) reports a similar lesion on the side of the middle finger. Fonseca & Ferreira da Rosa (1930) report it from various sites on the hand, frequently causing hyperkeratosis of the Pig. 109. — Dematium hispidulum. (After Saccardo. ) palmar surface. The white race seems to be affected as well as the negro in Brazil. Cerqueira studied the condition as early as 1891 but did not publish his results. A. Sartory, R. Sartory, Rietmann & J. Meyer found subcutaneous and intraperitoneal injections into guinea pigs negative. Superficial inocula- tion in the inguinal region produced black plaques developing in 5-7 weeks, forming scaly crusts before spontaneous healing. The liquid from culture media produced death by anaphylactic shock. Lesions were easily reproduced on inoculation into human subjects. Mycelium dimorphic, some hyphae slender and branched, septate, other hyphae larger, varicose, thick-walled with many chlamydospores ; hyphae aris- ing from the massive mycelium bearing terminal pyriform conidia. In scales, hyphae long, septate, terminated by arthrospores ; others slender, solitary, rounded or narrowed at the tips, often flexuous, whole mycelium dark green- 678 MEDICAL MYCOLOGY ish, rarely branched ; spores variable in size, spherical or ovoid, not in chains. Fonseca & Ferreira Rosa (1930) report hyphae branched in scales. On glucose agar, the Fumago form with rounded cells, thick-walled, some- times deeply colored. On carrot, the Dematium form with mycelium of short cells, with simple blastospores ; yeast forms abundant, giving cultures humid or varnished aspect. In carrot decoction, delicate mycelium with abundant spores furnished with thickenings which serve as disjunctors. At first colony easily separable from medium, giving the appearance of a black yeast; later it becomes very hard with elevations and depressions similar to those of Trichophyton acuminatum. Colony covered with a greenish bloom which leaves a very black core. A. Sartory, R. Sartory, Rietmann & J. Meyer (1930) report that it grows best on lipoid-containing media. It utilized glycine, hip- puric acid, creatine, and creatinine freely, purines slowly and secondarily, urea and guanidine not at all. Dematimn Mansoni (Castellani) Dodge, n. comb. Microsporum Mansoni Castellani, Brit. Med. Jour. 2: 1271, 1905. Foxia Mansoni Castellani, Jour. Trop. Med. Hyg. 11: 260, 1908. Cladosporium Mansoni Pinoy, Ann. Derm. Syphiligr. V, 3: 341-343, 1912. Torula Mansoni Vuillemin, C. R. Acad. Sci. 189: 406, 1929. Isolated from cases of pityriasis nigra in India. Hyphae short, 18-20 x 2.5yu, in diameter, arthrospores spherical, 5-7.5/i,, frequently in clusters, uniguttulate. Pollacci & Nannizzi report that olivaceous hyphae, little branched, 2-2.5/1, are terminated by ovoid or ellipsoid arthrospores, 7-8.5 x 2-2. 5;*, one-celled or occasionally two-celled. On maltose agar, colonies hemispheric, dark greenish to black, coalescing, forming a jet black, rugose mass. On broth and peptone solution, sediment greenish black, no action and little growth on milk. Growth slow on gelatin with little or no liquefaction. Optimum temperature 30°-32° C. ; growth slow over 35° or under 25° C. Dem.atium Gougeroti (Matruchot) Grigorakis, Bull. Soc. Myc. France 40: 272-276, 4 figs., 1924. Sporotrichum Gougeroti Matruchot, C. R. Acad. Sci. 150: 543-545, 1910. Rhinocladium Gougeroti Verdun, Precis Parasitol. 1913. Early isolated from a gumma deep in a human leg muscle. Later found in an ulcer on the dorsum of the penis with inguinal adenitis in a young man in Madagascar (Fontoynont & Boucher, 1923) and a very similar case in Porto Rico (Kesten et al., 1932). Griitz (1925) reports it from more super- ficial lesions. Pathogenic to rats and monkeys (Kesten). Mycelium of moniliform filaments or isolated sprout cells. These give rise to slender hyphae, which collect and produce black coremia. Spores often borne in lateral tufts. Kesten et al. report spores 3.5-5.5 x 1.5-2.5/1. Colonies black from the first, elevated, with a velvety surface. TORULEAE 671) Several have referred Sporotriclium Lecante Beiinnaiiii & Gougerot to synonymy with this species, but I have been unable to locate the original description of Benrmann & Gougerot. Dematium Sakuranei (Jannin) Dodge, n. comb. Oidium sp. Sakurane, Arcli. Derm. Syphilis 78: 211-221, 1906. Mycoderma Sakuranei Jannin, Les Mycodemia . . . 247, 248, 1913. Cryptococcus Sal-uranei Froilano de Mello & Gonzaga Fernandes, Arq. Hig. Pat. Exot. 6: 293, 297, 1918. Monilia Sakuranei Vuillemin, Champ. Parasit. Homme Anim. 86, 1931. Geotrichum Sakuranei Almeida, Annaes Fac. Med. Sao Paulo 9: 12, 1933. Isolated from cold subcutaneous abscess in a nine-year-old girl in Japan. Neoformation of cell tissue, ulceration of dermis, and painless tumefaction of lymphatic ganglia. Mycelium apparent on direct examination. Pathogenicity not clearly established. Hypliae and ovoid to spherical cells. True arthrospores seen but no true conidia. Hyphae branched, septate on potato and sugar media. On ordinary agar, colonies brownish black, circular, more or less irregu- lar, becoming hemispheric, hard, surface spongy, Avithout special elevations, becoming moister in age, easily separable, the colony then consisting prin- cipally of arthrospores. On glucose agar, brownish black or dark green, hard, adherent, covered with a thin, gray mycelium which finally becomes brownish black. On potato, colony gray green with gray aerial mycelium at the per- iphery, growing deep into the medium. On gelatin, colony greenish brown, granular, growth slow, no aerial mycelium, no liquefaction. On glucose broth and milk, spongy flocci below, dark brownish black, hard pellicle on surface. Milk not coagulated. Doubtful Position The following species may belong in Dematium or in Torula, but it is too poorly described for certain determination. Aspergillus caxbonarius seu ater Meis & Parascandalo, Gaz. Ospedali 16: 769-772, 1895. Isolated from a case of diphtheria in a dog. Appears to be a contaminant and not pathogenic. Mycelium not septate, branched. Fertile hypha swollen at apex; spores in chains, "surrounded by a membrane," 5-7/a in diameter, black. Hyphae and mycelium also black. Optimum temperature for growth 30°-37° C, growth ceasing at 60° C. Growth good on agar, poor on potato, none on egg albumen. On gelatin, small colonies resembling coffee beans. Growth rapid on prune decoction and bread, also on Naegeli's solution. Glucose and sucrose fermented. Gas evolved with gelatin. This is obviously not a species of Aspergillus since it has black mycelium. It is too pooi-ly described to identify, but probably is a saprophytic Dematium. 680 MEDICAL MYCOLOGY MADURELLA Madurella Brumpt, C. R. Soc. Biol. 57: 999, 1905. Streptothrix mycetonii Laveraii is the type species. Mycelium white at first, parasitic on a variety of animal tissues; bones, muscles, connective tissue. The vegetative hyphae have a diameter always larger than 1/x, sometimes up to 8-10j«. These hyphae are septate, branch from time to time and secrete a brown pigment (Fig. 110). In age they agglomerate into sclerotia where one finds a variable quantity of spherical bodies, 8-30/x in diameter (chlamydospores). Fig. 110. — Madurella mycetonii. Mycelium and chlamydospore.s. (After Brumpt 1906.) Members of this genus have been found almost exclusively in mycetoma with black grains (maduromycosis, Madura foot, etc.). This disease has been known clinically for several centuries in India and has been repeatedly de- scribed (cf. Carter 1874, Brumpt 1906, Musgrave & Clegg 1907, Gammel 1926, Heitor Froes 1930 for excellent summaries of the literature). It seems to be more common in regions with hot dry seasons where the vegetation abounds in thorns and the natives go barefoot, a large proportion of the patients giving the history of a thorn in the foot some time before the lesion became serious. However, the cases are not confined to such regions. The lesions are usually restricted to the feet, but are occasionally found in other situations, such as TORULEAE 681 in the groin and in one patient behind the ear. The lesion begins by swelling of the affected member, nsnally accompanied by one or more nodules, pustules, or abscesses which, upon rupture, become fistulas. New lesions make their appearance near the first until a considerable portion of the member may be involved. The tumor mass begins to soften, and the lesions burst and dis- charge an oily, viscid, seropurulent fluid with small black grains (sclerotia). The overlying skin becomes discolored, usually darker, mottled and scarred, surmounted by discharging sinuses. Pain is usually mild or even lacking, but the resulting deformity often interferes with locomotion. In general, the bones are not affected by Madurella, although they frequently are when Actinomyces is the etiologic agent. Surgical removal of the tumorous mass or amputation is usually the only satisfactory method of treatment. In the bibliography at the end of this chapter, I have collected all the references to this disease where the organism was not presumably a species of Actinomyces, Monosporium, or Aspergillus. Key to Species Colony white. M. Toseuri. Colony grayish ashy on carrot, oehraceous on potato, and brown on Sabouraud agar. M. Ea/miroi. Colony greenish gray to purplish gray, becoming purplish when old; on Sabouraud agar yellowish, then purplish. M. Oswaldoi. Colony grayish white, zonate, center brown, folded, becoming dry membrane, deep chamois or darker. M. algiris. Colony copper yellow, margin greenish yellow, finally covered with a brown pubescence on Sabouraud agar; gelatin not liquefied. M. Tabarkae. Colony grayish white, becoming liglit brown, gelatin liquefied, sclerotia abundant on Sabouraud conservation agar. Colonies accuminate on Griitz and Sabouraud agar. M. IJcedae. Colonies tomentose and cerebriform on Griitz agar, little or no growth on Sabouraud agar. M. americana. Madurella Tozeuri (Nicolle & Pinoy) Pinoy, Ann. Derm. Syphiligr. V, 3: 341-343, 1912. Oospora Tozeuri Nicolle & Pinoy, Bull. Soc. Path. Exot. 1: 95-99, 1 fig., 1908. Isolated from a growth on the top of the foot, not involving muscles, joints, or bones. Epidermis and dermis only affected. No pain. Lesions eighteen years in developing. Hyphae straight or flexuous, l-4/i, in diameter. Cells have thick walls. Chlamydospores 2-6/x in diameter. Hyphae united by a brown pigmented mass, or follow blood vessels, branching at the capillaries. Grains 50-100/a in di- ameter, with outer layer of filaments yellow brown, l-2;tt in diameter. Organism cultivated on several media. On glucose or maltose agar, in 24 hours at 37° C, white colonies develop with a blackening of the substrate. Same on potato. Media not blackened in the absence of carbohydrates. 682 MEDICAL MYCOLOGY Madurella Ramiroi Piraja da Silva, Brasil Med. 33: 81-83, 1919; Mem. Inst. Butantan 1: 187-207, Pis. 34-38, 1918-1919. Isolated from a black grain Madura foot in Brazil. The author was unable to reproduce lesions in pigeons, rats, or bats. Mycelium moniliform, cells about 2.7/a in diameter; there are slender my- celium producing chlamydospores (intercalary) and moniliform mycelium. Large spherical cells, 22/* in diameter, seen in the sclerotia are probably large chlamydospores similar to those in the grains from lesions. On Sabouraud glucose, and maltose agars, growth rather rapid, colonies deep brown, cerebriform with abundant pigment formation. Sclerotia rare. Colonies on yam dark gray in the center, with ashy margins. On sweet potato, growth better than on banana (de terra). The culture media blacken as the colonies age. On carrot, growth is grayish ashy, with irregular verrucose surface. Growth best on potato, colonies ochraceous with medium darkening. In potato decoction, tufts of mycelium with small black granules. In hay infusion, there are formed white tufts which adhere to the walls of the tubes and later there are small grains (sclerotia) similar to those from the lesions. Madurella Oswaldoi Parreiras Horta, A Pathologia Geral Nos. 1-4, 1919, [quoted by Gammel 1926, Fonseca 1930, Froes 1930]. Organism shows a luxuriant mycelium which changes its color during growth from greenish gray to purplish gray, becoming purplish when old. No sclerotia seen in cultures. Cultures show a central knob with a deep depression and a zone of deep fissures. Growth rapid on all media. On Sabouraud media, colonies are white to dirty yellow and purple ; with sugar media, either liquid or solid, darkening. Madurella algiris (Blanchard) Dodge, n. comb. Oospora Tozeuri var. Brault & Masselot, Soc. Chirurg. April, 1911 ; Arch, de Parasitol. 15: 218-225, 1912. Discomyces algiris Blanchard. Oospora algirus Sartory, Champ. Parasit. Homme Anim. 777, 778, 1923. Isolated from a tumor on the foot of a young Arab fifteen years old. Tumor just behind the space between first and second toes, size of a large nut, surface thin and ready to burst. On puncture, there was exuded a reddish viscous liquid Avith black grains varying in size from that of a pinhead to that of a grain of wheat. Cavity lined with a thick, soft, brownish tissue. KI was not tolerated. Patient treated frequently with tincture of iodine until cured. Organism not pathogenic to laboratory animals. In grains, mycelium septate with large intercalary chlamydospores 7 x 12fx in diameter. Hyphae B/x in diameter, cells 5.5-8/x long. Growth good on glucose and glucose glycerol agar. Colonies grayish white with concentric rings of growth, center browning and folding. In age it appears as a dry yellow membrane from deep chamois to tinder, with medium becoming black or caramel in color. On potato, growth is poor, colony dirty white, with medium blackening. On carrot, especially with glycerol, growth TORUIiEAE 683 is similar to that on potato but more rapid. lu hay infusion, there are cottony tufts with slight sediment and no clouding of the medium. Madurella Tabarkae Blanc & Brun, Bull. Soc. Path. Exot. 12: 741-748, 1 pi, 1919. Isolated from a black grain mycetoma which was subcutaneous without involving muscles or bones. Grains removed and wounds healed normally. The sclerotia are composed of fine filaments, 2-4/a in diameter, with chlamy- dospores either terminal or intercalary, 12/x in diameter. Hyphae slender, cells short. Optimum temperature 22° C, growth slow at 35° C. Organism grows best on Sabouraud maltose agar. On simple Sabouraud agar, it produces good colonies, with the growth finally becoming rounded and folded. Color begins at the center and spreads outward. After one month, center is copper yellow and periphery greenish yellow, with amber 3'ellow guttation in the center. Culture finally becomes covered with a brown pubescence and medium turns brown. Similar, but faster, on Sabouraud maltose agar. On ordinary agar, hay infusion, or potato agar, in 24 hours, small white colonies appear, while medium darkens progressively. Colonies are never large. Similar on lead agar with growth sparse and white. On neutral red agar, growth good with no change or browning of medium. On potato, potato glycerol, or carrot, small white colonies which become yellow and folded; potato rapidly blackens. On coagulated serum, growth as an ordinary agar except that colony is rose at first, later becoming brown. On gelatin stab, there is abundant growth on the surface, which becomes in- tensely brown with arborization and sclerotium formation below. In beef peptone broth in 24-48 hours, small globular masses are especially abundant in contact with the tube. Medium progressively browned up to the fifteenth day. In peptone broth, growth is the same. No indol formation. No growth in hay infusion. In potato infusion as in beef broth, except that the floecu- lent colonies slowly collect at the bottom of the tube. Black sclerotia appear in about 25-30 days. In peptone sugar solutions, neither acid nor gas is formed (no fermentation). Growth as in beef broth. In peptone bile, small spherical colonies appear with browning of the medium. Milk is coagulated on the eighth day with browning of the medium and abundant growth on the surface. Gelatin not liquefied. Madurella Ikedae Gammel, Arch. Derm. Syphilol. 15: 241-284, 20 figs., 1927. Isolated in Minneapolis by Ikeda from a black grain maduromycosis. Nonpathogenic to rabbit or guinea pig. Mycelium grayish white, later light brown, darkening certain sugar media. Hyphae hyaline or subhyaline, rarely granular, varying in diameter from 1.5-5/i. Chlamydospores numerous only in Sabouraud glucose broth. In all liquid media many hyphae break up into chains of citriform or spherical spores. Acuminate light brown colonies on both Sabouraud and Griitz a gars. Sclerotia numerous on surface and in depth of Sabouraud preservation me- 684 MEDICAL MYCOLOGY dium. Pigment production abundant in glucose, galactose, or maltose, moder- ate in mannite or levulose, poor in dextrin, poor or absent in inulin, sac- charose, or lactose broth. Organism ferments milk and is thermophilic. Gelatin liquefied. Madurella americana Gammel et al., Arch. Derm. Syphilol. 13: 66-77, 6 figs., 1926; Ibid. 15: 241-284, 20 figs., 1927. Acladium americanum Ota, Jap. Jour. Derm. Urol. 28: [4], 1928. Isolated from black grain mycetoma in Cleveland, Ohio. Animal inocula- tions negative. Mycelium yellowish brown and darkening certain sugar media, grayish white when old. Hyphae subhyaline or brown and granular, varying in di- ameter from 1-5. 2/x according to the type and percentage of sugar in the medium. Chlamydospores numerous, especially on Sabouraud glucose broth; arthrospores with squarely cut off ends rare. Optimal medium for growth is 1% glucose agar. Growth on Sabouraud agar none or very poor. On Griitz agar, colony tomentose or cerebriform. Sclerotia, 1 mm. in diameter, form on surface of gelatin and Sabouraud con- servation agar. Pigment formation is abundant in glucose, maltose, dextrin, and galactose ; moderate in fructose ; poor in sucrose or lactose ; absent in inulin broth. Milk is fermented, organism being thermophilic. Gelatin is liquefied. Madurella mycetomi (Laveran) Brumpt, C. R. Soc. Biol. 57: 997-999, 1905. Streptothrix mycetomi Laveran, Bull. Acad. Med. Paris III, 47: 773-776, Fig. 1, 1902. One of the commonest species producing mycetoma with black grains. Laveran does not report spore or cultural characters, although he recognized the characters which separate it from Actinomyces. Brumpt, Bouffard, and Chabaneix give interesting case histories and clinical observations but were unsuccessful in their attempts to cultivate it. Chatterjee, Centralbl. Bakt. I, 61: 358-365, Pis. 1, 2, 1911, was the first to describe cultural characters. Noc & Jouenne, Ann. Inst. Pasteur 36: 365-385, 1922, give cultural characters in detail. Puyaubert & Jolly (1918, 1920) report an interesting case in the perineum extending from the base of the scrotum to within 3 cm. of the anus, involving muscular tissues only. They cultivated the organism which was not pathogenic for guinea pig. Doubtful Position The following species belong in Madurella but are too poorly described to place more definitely. Aspergillus mycetomi Villabruzzi & Gelonesi, Annali Med. Nav. Colon. 33: 283-308, 8 figs., 1927. Isolated from black grain mycetoma in Somaliland. No attempt was made to determine its pathogenicity. TORULEAE 685 Hypliae 3-4/i, in diameter, septate, rarely branched. Spherical or ovoid sporophore swells to form vesicle and is 4-6-spored. Spores spherical, Bis- marck brown. No trace of conidia or ascocarps. Grows rapidly only on banana, very slowly on other media. Colonies small yellowish white masses the size of a lentil or smaller, becoming zonate ; yellowish, orange, blackish brown with a white margin. Madurella Bouffardi (Brumpt) Dodge, n. comb. Aspergillus Bouffardi Brumpt, Arch, de Parasitol. 10: 525-533, 1906 [Case described and organism figured by Bouffard, Ann. Hyg. Med. Colon. 8: 579- 590, i 2?/., 1905]. Isolated at Djibouti, Somaliland, from tumors in the adipose tissues of the foot which did not affect the epidermis, muscles, or bones. The anterior third of the foot was globular, but there was no suppuration. The tumors were fibrous, each containing 3-7 black grains embedded in the center, and varying in size from that of a pea to that of a hen's egg. When these were surgically removed, healing was rapid and complete. Source of infection unaccounted for. Inoculations unsuccessful in monkey, dog, cat, and gazelle. Young mycelium silver white with a brown periphery forming a cortical zone ; conidiophore erect, simple, continuous, white, 2/x, in diameter with vesicle 4.5/a broad by 6^ tall. Conidia spherical, 1.33-2;u, in diameter, smooth, white, in chains. Chlamydospores spherical, 5-10;u, broad. Intercalary chlamy- dospores colorless, terminal ones brownish. Grains are similar to sclerotia except that they are sporiferous within. Madurella Bovoi Brumpt, Precis Parasitol. ed. 1, 1910. Aspergillus fumigatus Bovo, Policlinico Sez. Chirurg. 13: 97, 119, 6 figs., 1906. Isolated from a typical Madura foot with a few granules appearing in the groin. A grain from the foot was treated with 30% KOH for 12 hours, then kept for 24 hours in distilled water, stored in 80% alcohol for a year, and then treated for 24 hours with 25% KOH. On examination it showed hyphae 4-6/i in diameter, septate. Bovo figured something which looks like a young mucor sporangium, is ovoid and stains with methylene blue. He thought this was a head of Aspergillus. This organism probably should be referred to M. mycetomi. INDIELLA Indiella Brumpt, Arch, de Parasitol. 10: 547, 1906. This genus includes IMucedineae with Avhite thallus, parasitic in a variety of animal tissues (bone, muscle, connective), possessing vegetative hyphae which vary in diameter from 1 to 5 and even 10/*. These hyphae are septate and laterally branched, never secrete a pigment, form grains like the sclerotia which characterize the different species of the genus. In these grains there are present a fairly large number of chlamydospores, usually terminal. 686 MEDICAL MYCOLOGY The type species is Indiella Mansoni Brumpt. Indiella Brtunpti Piraja da Silva, Sobre uma nova maduromycose da graos branclios . . . Thesis Bahia, 1922. Following description by Gammel, Arch. Derm. Syphilol. 15: 256, 1927. Sterile white hyphae at first slender, scarcely septate and about 2/i. in diameter, then thicker, up to 7/x in diameter, and septate. Septa variably spaced from 3-15/x apart. White grains are soft, composed of densely packed hyphae, generally spherical and 0.06-0.08 mm. in diameter. Chlamydospores terminal, often spherical. Indiella Mansoni Brumpt, Arch, de Parasitol. 10: 545-549, Fig. 9, 1906. This organism is parasitic on man in India. Mycelium white, quite slender when young, 1.5-2/a in diameter, septate with walls 15-20/1, apart. Older hyphae irregular, 3-5/* in diameter, walls 5-10/i apart. Many chlamydospores present, terminal or intercalary, 5-12/* in di- Fig. 111. — Ithdiella Mansoni. (After Brumpt 1906.) ameter, generally spherical and unicellular, rarely segmented (Fig. 111). Grains in tissues are very small, flattened, varjdng from 200-250/* in diameter, often parasitized by bacteria. Indiella Reynieri Brumpt, Arch, de Parasitol. 10: 549-555, Figs. 10, 11, 1906. Organism found parasitic on man in Paris. Thallus white. Young mycelium septate and slender, 1-1.5/x in diameter, septa 10-15/1 apart. Peripheral hyphae 4-5/i in diameter, irregular, moniliform, walls closer than in central hyphae, each terminated by a 2-3-celled chlamydo- spore, from 5-20/a in diameter. Intercalary chlamydospores rare, grain or sclerotium rounded when young, later becoming like a twisted cord or the excrement of the earthworm and about 1 mm. in diameter. Doubtful Position It seems likely that the organism described as Halohyssus moniliformis var. parasiticus Maffei, Atti 1st. Bot. R. Univ. Pavia IV, 3: 19-28, 10 figs., 1932, belongs in Indiella, or if it proves to belong in Halohyssus, perhaps that genus TORULEAE 687 should be amended to include Indiella. This reference was received too late for critical study of the case history, which is recorded by Barco, P., Granu- loma del polso a sede sottocutanea da micete del genere Halobyssus Zukal, Arch. Ital. Chirurg. 29: 1931, but MafPei summarizes the case. Isolated from mycetoma of the wrist, Italy. Hyphae 1-5/* in diameter, hyaline, septate, branched finely granular, form- ing terminal chains of chlamydospores with a tendency for the terminal spore to be the largest and each successive one smaller until the size of the hyphae is reached. Chlamydospores with thick verrncose walls, 4-14/x, more commonly 10-12/i, in diameter. Growth good on Pollacci agar, a coarsely granular mass slightly floccose, Avhitish. Growth good in eitticr liquid or solid media. BIBLIOGRAPHY Adami, J. George &i B. A. Kirkpatrick. 1895. A case of Madura foot disease (mycetoma pedis, ochroid variety), Trans. Ass. Amer. Physicians 10: 92-100, 3 figs. Agostini, Angela. 1931. Coniosporium isole d'un cas d 'onyehomycose, Boll. Ses. Ital. Soc. Iniernaz. Microbiol. 3: 214, 215. — . 1931. Coniosporium onychophilum n. sp. causa di onicomicosi, Atti 1st. Bot. B. Univ. Favia IV, 3: 29-36. Aitchison, J. E. Tierney. 1861. Godfrey and Eyre's tubercular disease, Indian Ann. Med. Sci. 7: 517-520. Almeida, Floriano Paulo de. 1932. ConsideraQoes sobre o diagnostico histopathologico dos mycetomas, Eevista Biol. Syg. 3: 93-95, Pis. 3-4. Anderson, Charles. 1924. Sur un douzieme cas de pied de Madura observe en Tunisie, Arch. Inst. Paste^ir Tunis 13: 68-70. Anderson, Ch., E. Broc & Cassar. 1926. Deux nouvelles observations tunisiennes de mycetome. Arch. Inst. Pasteu/r Tunis 15: 332-335. Armand, Delille, P. Duhamel k Marty. 1923. Gommcs cutanees et manifestations pul- monaires dues a une mycose d'espece encore indeterminee. Bull. Soc. Pediatr. Paris 21: 190-193. Arwine, J. T. & D. S. Lamb. 1899. A fifth case of fungus foot in America, Am. Jour. Med. Sci. 118: 393-399, 5 figs. Audrain, L. C. 1924. A case of mycetoma, Jowr. Am. Med. Ass. 83: 1165. Barreto, Antonio Luiz B. & Carlos Burle Figueiredo. 1919. Da reacgao de fixagao de comple- mento no mycetoma podal ou pe de Madura, Brasil Med. 33: 403. Bassini, E. 1888. Un caso di micetoma al piede o Piede di Madura, Arch. Sci. Med. 12: 309-318, Pis. 10-12. Bays, James T. 1905. Madura foot in Cape Colony, S. African Med. Bee. 3: 75. Beldk, Alexander. 1919. Studien an zwei von v. Verebely aus Madurafiissen geziichteten Pilzstammen, Centralbl. BaTct. I, 83: 528-530. Berkeley, Miles Joseph. 1864. Observations on a peculiar mode of fructifications in Chionyphe Carteri Berk., Jour. Linn. Soc. London Bot. 8: 139-142, PI. 10. Beron, B. 1931. Zwei autochtone Falle von Mycetoma mit schwarzen Korner, Derm. Woch. 92: 265-272, 16 figs. Blanc, Georges & Gabriel Brun. 1919. Nouveau cas de mycetome a grains noirs observes en Tunisie, Bull. Soc. Path. Fxot. 12: 741-748, 1 pL Bloch, Bruno. 1916. iJber einen Fall von Mycetoma pedis, verursacht durch eine bis jetzt noch nicht beschriebene Streptothrix Art, Cor. Bl. Schweis Aerste 46: 270, 271. Bocarro, J. E. 1893. An analysis of one hundred cases of mycetoma, Lan<;et 2: 797, 798. — . 1895. Mycetoma, Lancet 1: 70, 71. MEDICAL MYCOLOGY Bockenheimer, Ph. 1909. Einiges iiber die Behandlung chirurgischer Krankheiten in Asien, Arch. Klin. Chirurg. 90: 174-190, S pis. Borja, Antonio. 1918. Sobre un caso dc mycetoma de graos negros, Brasil Med. 32: 221. Bouffard. 1902. Pieds de Madura observes a Djibouti, Ann. Eyg. Med. Colon. 5: 636-652. — . 1905. Mycetoma a grains noirs en Afrique, Ann. Hyg. Med. Colon. 8: 579-590. Boyd, Mark F. & Earl D. Crutehfield. 1921. A contribution to the study of mycetoma in North America, Amer. Jour. Trap. Med. 1: 215-289. Brault, J. 1912. Mycetome a grains noirs observe en Algerie, isolement du Madurella mycetomi, Ann. Derm. Syphiligr. V, 3: 333-343. — . 1913. Note sur les cultures de Madurella mycetomi. Bull. Soc. Path. Exot. 6: 407-409. Brault, J. & Lucien Masselot. 1912. Mycose du pied a grains noirs chez un jeune indigene algerien, Arch, de Parasitol. 15: 218-225, 4 figs. Brown, Grafton Tyler. 1932. Sensitization to fungi, Ann. Intern. Med. 6: 655-671. Bruas. 1903. Pied de Madura a grains noirs observe k Madagascar, Ann. Hyg. Med. Colon. 6: 602-605. Brumpt, Emile. 1902. Notes et observations sur les maladies parasites. 10. Mycetome a grains noirs, Arch, de Parasitol. 5: 151-156, 460. 11, Mycetome a grains blancs, Ibid. 5: 156-158. — . 1905. Sur le mycetome a grains noirs, maladie produite par une mucedinee du genre Madurella n.g., C. R. Soc. Biol. 57: 997-999, — . 1906. Les mycetomes. Arch, de Parasitol. 10: 489-572, 10 pis. [These Fac. Med. Paris, 94 pp., 10 pis.] Brumpt, Emile, Bouffard & J. A. Chabaneix. 1901. Notes sur quelques cas de paludisme et sur un cas de mycetome observes h Djibouti, Arch, de Parasitol. 4: 564-567. Burnet, Etienne & Gabriel Brun. 1926. XVIIe observation tunisienne de myceetome, Arch. Inst. Pasteur Ttmis. 15: 151-153. Canal Feijoo, E. J. «fc R. Barrionuevo. 1932. Un nuevo caso de micetoma con granos negros en Santiago del Estero, Eeunlon Soc. Argentina Patol. Reg. del Norte en Tucv/mdn 7: 496-500, 6 figs. Carter, Henry Vandyke. 1863. On mycetoma, Brit. Foreign Med. Chir. Rev. 32: 198-203. — . 1873. The parasitic fungus of mycetoma, or the fungus of India, Trans. Path. Soc. London 24: 260-263. — . 1874. On mycetoma or the fungus disease of India, London xi, 118 pp. 11 pis. — . 1874. On the nature of mycetoma, or the fungus disease of India, Lancet (London) 2: 44-46; 113-115. Catanei, A., Grosdemange & Cli. Legroux. 1927. Sur un cas de mycetome du pied observe en Algerie, Bull. Soc. Path. Exot. 20: 11-13. Castellani, Aldo. 1925. Notes on non-gonorrhoeal urethrites. Jour. Trop. Med. Syg. 28: 250-252. Catsaras, Johannes. 1912. Zwei Falle von Madurafuss (Mycetoma pedis) in Griechenland, Arch. Schifs-Tropenhyg. 16: 461-474, PI. 2. — . 1915. Bemerkungen iiber neue Falle von griechischen Myzetom, Arch. Schiffs-Tropenhyg. 19: 617-625. Chabaneix, J. A. & Bouffard. 1901. Pieds de Madura observes a Djibouti, Ann. Hyg. Med. Colon. 4: 452-456. Chalmers, Albert J. & E. G. Archibald. 1918. The classification of the mycetomas, Jour. Trop. Med. Hyg. 21: 121-123. — . 1916. A Sudanese maduromycosis, Ann. Trop. Med. Parasitol. 10: 169-222, PI. 4-7. — . 1917. Mycetoma and pseudomycetomatous formations, New Orleans Med. Surg. Jour. 70: 455-475. Chatter jee, G. C. 1911. On the biology of black mycetoma, Indian Med. Gas. 46: 376-378. — , 1911. On the cultivation of black variety of mycetoma, Centralhl. BaJct. I, 61: 358- 365, Pis. 1, 8. TORULEAE 689 Chmielewski. Paul. 1921. Contribution a 1 'etude des mycetomes de I'afrique du nord et du diagnostique general des Mycetomes, These Univ. Alger. Fac. Med. Pharm. 16: 1-64. Ciferri, Eafaelle and Bailey K. Ashford. 1929. Two strains of Pullularia puUulans (de Bary) Berkhout isolated from the human skin, Porto Eico Jour. Public Health Trop. Med. 5: 188-195. Clamann. 1924. Charakteristisdie Pilzformen (BefJillungspilze, Schwarzepilze) auf der Haut der Landbevolkerung, Derm. Woch. 78: 584-586, ^ figs. Clemow, Frank G. 1906. Mycetoma (Madura foot) in the Yemen, Brit. Med. Jour. 1: 918, 919, 3 figs. Collaa. 1883. La maladie de ballingall (pied du Madure) d'apres des notes inedites du docteur Collas ... a Pondichery mis en ordre completees et annotees par ... A. Corre, Arch. Med. Nav. 39: 81-137, 204-224. Coquerel, Charles. 1866. Note sur I'examen microscopique des lesions que Ton observe dans 1 'affection connue sous les noms de perical, pied de Madura, Mem. Soc. Biol. 17: 191-196. Corre, A. 1883. See also Collas. 1883. Cunningham, D. D. 1895. Is mycetoma primarily owing to the action of tlie fungal elements ordinarily associated with its products? Sci. Mem. Med. Off. India 9: 67-74, Pis. 1, 2. Delamare, G. & C. Gatti. 1929. Hyphomycete cultivable a grains blancs reniformes et durs, Indiella americana, C. B. Acad. Sci. 188: 1264-1266, S figs. — . 1931. Mycetoma du pied a grains noirs, Bull. Soc. Path. Exot. 24: 80-84, S figs. Etienne, G. 1920. Mycoses et mycetomes, Bev. Med. de I'Est (Nancy) 48: 1-11. Falchi, G. 1925. Onicomicosi da Hemispora stellata, Giorn. Ital. Derm. Sifilol. 66: 650- 656, 1 pi. Ferrari, Angelo. 1932. Ricerche sul Cryptococcus metaniger Cast., Atti 1st. Bot. B. Univ. PoAjia IV, 3: 175-184, 3 figs. Fonseca Filho, Olympio Oliveira Eibeiro da. 1930. O genero Madurella e os mycetomas produzidos pelas especies nelle contidas, Bev. Med. Cirurg. Brasil 38: 262-269, 2 figs. — . 1930. As chromoblastomycoses, Bev. Med. Cirurg. Brasil 38: 197-216, 3 pis. [English translation 216-236.] Fonseca Filho, Olympio Oliveira Eibeiro da % A. E. da Area Leao. 1927. Contribuigao para 0 conliecimento da Hemispora stellata, Sciencia Med. 5: 584-587, 1 pi., 2 figs. [A contribution to the knowledge of Hemispora stellata, Ihid. 5: 587, 588.] — . 1927. Contribution a I'etude d 'Hemispora stellata, C. B. Soc. Biol. 97: 1790-1792, 2 figs. Fonseca Filho, Olympio Oliveira Eibeiro da, A. E. da Area Leao & J. J. C. Nogueira Penido. 1927. Mycose de type ulcero-nodulaire, semblable a la sporotrichose et produite par Hormodendrum Langeroni, C. B. Soc. Biol. 97: 1772-1774, 2 figs. Fonseca Filho, Olympio Oliveira Eibeiro da &t A. Ferreira da Eosa. 1930. Sobre a 'keratomy- cosis nigricans palmaris, ' Bev. Med.-Cir. Brasil 38: 337-344, 1 pi. — . 1930. Sur Cladosporium Wernecki et la keratomycose nigricans palmaire, C. B. Soc. Biol. 105: 785, 786. Fox, Tilbury. 1870. Fungus foot of India, Trans. Path. Soc. London 21: 411, 412. — . 1871. Fungus foot of India [pathology by J. S. Bristowe], Trans. Path. Soc. London 22: 320-326, Figs. 29-32. — . 1876. The so-called "Fungus-foot" of India, Lancet 1: 190, 191. Froes, Heitor Praguer. 1930. Do mycetoma pedis no Brasil . . . Monographia contendo varios quadros e mappas estatisticos e documentada con 60 microphotograpliias ori- ginaes, alem de 36 photograpliias diversas, algumas tambem originaes e em grande parte ineditas. A Nova Graphica, Bahia, 300 pp., 47 pis. — . 1931. Mycetoma pedis (Madura foot) and its incidence in Brazil, Jour. Trop. Med. Hyg. 34: 376-378, 2 pis. ■ — , 1932. Mycetoma pedis ou Mycetoma podal, Jornal dos Clinicos, 3-17. 690 MEDICAL MYCOLOGY Fulleborn, F. 1911. Madurafuss aus Deutsch-Siidwestafrica, Arch. Schiffs-Tropenhyg. 1: 131-133. Gammel, J. A. 1927. The etiology of maduromycosis, Arch. Derm. Syphilol. 15: 241-284, 20 figs. Gammel, J. A., Hagope Miskdjian & Harvey S. Thatcher. 1926. Madura foot (mycetoma), the black grain variety in a native American, Arch. Derm. Syphilol. 13: 66-77, 6 figs. *Gastaminza. 1929. Med. de los Paiscs Calidos 2: 445. Gelonesi, Gregorio. 1927. Due nuovi parassiti del piede di Madura. Studio sui micetomi della Somalia meridionale, Ann. Med. Nav. Colon. 33: 283-308, 8 figs. Ouiart, J. 1920. Considerations sur le mycetome a propos d'un cas nouveau, C B. Soc. Biol. 83: 277, 278. Hatch, William K. & F. L. Childe. 1894. A remarkable case of mycetoma, Lancet 72: 2: 1271-1273. Hirsch, August. 1863. Der Madura-Fuss. Ein Beitrag zur geschichte des pflanzlichen Parasitismus, Arch. Path. Anat. Physiol. [Virchow] 27: 98-115. *Hoffmann, W. H. 1928. La cromoblastomicosis en Cuba y la enfermedad de Guiteras o "Chappa," Bev. Med. Cui. 39: 420. Hogg, Jabez. 1872. Mycetoma: the fungus-foot of India, Monthly Microsc. Jour. 7: 98-100. — . 1872. Fungus-foot disease of India, Trans. Path. Soc. London 23: 294-297. Horta, P. See Parreiras Horta, P. Huntly, William. 1889-1890. Case of Madura foot in its initial stage, Glasgow Med. Jour. 32: 344-346; 33: 339-342. Jeanselme, E., L. Huet & Lotte. 1928. Nouveau type de mycetome a grains noirs du a une Torula encore non decrite, Bull. Soc. Frang. Derm. Syphiligr. 35: 369-375, 3 figs. Jones, Jack W. & Herbert S. Alden. 1931. Maduromycotic mycetoma (Madura foot). Re- port of a case occurring in an American negro, Jour. Amer. Med. Ass. 96: 256-260. Kemper, G. W. H. 1876. A case of podelcoma: with microscopic examination of the dis- eased structure by Henry Jameson, Am. Practitioner 14: 129-135. Kobner, 1891. Pilzpraparat von Madurafuss (Mycetoma pedis) aus Italien, Berlin. Klin. Week. 28: 132. Kyriasides, K. N. 1929. Die in Griehenland boi Madurafuss (Mycetoma pedis) gefundenen Pilzarten, Arch. Schiffs-Tropenhyg. 33: 656-660. — . 1930. Champignons des mycetomes observes en Grece, Ann. Parasitol. Sum. Comp. 8: 194-200. Langeron, Maurice. 1928. Mycetome a Torula Jeanselmei Langeron, nouveau type de my- cetome h, grains noirs, Ann. Parasitol. Hum. Comp. 6: 385-403, 12 figs. Langeron, Maurice So P. Parreiras Horta. 1922. Note complementaire sur le Cladosporium Wernecki Horta 1921, Bull. Soc. Path. Exot. 15: 381-383. Laveran, Alphonse. 1902. An sujet d'un cas de mycetoma a grains noirs. Bull. Acad. MM. Paris III, 47: 773-776, 1 fig. LeDantec, A. 1894. iKtude bactcriologique sur le pied de Madura du Senegal (variete trufPoide), Arch. Med. Nav. Colon. 62: 447-454. Mackenzie, K. W. 1911. A case of Madura foot, Indian Med. Gas. 46: 378-380. *McMurray, W. & Norman Paul. 1914. Tinea albigena, a fungus near Trichophyton albiscicans Nieuwenh, Australasian Med. Cong. 10th Sess. Auckland, 629. Madden, Frank Cole. 1902. Two cases of the pink variety of mycetoma. Jour. Trop. Med. Hyg. 5: 243, 244. Maffei, L. 1931. Micosi causata da una varieta di Halobyssus moniliformis Zukal, Atti Jst. Bot. B. Univ. Pavia TV, 3: 19-28, 10 figs. Magalhaes, Pedro Severiano de. 1921. Hyphomycetoma (nova mycosa) Rio Janeiro 1914, CentralU. Bakt. 72: 94, 95. TORULEAE 691 Maitland, J. 1898. Case of mycetoma of the abdominal wall, Indian Med. Gaz. 33: 57-59, 1 fig- Mazza, Salvador & E. J. Canal Feijoo. 1931. Micetoma de granos negros por Madurella sp. del Chaco santiagueiio, Bol. Inst. Clin. Quirurg. Univ. Buenos Aires 6: 244-254, 10 figs. [6ta Reunion Soc. Argentina Patol. Beg. del Norte, Salta, 1930]. Minas, P. 1861. [Two] cases of tubercular disease of the foot, Indian Ann. Med. Sci. 7: 521, 522. Monnier. 1918. Au sujet d'un pied de Madura observe a Fort Dauphin, Madagascar, Bull. Soc. Path. Exot. 11: 407-416. Montpellier, Pierre Jean Marie, 1914. Les mycoses et les pseudomycoses du pied en Algerie, These Bordeaux 101: 90 pp. Montpellier Jean, P. Gouillon, & A. Lacroix. 1921. Nodule mycetomique du bras; inocula- tion par epine, Bull. Soc. Frang. Derm. Syphiligr. 28: 347-349. Musgrave, W. E. &, M. T. Clegg. 1907. The etiology of mycetoma: report of a case of the ochroid variety occurring in the Philippine Islands and caused by a new species of Streptothrix (Streptothrix Freeri), Philippine Jmir. Sci. B. 2: 477-511, Pis. 1-4. Newman, J. H. 1874. [Four] cases of Madura foot disease, Indian Med. Gas. 9: 322, 323. Nicolle, Charles & Georges Blanc. 1920. Sur les divers cas de mycetoma (pied de Madura) observes jusqu'a ce jour en Tunisie, Arch. Inst. Pasteur Tunis 9: 183-224, 7 figs. Nicolle, Charles & E. Pinoy. 1907. Sur la fructification des pathogenes a I'int^rieur meme des tissues chez I'homme, C. R. Acad. Sci. 144: 396, 397. — . 1908. Un cas de mycetome a grains noirs ; culture et inoculation experimentale. Bull. Soc. Path. Exot. 1: 95-99. Noc, F. & Jouenne. 1922. Les mycetomes a grains noirs du Senegal, Ann. Inst. Pasteur 36: 365-385. Onorato, Kaffaele. 1926. I micetomi in Tripolitania, Arch. Ital. Sci. Med. Colon. 7: 1-8, 33-50, 65-83, 97-110, 137-152. Oppenheim, Moriz. 1904. Die pathologisclie Anatomic des- indischcn Madurafusses (Mycetoma pedis), Arch. Derm. Syphilis 71: 209-242, Pis. IT -20. Palmer, F. J. 1926. A case of Madura foot treated by chemotherapy; apparent cure, Indian Med. Gaz. 61: 74. *Parreiras Horta, P. 1919. Urn novo mycetoma de graos negros produzido pela "Madurella Oswaldoi, n. sp. " A PathoJogia Geral, No. 1-4. — . 1921. Sobre um case de tinha preta e um novo cogumelo (Cladosporium Wernecki), Rev. Med. Cirurg. Brasil 29: 269-274. Patton, W. S. 1906. Mycetoma (Madura foot) in the Yemen, Brit. Med. Jour. 1: 1401. Pelletier, J. F.: 1906. Mycetome a grains rouges observe a Saint Louis (Senegal), Ann. Eyg. Med. Colon. 9: 578-580, 1 fig. Pena, Eaul. 1930. A propos d'un cas de mycetome a grains blancs observe a Assoniptioii (Paraguay) et produit par le Seedosporium apiospermum, C. R. Soc. Biol. 104: 689, 690. — . 1930. A proposito de um caso de mycetoma podal de graos brancos observado em Asuncion, produzido pelo Seedosporium apiospermum, Rev. Med. Cirurg. Brasil. 38: 142-145, 1 fig. [English translation 145-147]. Pinoy, E. 1912. Note, Ann. Derm. Syphiligr. V, 3: 341-343. — . 1913. Actinomycoses et mycetomes. Ball. Inst. Pastexir 9: 929-938, 977-984. Pirajd da Silva [Paulo]. 1918-19. Contribuigao a micologia parasitaria do Brasil. Duas novas especies de fungos productores de maduromicose, Mem. Inst. Butantan 1: 187- 207, Pis. 34-38. — . 1919. Duas novas especies de fungos productores de maduromycose no Brasil, Brasil Med. 33: 81-83. * — . 1922. Sobre uma nova maduromycose de graos brancos produzida pelo Indiella Brumpti, n. sp. Tesis Bahia. k 692 MEDICAL MYCOLOGY Plehn, A. 1927. Madurafuss (Mycetoma pedis), Hcmdbuch Path. Mikroorg. [KoUe & Wassermann, ed. 3] 5: 113-132, 1 pi, 4 figs. Polverini, G. 1903. Eicerche e ossevazioni sul piede di Madura, Lo Sperimentale 57: 659- 684. Pope, Benjamin T. & D. S. Lamb. 1896. Mycetoma, the fungous foot of India, N. Y. Med. Jour. 64: 386-388. Porcelli, Eodolfo. 1922. Hemisporosi cutanea. Dimostrazione della Hemispora stellata nei tessuti, Pathologica 14: 496, 497, 1 pi. — . 1922. Micosi cutanea da Hemispora stellata (Vuillemin). Primo caso in Italia, Giom. Ital. Mai. Yen. Telle 63: 698-709, 1 pi. Prado Valladares. 1916. Pityriasis nigra centro albicans (Pitirase negra de despigmentacao central), Brasil Med. 30: 137, 138, 1 fig. Prasad, K. 1906. Notes on a case of fungus disease of India (Mycetoma or Madura foot), Indian Med. Gas. 41: 139, 140. Puyhaubert, A. & Eobert Jolly. 1918. Note sur un cas de mycetome a grains noirs provoque par un champignon du genre " Madurella, " Arch. Med. Exper. 28: 441-445. — . 1919. Note sur un eas de mycetome k grains noirs. Bull. Soc. Path. Exot. 12: 57-60, Figs. IS. — : 1920. Note sur un cas de mycetome a grains noirs provoque par un champignon du genre Madurella, Bev. Med. de I'Est 48: 11-15, 6 figs. Eeenstierna, J. 1926. Eeproduction experimentale du mycetome. Arch. Inst. Pasteur Tunis 15: 101-107, 1 pl. Eeynier, Paul. 1906. Pr&entation d'un moulage, de coupes microscopiques et de photo- graphies d'un pied de Madura, Bull. Mem. Soc. Chir. Paris 32: 618-623, 5 figs. ♦ Eeynier, Paul & Emile Brumpt. 1906. Observation parisienne de Pied de Madura, Bull. Acad. Med. Paris III, 55: 709-723, 9 figs. Eietmann, Br. 1930. Note preliminaire sur une epidermomycose palmaire noire observe dans I'Etat de Baliia au Bresil. Btoll. Soc. Frang. Derm. Syphiligr. 37: 202-207, 5 figs. Eobin, Charles. 1863. Note sur le pied endemique de Madura, Gas. Med. Paris III, 18: 461. — . 1864. Note sur le pied endemique de Madura, C. K. Soc. Biol. 15: 34. Euelle, Edmond. 1893. Contribution a 1 'etude du mycetome. These Fac. Med. Pharm. Bor- deaux 6: 1-83. Sakurane, J. 1906. Ein Fall von Oidiomycosis der Haut und des Unterhautzellgewebe, Arch. Derm. Syphilis 78: 211-221. Sartory, Antoine, M. Meyer & Jacques Meyer. 1930. Contribution a 1 'etude des mycoses osseuses; trois cas d'osteites dues d'une part a 1 'actinomyces aster oides (Eppinger) et d 'autre part a 1' Hemispora stellata, Ann. Inst. Pasteur 44: 298-329, 8 figs. Sartory, Antoine, E. Sartory, Br. Eietmann «fc Jacques Meyer. 1930. Contribution a 1 'etude d'une epidermomycose bresilienne palmaire noire, provoquee par un Cladosporium nouveau, C. B. Soc. Biol. 104: 878-881. Semon, H. C. 1915. Mycetoma pedis, Brit. Jour. Derm. Syphilis 27: 299-303, PI. 8. Shattock, Samuel G. 1897, 1898. Mycetoma papiUomatosum, Trans. Path. Soc. London 49: 293-296, 1 pl.; also Brit. Med. Jour. 1: 622. Silva, Flaviano. 1929. Contribuigao para o estudo do mycetoma podal na Bahia, Sciencia Med. 7: 153-164, € figs. [Eev. Brosil Med. 43: 635, 636]. — . 1929. Tinea nigra (Cladosporose epidermica), Brasil Med. 43: 924-926. Silva, Piraja. See Piraja da Silva. Smyth, J. 1898. Notes on a case of mycetoma of the neck, Indian Med. Gas. 33: 56, 57, Ifig. Surveyor, Nusserwangi F.r 1892. Madura foot of India, Brit. Med. Jour. 2: 575, 576. Thompson, Harold L. 1928. The present status of mycetoma, Arch. Surgery 16: 774-788. Thompson, Harold L. & Kano Ikeda. 1928. Maduromycosis: fourth case reported in the United States, Arch. Surgery 16: 764-773, 10 figs. TORULEAE 693 Towey, John W., Henry C. Sweany & Willis H. Huron. 1932. Severe bronchial asthma ap- parently due to fungous spores found in maple bark, Jour. Amer. Med. Ass. 99: 453- 459, 7 figs. Vuillemin, Paul. 1906. Un nouveau genre de Mucedinees: Hemispora stellata, Bull. Soc. Myc. France 22: 125-129, PI. 7. — . 1919. Remarques sur les mycetomes: hommage a la memoire de M. [R.] Jolly, Arch. Med. Exp. Nat. Path. 28: 446-451; also Bev. Med. de I'Est 48: 15-20. Wright, James H. 1898. A case of mycetoma (Madura foot), Jour. Boston Soc. Med. Sci. 2: 128-130, 1 pi. — . 1898. A case of mycetoma (Madura foot), Trans. Ass. Amer. Physicians 13: 471-482, ^ fold. pis. — . 1898. A case of mycetoma (Madura foot). Jour. Exp. Med. 3: 422-433, Pis. 39, 40. *Yazbek, Alexandre Kalil. 1920. Do.s Mycetomas, These Inmigural Secgdo de Ohras do "Estado de Sao Podilo," 164 pp., ^6 pis. I CHAPTER XX ACTINOMYCETEAE The systematic position of this group of organisms has long been ques- tioned. Since the mycelium is usually much more slender than that of other groups of fungi and the arthrospores are often bacilliform and of approxi- mately the same magnitudes as bacteria, many bacterial systematists have included this order in the Schizomycetes, often adding Mycobacterium and Corynehacterium to the group. On the other hand, mycologists who have had occasion to work with the group (Bary, Thaxter, Sauvageau & Radais, Gasperini, Drechsler) have always included them in the Fungi Imperfecti along Avith the Hyphomycetes. The arguments of the bacteriologists have usually been based on the staining reactions, development of elavate elements in the animal body with Actinomyces, representing either the primitive stage from which Mycobacterium and Corynel>acterium form degeneration stages or the culmination from increased production of mycelium. Witter (1933) points out that he was unable to find either chitin or a definite nucleus in the species he studied. On the other hand, the mycologists have held that the spore production of Actinomyces is essentially similar to that of the conidial production of the imperfect fungi, which has nothing in common with bacterial structures, and that many of the apparent resemblances are the result of careless technic. Drechsler (1919) has perhaps given the most careful attention to the mor- phology of the group, and we owe much of our knowledge of the group to his work. It is to be regretted that his subsequent duties in the Department of Agriculture have prevented his continuing the study of this group and that he has been unable to turn his rich experience in the morphology of the group to the presentation of a systematic account. Only one who has worked Avith the group from the standpoint of its morphology for a considerable time is really competent to evaluate the conflicting accounts in the literature and to make the necessary assumptions and correlations in filling the lacunae of imperfect descriptions. In the following account, the material is frankly a compilation of the literature, in the hope that this may lead to a keener realization of the lacunae in our present knowledge. The mycelium of Actinomyces has very slender filaments, commonly 0.2- 1.0/i, in diameter, rarely up to about 1.5/a. It is generally sparsely and irregu- larly septate, occasionally somewhat regularly septate, but even then less so than in the other Fungi Imperfecti. Branching is common, probably never truly dichotomous, although in a few species it is impossible to be certain. Usually a lateral bud develops at some distance back of the growing tip and 694 k ACTINOMYCETEAE 695 may give rise to other branches by lateral proliferation. If these prolifera- tions start close to the tip so that it is difficult to decide which is the main axis, the branching is said to be dichotomous. The branches forming the periphery of the actively growing pellicle or the voiino: sporogenons branches attached at intervals to the superficial mycelium, are filled with a dense protoplasm which takes a deep homogeneous stain with hematoxylin. Nearer the origin of the hyphae the contents appear vacuolate. When the vacuoles are excessively large and extend throughout most of the cross-section of the cell, the cytoplasm is confined largely to the walls, leading some of the earlier writers to refer to the Aussenplasma and the Innenplasma. The presence of large vacuoles is often associated with local distentions of the hyphal wall, each swollen segment being largely oc- cupied by a single ellipsoid vacuole, separated from the vacuole of the neigh- boring distention by a protoplasmic partition at the constriction. In other species and in the nutritive mycelium generally there is no marked regularity in the alternation of inflated portions and constrictions, but pronounced devia- tions in the diameter of the filaments may occur with more or less variable frequency. Much importance has been attributed by earlier writers to a variety of abnormalities and products of degenerative changes occurring in the myce- lium. In the publications of Israel (1878), Johne and MacFadyean (1889), bodies described as micrococci, cocci, etc., were given minute attention and assigned an important role in the complex ontogeny ascribed to the parasite, supposed to be a pleomorphic bacterium. Wolff and Israel (1891) confused them with the spores of other authors, and as the structures did not have the heat resistance of bacterial spores, they questioned the formation of true spores by Actinomyces. Since they figure only sterile mycelium, it is probable that they were correct in their observations. Bostroem, on the other hand, had both true spores and endogenous granules, which he indiscriminately referred to as spores. Round granules, deeply stained in the living filament by very dilute methylene blue, are variable in size and have a method of multiplication and orientation related to the regions of growth in the mycelium. These were called nuclei by Neukirch. Schiitze, using Neukirch's methods, designated them as metachromatic granules. This material is easily distinguished by its powerful affinity for most of the ordinary laboratory stains. In material fixed in alcohol and stained with Delafield's hematoxylin, these granules retain the stain after it has been washed from all other portions of the cell. These granules are rare in the regions of active growth, but in the vacuolated por- tions they are found as minute bodies widely separated from one another. Farther from the tip of the hypha the granules increase in size and frequency, and their arrangement becomes more regular. The individual spherical bodies are nearly equal in size, exactly filling the lumen of the hypha, and are separated by nearly equal spaces. In other cases, the granules seem to coalesce 696 MEDICAL MYCOLOGY to form incomparably larger masses, making it difficult to believe that they have any relation to spores or to nuclei. Quite possibly this represents a waste product, since its abundance is always connected with advanced degeneration. While the mycelium is very uniform throughout the genus, there is very great diversity in spore formation. The spores are developed by a transfor- mation of more or less specialized hyphal branches, early distinguishable from sterile hyphae of the aerial mycelium. In general, the diameter of any por- tion of sterile mycelium is attained at the time it arises through the elongation of the growing hyphal tip. The sporogenous branches are, in the beginning. Fig. 112. — Actinomyces II isolated from soil. 1, portion of aerial mycelium, showing conspicuous septa in fertile branches and the relation of the latter to axial filaments ; 2-6, stages in the development of a fertile hypha (X2,750). (After Drechsler 1919.) conspicuously thinner than the axial hyphae from which they are derived. Later, when they have reached nearly their full length, they increase in thick- ness, the extent of this varying much with individual species. In most species the maturation of the sporogenous hyphae is associated with a peculiarity in growth by which they become coiled in more or less characteristic helices. The tendency toward a coiled condition is usually clearly manifest before the branch has grown to half its length through the open flexuous habit of the young filament. As elongation continues, the turns become increasingly definite, but the contraction leading to the final condition, which ranges from Actinomyces XIII, with its open, barely perceptible turns, ACTINOMYCETEAE 697 to one in which the helix is so strongly compressed that its adjacent turns are in continuous contact, is usually delayed until the later growth in thickness by the sporogenous filament. Specific differences may not only be indicated by the obliquity of the helix, but involve also the number and diameter of its turns and its construction with reference to a dextrorse or sinistrorse condi- tion. The range in different species extends from two or three turns in Actinomyces II (Fig. 112) to over twenty turns in others, the range in an individual species being much smaller. Coils of over twelve turns are very rare. The diameter of the helix is more or less inversely proportional to the number of turns characteristic of the species. Rotation in the formation of the helix is specifically sinistrorse or dex- trorse in different species, the sinistrorse condition being the more common as is general in plants. For a given species the condition of rotation of the helix is constant. Two main types of branching of sporogenous filaments from the main axis are evident ; the erect or dendroid type, in which the sporogenous hyphae are successively developed, and the prostrate racemose type, in which develop- ment is more or less simultaneous. In the erect type, the development of the fructification starts from a single erect hypha with a helical termination. Sporogenesis starts at the tip by the insertion of regularly placed septa and proceeds downward toward the base of the filament. Usually, before much of the hypha has been involved, a single septum will appear well toward this base, and immediately below it a bud of a new sporogenous hypha appears. As the latter is attaining its growth in length and thickness and its helical disposition, the basipetal septation in the axial filament proceeds to the septum above the insertion of this first branch, the young spores thus delimited un- dergo maturation processes, the helix becomes relaxed and the chain of spores subject to disruption. The branch now passes through the same stages of development as the axial hypha and in turn gives rise to a sporogenous branch below a septum a little above its own insertion. The number of sporogenous branches developed below a single septum is generally increased by subse- quent proliferations, and the initiation and development of successive orders may be indefinitely repeated. Complex fructifications are frequently developed in which a succession of the processes described are occurring simultaneously at many points. In the second type, there is no such clearly defined relation between younger and more mature sporogenous hyphae. Development of a fructifica- tion is initiated by the proliferation of branches at irregular intervals on the distal portion of a prostrate axial filament which often exceeds 1 mm. in length (Fig. 113). The branches may either cease their more extensive development after forming a helix or themselves proliferate a secondary branch a short distance above their own insertion, and this in turn may form a helix and give rise to a tertiary branch. By repetition of this process, each lateral ele- 698 MEDICAL MYCOLOGY ment may become branched several times, the whole apparatus as well as its insertion on the axial filament being characterized by an absence of septa. Sporulation, instead of beginning- in any individual helix as soon as it is formed, is usually delayed until the branching and growth of helical hyphae in the same lateral process have come to an end, when it will often proceed rapidly and almost simultaneously in all the helices. The termination of the axial filament itself develojDS into a helix and behaves essentially like a primary lateral branch. Occasionally, the axis of one of these racemose arrangements may be comparatively short, resulting in a rather intricate structure where one lateral branch may be entangled with another. The tendencies characteristic of the type, namely, the absence of a septum above the insertion of branches, and Fig. 113. — Actinomyces XVIII. Portion of fructification bearing aberrant fertile branches with- out spiral terminations (X2,750). (After Drechsler 1919.) the delay in sporulation in the helices first formed, are maintained, however, until the growth of the last order of sporogenous branches is more or less complete. Besides species in which these two types are distinct, there are a large number of species in which there is a combination of these two methods. Frequently the open racemose arrangement of the lateral branches on the main axial filament is associated with a successive order of development in the further ramification. The presence of a septum above the insertion of a branch is characteristic of more species than is its absence, and in some species both conditions prevail. In a few species there are formed, besides the more regular fructifications, others with relatively thick branching axial hyphae which are densely filled with protoplasm and bear, at very close and irregular intervals, a short thick AGTINOMYCETEAE 699 unbranched sporogenous hyplia with little or no helical modification. This seems to be associated with excessively rapid growth (e.g., Actinomyces alhus, Fig. 114). The prevailing idea, that most of the mycelium is converted into spores, is incorrect. Spomlation is strictly confined to terminal elements, never as a rule passing beyond the first junction with another element (Fig. 115). The proliferation of a branch nearest the end of the axial filament limits spore production in this filament to the portion beyond the insertion of the first Fig. 114. — Actinomyces albus (A. griseus Krainsky ?). 1, 2, portions of aerial mycelium; 4, germinating spore (X2,750). (After Drechsler 1919.) branch ; in the same manner, the proliferation of a secondary from a primary lateral branch results in a sterilization of the portion of the hypha below the insertion of the new branch. In Actinomyces Y, sporulation is even further restricted by the apparent abortion of a number of potential spores at the proximal end of the unbranched lateral branches. The hj-phal portion which is involved first develops as usual, but when the characteristic septation asso- ciated with the delimitation of spores in this species appears in the helix, it is not extended to the base of the branch, although indications of regularly spaced membranes may usually be distinguished. Later, the unsegmented 700 MEDICAL MYCOLOGY portion is gradually evacuated and converted into a sterile stalk devoid of protoplasm. It is interesting to note that the basal septum which in an allied and very similar form, Actinomyces VI, delimits the lowest spore from the axial filament, here also is present as a well-developed septum. The delimitation of the ultimate cells in the process of sporulation occurs usually as the growth in thickness and the contraction of the helix are ap- proaciiing completion. It has usually been believed that the details connected with the spore formation are uniform throughout the genus, but this is not universally true. In most species, the sporogenous hyphae become divided into regular cylindric cells separated by septa ; the latter generally stain deeply with Dclafield^s hematoxylin, probably as a result of an association with metachromatic or possibly nuclear material. These species may be subdivided into three groups. Fig. 115. — Actinomyces XVIII. Degenerate axial filament containing large vacuoles and spherical structures and bearing a fertile helix (X2,750). (After Drechsler 1919.) In the first group (e.g., Actinomyces /), the cross walls in the sporogenous hyphae remain without any very pronounced change, continuing to separate the adjacent cells until these have developed into a chain of mature contiguous spores. The insertion of these septa progresses from the tip toward the base and does not break the physiologic continuity of the hyphae, for food material apparently is transmitted through them to the young spores at the termina- tion, since these subsequently increase in size and may deposit a wall of measurable thickness. In the second group, the septa split into halves, which are then drawn apart by the longitudinal contraction of the individual protoplasts. In Actinomyces II, the very pronounced growth in thickness of the sporogenous hyphae, following the insertion of septa, indicates that in this species also septation brings about no impediment in the transfer of food material. This is remarkable in view of the extraordinary thickness of the septa character- ACTINOMYCETEAE 701 izing this species. Actinomyces scabies probably represents the more usual condition where the segment of tlie hyphal wall evacuated by the contraction of each two successive spores undergoes no change until fractured by the disruption of the chain of mature spores. In the third group, as in Actinomyces aureus, the septa first undergo a deep constriction which, by involving the ends of the young cylindric spores, Fig. 116. — Actinomyces aureus. 1-5, stages In the development of a sporogenous hypha ; 6, erect fructifications terminating long, prostrate aerial hyphae, exhiibiting a pronounced tend- ency toward the successive type of development and showing cuneate hyphal enlargements below the Insertions of branches (X2,750). (After Drechsler 1919.) gives to the latter an elongated ellipsoid shape (Fig. 116). The constricted septum now gradually loses its staining properties and appears to be slightly drawn out in a longitudinal direction. A preparation stained with Delafi eld's hematoxylin usually shoAvs many old spore chains in which the individual 702 MEDICAL MYCOLOGY spores are thus connected by hyaline isthmuses. Occasionally an isthmus may be found with a remnant of the old deeply staining septum still unchanged in its center. Besides these three, there is another group in which the septa are not to be demonstrated by the ordinary stains. The protoplast appears to contract at regular intervals, yielding a series of noncontiguous spores, held together for a time by the connecting segments of the evacuated hyphal wall (Fig. 117). Drechsler believed that cross walls appear in the development of sporogenous hyphae throughout the genus but are too thin to be seen in these forms. Owing to the very small size of the cells, it is not always possible to dis- tinguish nuclei from metachromatic granules. The spores germinate in dilute nutrient solutions by swelling and emitting 1-4 germ tubes, the number being quite characteristic for each species. Fig. 117. — Actinomyces Lavendulae. Portion of aerial liypha (X2,750). (After Drechsler 1919.) Methods for Study. — Drechsler recommends the following methods as be- ing especially suitable for studying Actinomyces. The fungus is grown on a suitable substratum, such as potato or glucose agar. Growth on potato agar is more prompt and productive of mycelium on most species; but as its use, especially with species exerting a strong tyrosinase reaction, stimulates to excessive guttation and disruption of the sporophores by the extruded drop- lets, a medium not possessing this property is advantageous. After the cul- tures have attained the proper degree of maturity, the whole growth is cut from the agar and removed from the tube as carefully as possible. A slide, smeared with albumin fixative, is now brought into firm contact with the mycelium and then separated, precautions being taken to avoid any sliding of the two surfaces on each other. If the growth is not too young, this pro- cedure will leave the upper portions of the aerial mycelium adhering to the slide without serious disarrangement, and killing and fixation may be at once effected by the use of strong alcohol. The material is subsequently stained and mounted in balsam. The quality of preparations in which the spore chains ACTINOMYCETEAE 703 have begun to disintegrate in large numbers is impaired by tlie presence of large masses of free spores which retain their staining properties for some time after maturing. Later the spore walls seem to become entirely impervious to stains and, as a result, when the secondary mycelium develops, no difficulty is encountered from this source beyond a slight clouding effect. The best results are obtained when the print is made soon after the mycelium begins to adhere readily to the smeared slide. The nature of the killing agent em- ployed was found to have no noticeable effect on the preparation. Flemming's weak and strong, picroformal, picro-acetic, Camoy, and 95% alcohol were tried. To save time in washing, alcohol is most frequently employed. Delafield's hematoxylin, which is the most satisfactory stain, is allowed to act 24 hours and then the preparation is decolorized. Vacuoles, metachromatic and nuclear structures, as well as septa, show clearly. Potron & Thiry (1913) recommend the following toluidine blue stains in pus from a case of actinomycosis. The smear is dried, fixed in the flame, cooled. A small amount of the powdered dye is placed on the smear and a drop of water added. After the stain has acted a short time it is rinsed off, and the preparation is dried and mounted in balsam, or the toluidine blue may be rinsed with a solution of 1% eosin in 90% alcohol to partially decolorize, then rinsed in water, dried, and mounted. Or the slide may be left in Lugol 's solution a few minutes before decolorizing with alcoholic eosin. Still another variation consists of partial decoloration with aqueous orange " de Gole." This order is composed principally of saprophytes living in soil, more rarely of facultative parasites of plants, e.g., A. scabies on the tubers of the potato (Pig. 118), or parasites on man, where a few species are fairly common and well known, and many have been briefly described with little regard for their morphology and little data regarding their physiology which would en- able them to be recognized when they are again encountered. The physiology of this group has been extensively studied for the sapro- phytic species by Lieske (1921), and especially the soil organisms by Waks- man and coworkers during the last decade. The systematic position of Actinomyces has long been subject to debate, many of those working with pathogenic species placing them among the pleomorphic bacteria, some, e.g., Lieske (1921), 0rskov (1923), and Jensen (1932), placing them in a separate group intermediate between bacteria and fungi; and mycologists, such as Saccardo, Thaxter, and Drechsler (1919) plac- ing them in the Hyphomycetes. In this connection, it is interesting to note that Lieske states that they are very closely related to Geotrichum candidum (Oidium lacfis), an undoubted member of the Hyphomycetes. Claypole (1913) regarded them as an ancestral type of microorganism, giving rise to yeasts and higher filamentous fungi and to mycobacteria, corynebacteria, and the other bacteria. While most authors have kept the majority of the organisms of this group in a single genus, various attempts have been made to erect more or less 704 MEDICAL MYCOLOGY natural groups of species. Wright (1905) would reserve Actinomyces for anaerobic organisms, placing the aerobic forms in Nocardia. This separation has been widely adopted with varying nomenclature, but is often difficult to apply in practice. I have used this character in my key, although I have kept all the species in Actinomyces. Chalmers & Christopherson (1916), followed Fig. 118. — Acivnomyces scabies (Thaxter) Gussow. 1-6, successive stages In the develop- ment of sporogenous branch ; 7, portion of aerial mycelium, some lateral elements bearing secondary branches developed successively, with an unusually long chain of spores (1-6 X 8,000 ; 7 X2,750). (After Drechsler 1919.) by Castellani & Chalmers (1919) and Froilano de Mello and coworkers (1918, 1919) have divided the genus into saprophytic and parasitic species. While this might be useful, if only the obligate parasites are included in the parasitic ACTINOMYCETEAE 705 groups, it seems extremely artificial as they use it. Their next subdivisions, which are based on comparative pliysiology and staininji' reactions, are more natural and have been followed in my key. Lieske (1921) provided much information but did not attempt any classi- fication. 0rskov (1923) proposed to redefine Cohnistreptothrix to include organisms which form a unicellular, nonseptate vegetative mycelium, and an aerial mycelium composed of hyphae, thicker than those of the vegetative mycelium and divided into spores of uniform size and shape. He retained Actuiomyces for organisms in which the vegetative as well as the aerial myce- lium divide by septa into pieces of irregular size and shape without any spores, as in hi.s Cohnisireptoihrix. He created a new genus, Micvomonospora, for organisms which form a unicellular mycelium without aerial hyphae but with spores borne singly on the tips of short branches of the vegetative hyphae. This classification by 0rskov is a step in the right direction, but it is obviously impossible to apply it to the pathogenic species in the present state of our knowledge, almost none of which have been described morphologically with sufficient care to place them. 1 suspect that when the morphology has been carefully studied, the Majoves of Chalmers & Christopherson will be found to belong in Cohnistreptothrix of 0rskov not of others, their Minores in his Actinomyces, and their Breves in his Micvomonospora with a certain amount of redefinition and readjustment. Jensen accepts 0rskov's classification for his soil organisms, adding to the nomenclatural confusion by replacing Cohnistreptothrix 0rskov by Actinomyces, and Actinomyces 0rskov by Proacti- nomyces. He includes Corynehactevium, Mycohacterium, and his Proactinomyces in his Proactinomycetaceae, leaving Actinomyces and Micvomonospora for his Actinomycetaceae. While is it quite possible that Corynehacterium and Mycohacterium may sometime be shown to belong to this group, the literature on these genera is so extensive and well known to the medical man that no attempt will be made to cover it in this work. Since so little morphologic work has been done on this group, it seems wiser to leave all the species in Actinomyces than to attempt a separation. Cohnistreptothrix has been used for the anaerobic species, but in the present chaotic state of the literature it is difficult to separate strict anaerobes from partial anaerobes. ActinobacilUis is recognized by Buchanan, Bergej^ and others as a distinct genus. Its morphology needs further study before its systematic position will be clear. Malhranchea was proposed many years ago and recently revived by Vuillemin. It is not distinguishable from many species of Actinomyces, in fact it shows the characteristic morphology of that genus. ACTINOMYCES Actinomyces Harz, Jahresber. Miinchener Central-Thierarzneischule 1877: 125-140, 1877. Streptothrix Cohn, Beitr. Biol. Pflanzen 2: 186, 1875, non Corda, Pracht- flora Europ. Schimmelbildungen 27, 1839. 706 MEDICAL MYCOLOGY Cladothrix Cohn, Beitr. Biol. Pflanzen 2: 185, 1875, non DeCandolle 1849. Discomyces Rivolta, Clin. Vet. Milano 1: 208, 1878. Malhranchea, Saccardo & Penzig, Michelia 2: 639, 1882. Nocardia Trevisan, I genera e le specie delle Batteriacee. 9, 1889. Micromyces Gruber, Centralbl. Bakt. 10: 648, 1891, not Dangeard, 1888. Oospora Sauvageau & Radais, Ann. Inst. Pasteur 6: 242-273, 1892, non Wallroth, 1833. Cohnistreptothrix Pinoy, Bull. Inst. Pasteur 11: 929, 1913. The type species is Acti7wmyces hovis Harz. Mycelium usually branched, very slender, usually less than 1/a in diameter, seldom more than 1.5/a, probably septate, but the septa are often very difficult to observe; conidia are borne on specialized conidiophores, usually in coiled chains which are very fragile and break up so easily that they are difficult to observe. Primarily soil saprophytes; a few species are parasitic on ani- mals, and still fewer on plants. Key to Species Cultures unknown. Seen in juxta-articular nodules. A. Carougea/ui. Seen in calcareous deposits in muscles of swine. A. musculorum. Cultivated. Obligate anaerobes. Bacillary forms only seen in cultures. Growth on gelatin which is liquefied; from dacryocystitis. A. Silherschmidti. Culture difficult, no liquefaction of gelatin; ulcers on muzzles of rabbits. A. cuniculi. Hyphae seen in cultures. No growth on gelatin or potato, but good on ascitic agar. Isolated from pus. A. Neschcsadimenhi. Isolated from axillary and pubic hair. A. tenuis. Good growth on gelatin. Gelatin liquefied; isolated from blood in generalized infection. A. Thjoettae. Gelatin not liquefied; calf diphtheria. A. necroplwrus. Facultative aerobes, growth better under partial anaerobic conditions. Growth on usual media, clavate bodies present. Not pathogenic for laboratory animals, from concretions in lacrimal canal, colonies grayish. Growth on potato, starch digested. A. Foersteri. No growth on potato, starch not digested. A. discofoliatus. Pathogenic for laboratory animals. Colony yellowish; human actinomycosis. A. Israeli. Colony white; subcutaneous and intramuscular nodules. A. Thihiergei. Growth absent on usual media, gram-negative. Colony translucent on ascitic agar; isolated fiom pubic and axillary hair. A. tenuis. ACTINOMYCETEAE 707 Colony opaque. Not pathogenic for laboratory animals; isolated from calves with pneu- nioiiia. A. actinoides. Pathogenic for guinea pig; isolated from liver abscesses and pulmf)nary infection. A. americanus. Patliogenicity not stated; isolated from mycetoma of the outer ear. A. cylindraceus. Aerobic, at least growth much better under these conditions. Gelatin liquefied, often serum also, not acid-fast, good growth on potato, diastase often present, odor earthy or moldy, efflorescence spreading, bright, chalky, growth good at 22° and 37° C, branching abundant in mycelium. Serum not digested or action on serum unknown. Colonies deep green; from soil, pathogenic for experimental animals. A. viridis. Colonies deep red with white margin on potato, white on agar; from dermatosis. A. panginensis. Colonies red on agar and potato; isolated from sputum, not pathogenic for experimental animals. A. mineaceus. Colonies yellow to brown. No pellicle on broth; from corneal infection. A. cerehriformis. Pellicle often fragile on broth. Growth on potato; tumors in domestic fowl. A. tossicus. No growth on potato; from lesions on sheep, producing lumpy wool. Colonies white on most media. ^- dermatonomus. Milk not coagulated. On milk, growth slow, digestion in three weeks; from mycetoma pedis. A. mexicanus. On milk, forming a flesh-colored, folded pellicle with white ef- florescence, only fat assimilated; from bronchopulmonary infection. A. Eivierei. Milk coagulated. Growth on milk not stated, but no pellicle on other liquid media; from enlarged spleen. A. Gibsoni. No pellicle on liquid media, colony white becoming cafe-au-lait on potato; from black tongue. A. Guegueni. Pellicle on vegetable decoction only; colony chestnut color on potato; from erythrasma. A. Pinoyi. Pellicle on most liquid media (saprophytes). Colony white on potato, medium colored yellowish or brownish. A. goensis. Colony yellowish waxy on potato, medium not colored. A. Christophersoni. Pellicle on milk folded, dark green; from air, pathogenic for experimental animals. A. pyogenes. Pellicle on milk yellowish. From sputum. A. gypsoides. From water and laboratory contamination. Serum digested. ^- invulneralilis. Colony white or grayish on most media. Colony rose, cerebriform on glycerol agar. A. roseus. 708 MEDICAL MYCOLOGY Colony white, from sputum. A. candidus. Colony white, with an amber yellow margin, medium discolored ; from pseudotuberculosis. A. albus. Colony grayish, medium discolored; from abscesses. A. Garteni. Colony white becoming greenish on old cultures. From mycetoma. A. Nicollei. From conjunctivitis. A. Dassonvillei. Colony grayish, medium not discolored; secondary invader of ring- worm lesions. A. minimus. Colony grayish, star-sliaped or cruciform ; from infection of cornea. A. radiatus. Colony yellowish to brownish on most media. Pellicle on liquid media, milk coagulated. Colonies grayish violaceous with reddish brown margin; from an- gina. A. rubidauretis. Colonies white, later brownisli; from mycetoma. A. Avadi. No pellicle on liquid media. No growth on potato; from pus. A. Levyi. On potato, gray to canary yellow; medium darkened; lumpy jaw. A. bovis. On potato, straw yellow, medium not darkened, grows with dif- ficulty; from mycetoma. A. liquefaciens. On potato, yellow, thick, verrucose; from conjunctivitis. A. aureus. On potato, cafe-au-lait with no pigmentation of medium; from conjunctivitis. A. luteolus. Gelatin usually not liquefied ; scrum not liquefied, usually acid-fast, good growth on potato, but no diastatic action, odorless or nearly so; efflorescence cir- cumscribed, dull, powdery, growth good at 37°, fair at 22° C. ; hyphal branching poorly marked. No growth on potato, or growth very poor. Growth on gelatin. Colony white on gelatin. A. gedanensis. Colony grayish on gelatin; cacao brown on carrot; from sputum. A. catarrhalis. No growth on gelatin; from roots of decaying teeth. Sediment in broth yellowish, not acid-fast. A. Matruchoti. Sediment in broth white. A. Rodellae. Growth good on potato. Colonies white, pellicle on broth. Action on gelatin unknown, milk digested ; from saliva of horse. A. Chalmersi. Gelatin slowly liquefied; mycetoma pedis. A. Tarozzii. Gelatin not liquefied; mycetoma. A. Ribeyroi. Colonies pink or reddish, not yellowish or brownish. Pellicle on broth or other liquid media. Dissolving the horny layer of the skin in grooves (keratoderma plantare sulcatum). A. keratolyticus. Pathogenicity uncertain, salmon pellicle on milk; from blood of horse. A. pluricromogerms. ACTINOMYCETEAE 709 Producing mycetoma pedis. No growth on milk; color brick red. A. micetomae. Milk not coagulated; salmon color. A. verrucosus. Milk slowly peptonized after twenty days. Acid-fast. A. Sommeri. Not acid-fast. A. Madurae. Growth on milk not reported, colonies deep red. A. iahiensis. No pellicle but some floating colonies which sink before forming pellicle; from lungs. Colonies coral pink. A. Leishmani. Colonies flesh or orange red. A. carneus. Growth on liquid media unknown. Colonies wine red; from erythrasma. A. minutissimus. Colonies white with salmon mammillae; from case resembling bu bonic plague. A. Jollyi. Colonies with yellow center and pink periphery; from mycetoma pedis. A. Freeri. Colonies yellow to brown on most media, rarely also reddish on some one or two media. No pellicle on liquid media. No growth on gelatin; from sample of vaccine, pathogenic for experimental animals. A. Hofmanni. Growth on gelatin. Substrate blackened on sugar media ; from air. A. niger. Substrate not blackened. Gelatin very slowly liquefied. Growth slow on potato; from mouth. A. lingualis. Growth good on potato ; from granuloma. A. phenotolerans Growth good on potato ; from cereal dust. A. BelUsari. Gelatin not liquefied. Serum liquefied; mycetoma. A. convoluUos. Serum not liquefied. Broth colonies not described; actinomycosis of bones. A. serratus. Gas produced on broth ; pulmonary infection. A. Donnae. No gas on broth; colony on potato reddish; corneal ulcers. A. deBerardinis. Floating colonies which are not confluent. Gelatin slowly liquefied; cattle farcy. A. farcinicus. Growth poor in gelatin; lungs. A. japonicus. Pellicle on surface of liquid media. Milk coagulated; mycetoma. A. pretorianus. Milk digested without coagulation ; lumpy jaw of marsupials. A. macrodipodidarum. 710 MEDICAL MYCOLOGY Milk not coagulated or digested. Pellicle pink in milk. On dog. A. canis. On goat. A. caprae. Pellicle on milk yellowish. On potato, colony white, granular; brain and bronchial abscesses. A. hicolor. On potato, colony orange yellow; mycetoma. A. hrasiliensis. On potato, colony yellowish red or brick color. Pellicle on broth waxy and fragile; from pseudo- tuberculosis with cerebrospinal meningitis. A. a^ter aides. Pellicle on broth thick yellowish; from rat. A. Sanfelicei. Often slight indications of liquefaction of gelatin and serum, rarely acid-fast, growth difficult on most media at 37° C, none at 22° ; little or no growth on potato, no diastatic action, odor sometimes feculent, slight or no efflorescence, hyphal branching rare. Growth on gelatin. Eed pigment diffusing into gelatin; from sputum. A. spumalis. No red pigment diffusing into gelatin. Colony yellowish. A. fiavus. Colony white; lumpy jaw. A. Spitsi. No growth on gelatin. No growth on agar, sometimes on blood agars and similar media. Growth on potato white with brown efflorescence; bronchopneumonia. A. cruaris. No growth on potato. Colonies red or orange; mycetoma pedis. A. africanus. Colonies not red. No growth on carrot, growth on serum, A. Berestneffi. A. Ponceti Growth on carrot, probably none on serum; from stomatitis. /-, ,, A. huccalis. Growth on agar. Colonies yellowisli. No growth on potato; from abscess on jaw. A. Krmisei. Little growth on potato, not acid-fast; from sputimi in bronchitis. A. Fijperi. Good growth on potato, acid-fast ; pseudotuberculosis. A. Sartoryi. Colonies white. Pellicle on liquid media. Growth on potato; mycetoma pedis. A. transvalensis. No growth on potato. Eed brown pigment diffusing into medium ; from sputum. A. fuscus. Pigment not diffusing into medium, endocarditis of cattle. A. valvulae. b ACTINOMYCETEAE 711 No pellicle on liquid media. Slight fermentation of sugars; bronchomycosis. A. hrcmchialis. No fermentation of sugars; pulmonary infection. A. pulmonalis. No fermentation of sugars; nodules on jaw of horse. A. equi. Growth on gelatin not reported. Gram-negative; from blood, fever following animal bites. No growth on common media, slight growth in brotli to which patient 's blood was added. A. Taraxeri-cepapi. Growth on blood agar and blood serum. A. Putorii. A. muris-ratti. Gram-positive or staining not given; growth on agar without addition of blood. From abscesses. Clavate formations in broth; from tongue abscesses, pathogenic for rabbits. A. londinensis. Clavate formations in broth; from chest abscesses and sputum, not pathogenic to rabbits. A. Foulertoni. From whitish dry scaly lesions. A. decussatus. Actinomyces Carougeaui (Gougerot) Brumpt, Precis. Parasitol, ed. 4, 1206, 1929. Discomyces Carougei Gougerot, C. R. Soc. Biol. 67: 580, 1909. Nocardia Carougeaui Castellani & Chalmers, Man. Trop. Med. ed. 2, 817, 1913. Streptothrix Carougeaui Greco, Origine des Tmneurs, 724, 1916. Cohnistreptothrix Carougeaui Chalmers & Christopherson, Ann. Trop. Med. Parasitol. 10: 273, 1916. The lesion caused by this organism was referred to as a paramycetoma by Chalmers & Archibald, N. Orleans Med. Surg. Jour. 70: 466, 1918. The authors have been unable to cultivate the organism. It was named from my- celium seen in the tissues described as "nodosites juxta articulaires." Actinomyces musculorum Hertwig, Arch. Wiss. Prakt. Tierheilk. 12: 365- 372, PI. 6, 1886. Oospora musculorwn suis Lehmann & Neumann, Atlas Grundriss Bakt. 376, 1896. Not cultivated but seen in calcareous deposits in the muscles of swine. Actinomyces Silberschmidti (Chalmers & Christopherson) Dodge, n. comb. Cohnistreptothrix Silberschmidti Chalmers & Christopherson, Ann. Trop. Med. Parasitol. 10: 273, 1916. Streptothrix sp. Silberschmidt, Centralbl. Bakt. 27: 286-493, 1900; Zeitsch. Hyg. 37: 345-380. Pis. 3, 4, 1901. Nocardia SiUjerschmidti Froilano de MeUo & Femandes, Mem. Asiatic Soc. Bengal. 7: 111, 1919. 712 MEDICAL MYCOLOGY Isolated from cases of dacryocystitis. Not greatly pathogenic to rabbit. This organism is a strict aerobe. Colonies on agar deep, round, pinhead size, giving a grayish white uneven surface to the culture. No growth on potato or serum. In broth there is a deposit, yellowish or grayish white, in the bottom. Gelatin not liquefied. Actinomyces cuniculi (Schmorl) Gasperini, Centralbl. Bakt. I, 15: 684, 1894. Streptothrix cuniculi Schmorl, Deutsche Zeitsch. Thiermed. Vergleich. Path. 17: 375-408, Pis. 7, 8, 1891. Streptothrix necrophora Kitt, Bakterienkunde 1900. Cladothrix cuniculi Mace, Traite Bact. ed. 6, 2: 753, 1913. Cohnistreptothrix cuniculi Chalmers & Christopherson, Ann. Trop. Med. Parasitol. 10: 273, 1916. Nocardia cuniculi Froilano de Mello & Fernandes, Mem. Asiatic Soc. Ben- gal. 7: 107, 1919. Oospora cuniculi Sartory, Champ. Paras., 824, 1923. Isolated from ulcers in the muzzles of rabbits. They spread to other or- gans and are, eventually, fatal. Pathogenic to rabbit and mouse ; not to guinea pig, dog, cat, hen, or pigeon. Hyphae 0.15-1.5 fx in diameter and are up to 100/x long. The organism is gram-positive and a strict anaerobe; optimum temperature 37°, no growth below 30° or above 40° C. On serum agar, only deep colonies appear. They are dull white, spherical, very small. No development on potato. On serum, small, grayish white, finely radiate, deep colonies. No growth in upper centimeter. In broth, there is a slight development with uniform turbidity. Neither coagulated serum nor gelatin liquefied. Schmorl's organism is reported by Sanfelice to be a Corynehacterium. Actinomyces Neschczadimenki (Chalmers & Christopherson) Dodge, n. comb. Cohnistreptothrix Neschczadimenl-i Chalmers & Christopherson, Ann. Trop. Med. Parasitol. 10: 223-282, 1916. Streptothrix sp. Neschczadimenko, Centralbl. Bakt. I, 44: 573-578, 1 pi., 1908. Oospora sp. Sartory, Champ. Paras. Homme Anim., 827, 1923. Found in the pus from a lesion near the navel. An account of patho- genicity promised for a later paper. Hyphae 0.75-1.00/* in diameter, some irregular ones attaining 1.5/x. Strict anaerobe, gram-positive, optimum temperature 37° C. On agar, colonies whitish gray, becoming darker, then yellowish, espe- cially in the center. No growth on gelatin or potato. On coagulated serum, L ACTINOMYCETEAE 713 growth is similar to that on agar, with no pigmentation of the medium. In broth, containing egg yolk, there appear whitish grannies on the walls and at the bottom, the medium remaining clear. Granules measure 0.5 x 4.5-9/a, sometimes 1 x 18-20/a in the swollen portions. Actinomyces Thjoettae Dodge, n. sp. Cohnistreptothrix sp. Thj0tta & Gundersen, Jour. Bact. 10: 1-12, 9 figs., 1925. Found in a case of generalized infection with acute rheumatism, pleuritis, pericarditis, and bronchitis. Thought by authors to be a saprophyte close to that of Lohlein. Nonpathogenic to laboratory animals. On veal broth medium, colonies as described below, hyphae nonseptate, showing no clubs and no spores in bottom growth. In surface growth hyphae slender or thick, twisted, breaking up into ovoid spores. Generally gram- positive. On lactose agar, growth rapid, producing smooth, shining, circular, ef- florescent adherent colonies after a few days. Surface growth on agar gram- positive, coccoid forms occasionally gram-negative. Hyphae obserA^ed. When 10 c.c. of the patient's blood is added to veal broth to which has also been added 0.2 gm. secondary sodium phosphate instead of NaCl, with a pH 7.8, and this incubated at 37° C. for two weeks, there is a growth of small, woolly colonies on layer of blood at the bottom; surface colonies dry and scaly with the medium clear. In broth, the optimum pH is 7.4. Coagulated ox serum not liquefied. Gelatin rapidly liquefied. Actinomyces necrophorus (Fliigge) Lehmann & Neumann, Bakt. Diag. ed. 2, 2. Bacillus necrophorus Fliigge, Die Mikroorganismen 265, 1886. Sirepioihrix necropJwra Ernst, Monatsh. Prakt. Tierheilk. 14; 193-228, 1902. Isolated from calf diphtheria. Hyphae 100/x long, gram-positive, anaerobic. In gelatin and agar, fine, white to yellow spherical colonies, center opaque, margin showing filaments, no surface growth. In coagulated horse or ox serum, colonies reddish or brownish red. In milk, odor of stinking cheese. No growth in broth, etc. Pathogenic to experimental animals. Actinomyces Foersteri (Cohn) Gasperini, Centralbl. Bakt. 15: 684, 1894. Streptothrix Foersteri Cohn, Beitr. Biol. Pflanzen 1: 186, 1874, non Gasperini, 1890. Cladothrix Foersteri Winter, in Rabenhorst Kryptogfl. Deutschl. ed. 2, 1 : 60, 1884. Cladothrix dichotoma Mace, Traite Pratique Bact., 1888. Nocardia Forsteri Trevisan, in Sacc. Syll. Fung. 8: 928, 1889. 714 MEDICAL MYCOLOGY Oospora Forsteri Sauvageau & Radais, Ann. Inst. Pasteur 6: 242-273, 1892. Discomyces Foersteri Gedoelst, Champ. Paras. Homme 176, 177, 1902. Isolated from concretions in the lacrimal canal. A facultative anaerobe, not pathogenic for laboratory animals. Hyphae straight, curved, or in irregular helices, 0.5-0.6/x in diameter, arthrospores ovoid, 0.8/x in diameter. On agar, small round colonies, confluent in a mammillate pellicle, folded, grayish, covered with whitish efflorescence. On potato, growth is rapid, chalky; starch is hydrolyzed to sugar. On gelatin, white spheres with short radiating hyphae at the periphery. In broth, there forms a sediment of small grayish spheres at the bottom, mouldy odor. Actinomyces discofoliatus Gmter, Zeitschr. Augenheilk. 79: 477-510, 13 figs., 1933. Isolated from concretions in the tear ducts, not very pathogenic for ex- perimental animals. Mycelium branching, gram-positive in concretions; in cultures spores in chains of 3-10 cells borne on tips of short branches, terminal spores somewhat larger; spores gram-negative at maturity. On agar, colonies grayish white, round, moist, 3-5 mm. in diameter, ad- herent to the substrate, no pigment formed. In anaerobic cultures, growth slower but good, yellowish white colonies, 5-8 mm. in diameter. In agar shake, growth similar, with leaflike outgrowths that become imbricate (like shingles on a roof). On gelatin, growth good along the stab. On Loeffler serum, colony thin, slightly granular. No growth on potato. On broth, abundant gray white sediment leaving liquid clear. Growth at both 22° and 37° C. No hemolysis, no digestion of starch, no coagulation of milk, and no liquefaction of gelatin. Actinomyces Israeli (Kruse) Dodge, n. comb. Streptothrix Israeli Kruse in Fliigge, Mikroorganismen 56, 1896. Discomyces Israeli Gedoelst, Champ. Paras. Homme 163-167, 1902. Discomyces iovis Brumpt, Arch, de Parasitol. 10: 489-572, 1906 non al. Nocardia Israeli Castellani & Chalmers, Man. Trop. Med. ed. 2, 814, 1913. Cohnistreptothrix Israeli Pinoy, Bull. Inst. Pasteur 11: 932, 1913. Brevistreptothrix Israeli Lignieres, Ann. Parasitol. Hum. Comp. 2: 19-22, 1924. Found in a case of human actinomycosis. Wolff-Israel, Arch. Path. Anat. Physiol. [Virchow] 126: 11-59, Pis. 1-8, 1891. Mycelium bacillary, grows best anaerobically. Coccoid forms in old cul- tures. Serum-agar colonies resemble drops of dew, finally becoming yellow- ish. In broth, there appears a deposit in the bottom of the tube. No growth on gelatin. Fungi of Doyen 1891, Jurinka 1896, Silberschmidt 1901, Schukevich 1902, Doepke 1903, and Wright 1904 were referred here by Froilano de Mello. L ACTINOMYCETEAE 715 Actinomyces Thibierg-ei (Ravaut & Pinoy) Greco, Origine des Turaeurs .... 723, 1916. Discomyces Thibiergei Kavant & Pinoy, Ann. Derm. Syphiligr. IV, 10: 417-432, Pis. 2, 3, 1909. Nncardia Thibiergei Castellani & Chalmers, Man. Trop. Med. ed. 2, 817, 1913. Colinistre'ptothrix Thibiergei Chalmers & Christopherson, Ann. Trop. Med. Parasitol. 10: 273, 1916. Oospora Thibiergi Sartory, Champ. Paras. Homme Anim. 792, 1923. Isolated from subcutaneous and intramuscular nodules showing very small white grains in the pus. Medicated with KI. Nonpathogenic to laboratory animals. Hyphae 0.2yu, in diameter, branched, producing bacillary arthrospores, 2 X 0.2/A, curved. Sclerotia 80/a in diameter, claviform formations present. Organism is gram-positive and aero-anaerobe. Aerobic cultures give the more filamentous forms. Optimum temperature 37°-38° C. On ordinary agar, colonies elevated, rugose, rooting, adherent. On mal- tose agar, small white colonies. On potato-gJycerol, small, white, translucent granules form. In broth, bottom growth shows white grains. Growth is best when broth is enriched with egg albumen or serum. Perhaps the following unnamed species of van Loghem should be re- ferred here : Streptothrix sp. van Loghem, Centralbl. Bakt. I, 40: 298-305, 1906. Oospora sp. Sartory, Champ. Paras. Homme Anim. 828, 1923. Isolated from the pus of the abscesses in a fatal generalized infection, also from the sputum. Pathogenic to guinea pig and rabbit. Hyphae up to 40/x long, branched. Spores 4 x 0.5/x. Ends of the hyphae thickened. Organism is both aerobic and anaerobic, optimum temperature being 37° C, no growth at 22° C. ; gram-positive. In agar, colonies white, adherent, round. Anaerobic growth on glucose agar shows slight, amber, irregular, convoluted colonies, 3-4 mm. in diameter. No fermentation. In broth, there is a sediment of white masses up to 5 mm. in diameter, no pellicle, and medium remains clear. No growth on serum, milk, or potato. Actinomyces tenuis (Castellani) Dodge, n. comb. Nocardia tenuis Castellani, Brit. Jour. Derm. Syphilis 23: 341, 1911. Discomyces tenuis Castellani, Proc. Roy. Soc. Med. 6: Derm. 23-27, 1912. Cohniistreptothrix tenuis Ota, Jap. Jour. Derm. Urol. 28: [4], 1928. Isolated from cases of trichomycosis axillaris, flava, rubra, and nigra. This species really causes only trichomycosis flava. The other colors are de- veloped by bacteria living with the fungus. Bacillary bodies 4-10 x 1-1.5|U,, straight or bent, variously branching. Gram-positive, not acid-fast (not cultivated in 1912) 716 MEDICAL MYCOLOGY Macfie, Ann. Trop. Med. Parasitol. 10: 283-289, 1916, claims to have cul- tivated this species on ascitic agar. Colonies translucent, colorless, with a slightly opaque center ; on transplants, hyphae 0.6/* in diameter. Gram-nega- tive, not acid-fast. On old colonies, hyphae gram-positive ; bacillary and cocciform bodies found. On parasitized hairs, gram-negative at first, but gram-positive on old infections. Actinomyces americanus (Chalmers & Christopherson) Dodge, n. comb. 8treptothrix sp. Bloomfield & Bayne-Jones, Johns Hopkins Hosp. Bull. 26: 230-233, 1915. Cohnistreptothrix amcricana Chalmers & Christopherson, Ann. Trop. Med. Parasitol. 10: 273, 1916. Isolated from a liver abscess, also from a chronic, low grade pulmonary infection. No granules in the pus. Organism shows low pathogenicity for guinea pigs, no general infection but local chronic infections which tend to heal spontaneously. Organism grows best under a partial oxygen pressure, 1-2 cm. below the surface. No growth at room temperature, optimum temperature 37° C. No elavate forms. Hyphae 0.5ju. in diameter. Gram-positive, not acid-fast. Small gray colonies appear on ascitic glucose agar, 1 mm. in diameter, opaque with a dark center and irregular edges. No growth on usual media. Actinomyces actinoides (T. Smith) Bergey, Man. Det. Bacteriol. 346, 347, 1923. Bacillus actinoides T. Smith, Jour. Exp. Med. 28: 333-344, Pis. 25-28, 1918. Isolated during a mild epidemic of pneumonia in calves over four weeks old. Not pathogenic for laboratory animals. Mycelium and spherical spores reported ; elavate forms in lungs of calves. Gram-negative. Optimum temperature 37° C. Organism a partial anaerobe. On agar, colonies minute, pale, straw-colored, with flocculent growth in water of condensation. In blood serum, there is flocculent growth in the water of condensation, nitrates not reduced. No growth in gelatin, broth, milk, or potato. Actinomyces cylindraceus (Korte) Brumpt, Precis Parasitol. ed. 4, 1206, 1927. Nocardia cylindracea Korte, Ann. Trop. Med. Parasitol. 11: 205-278, Pis. 8, 9, 1918. Discomyces cylindraceus Neveu-Lemaire, Precis. Parasitol. Hum. 44, 1921. Oospora cylindracea Sartory, Champ. Paras. Homme Anim. 774, 775, 1923. Isolated from an infection of the outer ear, resembling mycetoma and con- taining grains. Aero-anaerobic. On glucose agar shakes, cylindric or hollow spherical colo- nies form in the medium, semitransparent, very adherent hyphal tips sometimes enlarged but no free spores seen. Unable to subculture, acid-fast. On blood smear agar, to which sterile cerebrospinal fluid has been added, small, pale lemon yellow, adherent colonies after smearing, appear like acid- ACTINOMYCETEAE 717 fast bacilli, 0.3-0.5 x 1-4/a; rods allantoic!, mostly curved, stain unevenly. No clavate forms, gram-negative ; in subcultures, gram-positive ; dots in the gram- negative rods. Aerobic culture produces oidial form (bacilliform) in sterile ovarian fluid, development with smell of old boot leather or of a tannery, as in patient's ear. Not acid-fast. Grayish, tough, slimy growth. Finally, after 7 months in broth, oidal converted to arthroidal form; colony spheroidal. Mycelium transferred into glucose agar shake yielded surface growth, gram-negative nonacid-fast, bacilliform hyphae and terminal or intercalary chlamydospores. On glycerol potato, oidial form shows no growth, arthroidal form thin, scanty, yellow, discoloring potato and liquid. On gelatin, no growth. On glucose agar shake, oidial form shows no growth in depth, arthroidal form profuse, dirty yellow, slimy growth. On blood agar, oidial form solid, heaped up, discrete, pale orange, no digestion of hemoglobin. Arthroidal form dirtj yellow, slimy, collecting in the water of condensation as oily balls, hemoglobir digested. In litmus milk, fine fawn-colored flocculus in the oidial form, coloi partly dissipated with slight digestion of curd, arthroidal form showing com plete decolorization and more marked digestion. In inspissated sheep serum the oidial form shows slight digestion, forming a clear fluid with a slighl scum, while the arthroidal form shows more marked digestion, opalesceni fawn-colored fluid with a slight dirty yellow scum. The following organism belongs in this group, but in the absence of cul tural characters, it cannot be placed more definitely. Actinomyces anaerobies Plant. Actinomyces sp. Butterfield, Jour. Infect. Dis. 2: 421-430, 1 pi., 1905. Oospora anaerobies Sartory, Champ. Paras. Homme Anim. 830-832, 1923 Isolated from the lung of an American male, 19 years old, primarily suf fering from diabetes mellitus. Urine showed 3.5% to 8% glucose, also acetone and diacetic acid. Filaments of Actinomyces isolated from pus in wall cavity of right lung, also from bronchopneumonic tissue constituting the walls of the cavity. In the latter, the organism was present in the small bronchi, bron- chioles, and in the air sacs filled with leucocytes. Best demonstrated by stain- ing with Weigert's modification of Gram's method. No other organisms found In tissues, organism usually appears as tangled masses of branching hyphae, sometimes in a distinct radiate arrangement with beaded hyphae in the more central portions. No clavate forms observed in examination of hundreds of such colonies. Organism acid-fast. Actinomyces viridis (Lombardo-Pellegrino) Dodge, n. comb. Streptothrix viridis Lombardo-Pellegrino, Riforma Med. 19: 1065-1068 1903. Isolated from soil, found pathogenic to rabbit, guinea pig, and cat on ex- perimental inoculation. Hyphae long, slender, flexuous, branched, acid-fast, aerobic or anaerobic 718 MEDICAL MYCOLOGY Colonies green, ranging- in shade from that of pistachio to dark green or almost black. Pigment diffusing. Good growth on gelatin and agar, convex, adherent, margin finely lacy ; colony dirty white at first, later becoming green. In broth, no turbidity or pellicle, although floating colonies form and settle. Milk colored green and coagulated. On potato, colonj^ whitish, medium be- coming very dark violet. Actinomyces panginensis (Froilano de Mello & St. Antonio Fernandes) Dodge, n. comb. Nocardia panginensis Froilano de Mello & St. Antonio Fernandes, Mem. Asiatic Soc. Bengal 7: 130, 1919. A saprophyte isolated from a dermatosis. Cells 40-60 X 0.8-1/a. Gram-positive, not acid-fast. On agar, colony white, opalescent, dry granular, margins regularly den- tate, efflorescence white. On sugar agars, better developed, yellowish. On potato, colony cinnabar red in the center with white margin, becoming brown- ish yellow and greenish, medium dirty brown. On broth, pellicle white, ad- herent, folded, turbidity and sediment present ; efflorescence white. Milk coagulated and digested, litmus decolorized. The following organism, incompletely described, perhaps belongs here. Actinomyces sp. Bumier & Duche, Bull. Soc. Fran^. Derm. Syphiligr. 40: 581, 1933. Isolated from a case of superficial eczema, dry, scaly, yellowish red on face, scalp, behind ear, and in inguinoscrotal, abdominal, and genital folds. Strictly aerobic, gelatin liquefied, efflorescence early, white, abundant; odor moldy; slight red pigment on glucose, peptone, or asparagin media. Actinomyces mineaceus (Kruse) Lachner-Sandoval, tJber Strahlenpilze, 66, 1898. Cladothrix sp. Ruiz Casabo, Cron. Med. Quirurg. llabana 20: 340-343, 1894. Streptothrix mineacea Kruse in Fliigge, Mikroorganismen, 63, 1896. Streptothrix rubra Petri, N. Giorn. Bot. Ital. n. s. 10: 591, 1903. Nocardia rubra Chalmers & Christopherson, Ann. Trop. Med. Parasitol. 10: 265, 1916. Actinomyces ruber Sartoiy & Bailly, ]\Iycoses Pulmonaires, 252, 1923. Not Actinomyces ruber Krainsky, 1914, nor Nocardia KrainsMi, Chalmers & Chris- topherson, Ann. Trop. Med. Parasitol. 10: 268, 1916. Isolated from sputum. Nonpathogenic to laboratory animals. Sporulation on the fifth day, spores spherical. On glycerol agar, colony bright red and superficial. On peptone agar, also bright red, granular, dry, rugose, not adherent. Colonies on potato red, spreading rapidly. In broth and milk, red colonies form at the surface and sink to the bottom. Coconut milk becomes turbid, but remains colorless. Gelatin is liquefied, without pigment formation. White pellicle also formed. ACTINOMYCETEAE 719 Actinomyces cerebriformis Namystowski, Bull. Internal. Acad. Sci. Cra- covie, CI. Sci. Math. Nat. 1909: 418-426, PI. 21, 1910. Isolated in a case of infection of the cornea. llyphae up to 1/a in diameter, spores ellipsoid, formed only on old potato and beet cultures. Colonies on ordinary agar, either yellowish, pulvinate, cerebriform, con- fluent or waxy, white, slightly elevated, with concentric growth rings. On glycerol agar, colonies are flat, round, margin wavy, often irregular with definite growth zones, wax white and of a waxy consistency. Colonies on potato, ochre yellow to orange yellow, with a white efflorescence. Colonies often almost spherical, 1 mm. in diameter or 2.5 mm. long and 0.5 cm. high, folded. On beet and carrot, bright dirty yellow, pulvinate, confluent colonies. In broth, colonies small, forming flocculent precipitate, medium remaining clear. In liquid blood serum, a deposit of flakes, 1-2 mm. in diameter. Coagu- lated serum not liquefied. Gelatin is liquefied, the colonies being smaller than in blood serum. No growth on sugar agar, pear, or bread with glucose solution. It seems probable that the following unnamed species is the same as, or closely related to, A. cerebriformis. Streptothrix sp. Bernardinis & Donna, Ann. Ig. Sperim. 15: 489-497, Pis. 8, 9, 1905. Oospora sp. Sartory, Champ. Paras. Homme Anim. 813, 1923. Isolated from a corneal ulcer caused by getting a piece of straw in the eye. Not very pathogenic to rabbit 's eye. Colonies small, not over 1.5 mm. in diameter, slightly elevated, yellow, with a regular margin. Deep colonies brown, granular, regular. On gelatin, growth is similar but slower. On peptone agar, a yellow orange pigment de- velops. Broth, with 1% glucose, becomes turbid, with a mass of flakes set- tling out. Discrete growth on coagulated rabbit serum. Gelatin liquefied, liquefaction beginning in 3-4 days. Actinomyces tossicus (Rossi) Dodge, n. comb. Actinomyces alhus var. tossica Rossi, Ann. Ig. Sperim. 9: 693-708, 1905. Oospora alba var. toxique Sartory, Champ. Paras. Homme Anim. 829, 830, 1923. Isolated from tumors in the abdominal cavities of domestic fowl. Patho- genic to guinea pigs, rabbits, and chickens. On agar, grow^th dirty Avhite, round, adherent, margin areolate, opaque, granulose, yellow brown, covered w^ith a snoAv white efflorescence. On potato, growth same as on agar. On gelatin, colonies round and dirty yellow with margin fringed. Broth remains limpid with round colonies adhering to the glass, and a farinose, fragile pellicle. Starch liquefied but sucrose not in- verted. Milk coagulated. Gelatin liquefied in one week. Actinomyces dermatonomus Bull. Australian Jour. Exp. liiol. 'Sled. Sci. 6: SOl-SU, 2 pis., 1929. 720 MEDICAL MYCOLOGY Isolated from lesions on sheep, producing copious exudates, which hold the overlying wool together in hard lumps, making shearing of these areas impossible and rendering much wool unmarketable ; usually attacks the younger merino animals. Infection varies from 10% to 25% on some farms in Australia. It is uncertain whether the similar condition, reported by Becker, is due to the same organism. In the early stages, a serofibrinous fluid, with a moderate number of leucocytes, exudes, is absorbed by the overlying wool, and then dries. In the later stages, this exudate is mixed with a relatively large num- ber of squamous epithelial cells from the surface of the skin. Chronic lesions become less exudative and more keratogenous or desquamative. In the early stages, the follicle is not involved (although it may be involved later), allow- ing the wool to pull away from the skin. The horny layer is diffusely invaded by neutrophile polymorphonuclear leucocytes. There is vesiculation of the epithelium in the more superficial parts of the rete mucosum, the vesicles being early invaded by polymorphs. In the cutis vera, surrounding the fol- licles, there is a well-developed accumulation of leucocytes and plasma cells, with an occasional polymorph. Sometimes there are areas of edema between the follicles, with some hemorrhagic extravasation. The crusts consist of keratinized epithelial cells and polymorphs bound together in a serofibrinous exudate, along with branching mycelium. Organism pathogenic to rabbits where skin has been injured by pulling out hair but not on uninjured skin. Local abscesses caused on subcutaneous inoculation, lesions in rabbit and guinea pigs being very close to those in sheep. Similarly pathogenic to cow and horse, in the latter very desquamative. The hyphae are short, easily broken into arthrospores. Mycelium is also present in the horny layer, and more sparsely in and around the edematous portions of the corium. Branching lateral, the branches usually of smaller diameter than the main trunk. Mycelium 0.5-1/* in diameter, conidia 0.5-1. 75/a, arthrospores 1.8-4.5/x long, usually 3.6/a, often thickened at one end with a wavy contour. Gram-positive, not acid-fast. Aerobic, growth in agar shake at all depths, but the organism is not a strict anaerobe. Optimum tempera- ture 37°, growth very poor at 22° C. On peptone agar, growth in 24 hours at 37° even, raised, convex to pul- vinate, yellowish, opaque, somewhat glistening, 1 mm. in diameter. After 48 hours colony more irregular, less raised, more tenacious, not spreading. No aerial hyphae. Color better developed at 22°, antimony yellow (Ridgway 16 'b), medium not colored, colonies flatter, center umbonate with raised lobate or lobulate margins. Deep blood agar plates show clear zone of hemol- ysis, 3.5/x in diameter, around colonies. No digestion in Loeffler's blood serum, good growth, more tenacious. Gelatin liquefied only at pH 7.6, growth sac- cate, 1 cm. in 14 days. At 37° C. liquefaction is complete in 9 days. No visible growth on potato. In peptone broth, pellicle appears in 24 hours, no clouding, growth on sides of tube with slight sediment, which becomes more abundant as pellicle falls, becoming mucoid. Litmus milk bleached in 4 days, medium digested in 7-8 days, medium yellow. Bromcresol purple milk digested in ACTINOMYCETEAE 721 7-8 days with increasingly alkaline reaction, ammonia produced. Glucose and fructose acidified but not fermented in 48 hours, glycerol more slowly. Little acidity with galactose and usual disaceharides, polysaccharides, pentoses, higher alcohols, and glucosides. Urea developed an alkaline reaction. No nitrate reduction. Protease active only in alkaline medium producing am- monia, no action on coagulated blood serum, egg white or fibrin. Actinomyces mexicanus Boyd & Crutchfield, Amer. Jour. Trop. Med. 1: 268-282, Figs. 15-18, 1921. Nocardia mexicana Ota, Jap. Jour. Derm. Urol. 28: [4], 1928. Isolated from mycetoma of the foot. Nonpathogenic to guinea pig. Hyphae 0.7/x, in diameter, tortuous, branching, tips sometimes slightly swollen. Chlamydospores formed in blood agar and Loefller's blood serum. Organism aerobic, optimum temperature being 37° C. Gram-positive, not acid-fast. On glycerol agar, growth very slow, colonies small, 1 mm. in diameter, more or less convex, tending to be wrinkled. On blood agar, colonies discrete, irregular, rectangular, 3-4 mm. on a diagonal, margins abruptly elevated 1-2 mm., center umbilicate, efflorescent, with a narrow zone of hemolysis. On Conn's synthetic citrate agar (citric acid 5 gm., NH^Cl 0.5 gm., KgHPO^ 0.5 gm., glycerol 6 c.c, agar 15 gm., water 1,000 c.c), colonies small, white, opaque convex, moist. On Waksman's modification of Czapek's agar, no growth visible. On meat infusion agar, colonies discrete, 2 cm. in diameter, with abrupt edges, 1-2 mm. high, center pointed or umbilicate, smooth, moist, slightly yellow. On Krainsky medium (glucose 10 gm., NH^Cl 0.5 gm., as- paragin 1.5 gm., agar 15 gm., water 1,000 c.c), colonies discrete, circular, not over 2 mm. in diameter, slightly convex, moist and white, mycelium penetrat- ing 2-3 mm. Growth on potato very slow, white and flat, with substrate un- altered. Growth also slow on gelatin. ]Meat infusion broth shows a flaky sediment, with liquid remaining clear. In carbohydrate peptone water, no fermentation, with colonies appearing as small white balls, not over 1 mm. in diameter, in the bottom of the tube. Medium remains clear. In litmus milk, growth is slow, no coagulation, digested in 3 weeks. Growth also very slow in Loeffler's blood serum as discrete, rectangular colonies 2-3 mm. in diam- eter, conical, center elevated 1-2 mm., colorless, moist, smooth, no liquefaction in 3 weeks. Gelatin liquefied. Faint fecal odor in cultures. Actinomyces Rivierei (Verdun) Brumpt, Precis Parasitol. ed. 4, 1201, 1927. Streptothrix sp. Riviere, Cong. Franc. Med. Bordeaux 1895, 2: 1003-1011, 1896. Actinomyces Sabrazes et Rivieri Berestnev, Diss. iMoskva 1897. Nocardia Rivierei Verdun, Precis Parasitol. 1912. Discomyces Rivierei Neveu-Lemaire. Precis Parasitol. Hum. 43, 1921, non Nocardia indica Kanthack fide Froilano de Mello & Pais, 1918. Oospora Rivierei Sartory, Champ. Paras. Homme Anim. 792, 1923. 722 MEDICAL MYCOLOGY Isolated from sputum and pus of a bronchopleuropuluionaiy infection fol- lowed by multiple miliary subcutaneous abscesses. Early symptoms those of tuberculosis, but Mycobacterium tuberculosis absent. Found pathogenic to lab- oratory animals by the use of a specialized technic. Hyphae Iju, in diameter, arthrospores 1.5/i, long by 1/x in diameter. Or- ganism a strict aerobe. Colonies on peptone agar small, conical, 1 mm. in diameter, powdery, growing to about 5 mm. in diameter, reverse yellow, medium blackened. On potato, growth slow, colonies appearing like little blocks of powder. Growth also slow on eg^ white, the substrate not digested. In liquid media, small grains size of a millet seed settling to the bottom, colonies floating, if not dis- turbed. Growth good on milk, forming a flesh-colored, whitish powdery pel- licle with furrows. Fat assimilated, casein and lactose not perceptibly at- tacked, apparently only glycerol used from fat. Grown on 6% glycerol pep- tone agar, colony mammillate. white, powdery, furrowed, cork color, finally becoming black. Lactose, maltose, glucose, and fi-uctose utilized. Gelatin liquefied. Actinomyces Gibsoni Dodge, n. sp. Streptothrix sjj. Gibson, Jour. Path. Baet. 23: 357, 358, 1920. Oospora sp. Sartory, Champ. Paras. Homme Anim. 776, 1923. Isolated from enlarged spleen. Nonpathogenic to guinea pigs, rabbits, and mice, pathogenic to Macacus monkej's, organism being reisolated in one case from an enlarged spleen. Gram-positive ; aerobic. Colonies on agar slants discrete, buff or white, hemispheric, becoming de- pressed and umbilicate. White efflorescence and musty odor. In peptone broth, there is a bottom growth of small, white "snowballs," with some even- tually rising and floating on the clear medium. Neutral red lactose broth is unchanged. Milk coagulated in 4-5 days. Gelatin slowly liquefied with growth near surface. Actinomyces Guegueni (Ota) Brunipt, Precis Parasitol. ed. 3, 1921; ed. 4, 1191, 1927. Oospora lingualis Gueguen, C. K. Soc. Biol. 64: 852-854, 1908; Arch, de Parasitol. 12: 337-360, 31 figs., 1909. Discomyces lingualis Brumpt, Precis Parasitol. ed. 1, 1910. Nocardia li^igualis Castellani & Chalmers, Man. Trop. Med. ed. 2, 819, 1913. Nocardia Guegueni Ota, Jap. Jour. Derm. Urol. 28: [4], 1928, excl. syn. Black pilose tongue by Gueguen along with Cryptococcus linguae-pilosae which later he thought to be the agent of the disease. Pathogenic for guinea pig. Mycelium slender, spirals present, sometimes terminating in a chlamydo- spore. Chlamydospores commonly intercalary, ovoid, 1-3/x in diameter, conidia in chains, spherical, O.Sfi, on simple clavate conidiophores, 4-5/x. Optimum tem- perature 37° C, partial anaerobe. ACTINOMYCETEAE 723 In broth, white flocci at the bottom of the tube. On carrot, small white points 1-1. .5 mm., later colored cafe-au-lait, with round protuberances, becoming powdery. On gelatin, colony white. Milk coagulated on third day, casein di- gested. Coagulated albumin digested. Nitrates reduced to nitrites, no gas. Actinomyces Pinoyi (Froilano de Mello & St. Antonio Fernandes) Dodge, n. comb. Nocardia Pinoyi Froilano de Mello & St. Antonio Fernandes, Mem. Asiatic Soc. Bengal 7: 130, 1919. Isolated from a case of erythrasma. Cells l-2yu, in diameter, gram-positive, not acid-fast. On agar, colonies moist, shining, whitish, margins irregularly dentate. On potato, colony dry, chestnut color, with small elevations, efflorescence chocolate color, medium colored deep brown. In broth, turbidity and sedi- ment ; slight pellicle on vegetable decoction. Milk coagulated in 72 hours, digested in 96 hours with odor of new leather, litmus decolorized. Actinomyces goensis (Froilano de Mello & St. Antonio Fernandes) Dodge, n. comb. Nocardia goensis Froilano de Mello & St. Antonio Fernandes. Mem. Asiatic Soc. Bengal 7: 130, 1919. A saprophyte, isolated from lesions of vitiligo. Cells 5-40 X 0.8-1/Li. Gram-positive, not acid-fast. No growth on agar. On glycerol agar, colony white, dry, with slight efflorescence, resembling a bacterial colony. On glucose and maltose agars, colonies granular, opaque, milky white, thick. On potato, colony white, dry, viscous, cerebriform, becoming clear gray, efflorescence white ; medium pig- mented yellow, becoming brownish. On broth, pellicle thin, white ; liquid turbid, sediment white. On sugar broths, pellicle better developed. Milk coagulated and digested, acidified. Actinomyces Christophersoni (Froilano de Mello k St. Antonio Fernandes) Dodge, n. comb. Nocardia Christophersoni Froilano de Mello & St. Antonio Fernandes, Mem. Asiatic Soc. Bengal 7: 130, 1919. Isolated from the air. Cells 30-70 X 0.6-0.7/*, gram-positive, not acid-fast. On agar, colony moist, creamy, without efflorescence ; on sugar agars, similar or slightly yellowish and waxy. On potato, colony elevated, cream yellow, becoming dry and whitish, waxy, no pigmentation of medium. On carrot, colonies similar, moist, granular, resembling butter. On broth, a strong white pellicle, not adherent, white sediment, slight turbidity. On glucose broth, similar but yellowish on vegetable decoctions, pellicle grayish, sedi- ment sandy. Milk coagulated and slightly digested, acidified. Actinomyces pyogenes (Chalmers & Christ opherson) Dodge, n. comb. Streptothrix pyogenes Chalmers & Christopherson, Ann. Trop. Med. Para- sitol. 10: 270, 1916. 724 MEDICAL MYCOLOGY Streptothrix sp. Caminiti, Ceutraibl. Bakt. I, 44: 193-208, 4 ph., 1907. Isolated from the air. Found pathogenic to rabbit, guinea pig, and dog. Generally hyphae are present in young colonies, in old colonies bacilli- form spores. Growth very slow under anaerobic conditions, diastase present. Colonies white, opaque, wavy, margin elevated, center umbilicate, ef- florescent. On 2% glycerol agar, colonies dry, gray white, elevated, adherent. Growth rapid. On potato, colonies small, umbilicate, soon confluent, white or yellowish. On blood serum, colonies white, adherent. On blood serum with glycerol, colonies opaque, gray green, with the medium tinted slightly violet. Colonies in broth, floating both at top and at bottom, the former forming a pellicle at first white, then darker, the medium remaining clear. In serum broth, colonies are gray white. Colonies brown ochraceous or dark brown in glycerol broth. The higher the percentage glycerol, the darker the colony color on all media. On milk, a greenish pellicle, opaque and folded, forms. It becomes dark green in 2 weeks. A flocculent coagulum settles. Gelatin liquefied. This organism is near to Actinomyces alhus, to which Chalmers & Christo- pherson reduce it. Actinomyces gypsoides Henrici & Gardner, Jour. Infect. Dis. 28: 232-248, 1921. Oospora gypsoides Sartory, Champ. Paras. Homme Anim., 802, 1923. Isolated from the sputum of a patient suffering from cough with pains in the chest. Pathogenic to rabbits and guinea pigs. Colonies star-shaped. On agar, they are thin, grayish, soon becoming thick, opaque, and chalky white. Surface dry and wrinkled, adhering to the medium, breaking away only in large flakes. Mycelium is buff-colored below the efflorescence. GroAvth on sugar agars more rapid, but otherwise similar. Similar also on potato. Stratiform superficial growth on gelatin. In liquid media, small snow white flakes appear, presently coalescing into a thick, wrinkled, snow white pellicle and broad ring. After 2 weeks the pellicle breaks and settles. No further growth occurs. In litmus milk, there is a similar pellicle but with a yellowish tinge. At first alkaline and curdled, then litmus reduced and casein digested in 2 weeks. Meat extract media, also Dunham's peptone, darken, becoming brownish. Organism grows also on Czapek agar and on soil infusion agar. No hemolysis of blood agar. Gelatin liquefied. Neither serum nor Dorsett's egg medium digested. Actinomyces invulnerabilis (Acosta & Grande Rossi) Dodge, n. comb. Cladothrix invulnerabilis Acosta & Grande Rossi, Cronica Med. Quirurg. Habana 19: 97-100, 1893. Isolated as laboratory contamination, and also from water ; not pathogenic for laboratory animals. On agar, colonies small round dirty white, becoming dull white, margins elevated, adherent to substrate ; surface with radial furrows ; reverse yellow. On glycerol agar and potato, growth similar. i ACTINOMYCETEAE 725 On gelatin, colony silvery white, center deprfssed, velvety, liquefying the gelatin slowly. On potato, colonies white, chalky, earthy odor, potato black- ening. On milk, coconut milk, and broth, yellow solid pellicle (tube can be inverted without flow) milk digested (?). Yery resistant to heat, resisting 6 discontinuous sterilizations at 100° C, grows in T"r boric acid. Actinomyces roseus Xamyslowski, Centralbl. I>akl. 1, 62: 564-;')68, 1912. Ac fi 7107711) ces sp. Lciwenstein, Klin. Monatsbl. Augenheilk. N. F. 10: 203- 207, 4 figs., 1910. Not Nocat'dia 7-osea Chalmers & ('hristopherson, Ann. Trop. Med. Parasitol. 10: 270, 191 (i. which is .1. roseus Krainsky, Centralbl. Bakt. IT. 41: 649, 1914. Ilyphae 6-8;a, richly septate, branched. Colonies on agar and potato, chalky white; on gl\'cerol agar, delicate rose. Colonies cerebriform and chalky white on gelatin. In broth no turbidity. Growth on lactose litmus the same as on sugar. All colonies have musty odor. No growth on milk or plum decoction, slight growth on egg white Coagulated ox blood serum liquefied after a few days. Gelatin not liquefied. Actinomyces candidus Petruschky in Kolle & Wassermann. Handb. Path. ]\Iikroorganismen 2: 832. Streytothrix geda7hcnsis IL Petruschky 1898, Froilano de Mello & Pais, Arq. Hig. Pat. Exot. 6: 158, 1918. StreptotJwix lathridii Petruschky, 1898. Nocardia Candida Castellani & Chalmers, Man. Trop. Med. ed. 2, 818, 1913. Discor7iijces candidus Verdun & Mandoul, Precis Parasitol. 754, 1924. Found in sputum. Aero-anaerobe. Surface of colony on potato vtlvety with chalky efflores- cence. Serum and gelatin liquefied. Actinomyces albus (Rossi-Doria) Gasperini in Lehmann & Neumann, Atlas Grundr. Bakt. 2: 383, 1896 excl. syn. ActinoTiiyces chTomogenes p. Gasperini, 18. St7-eptotlirix Foersteri Gasperini, Ann. Mierogr. 2: 449-474, Ph. 5-7, 1890, non aliorum. Streptothrix alba Rossi-Doria, Ann. 1st Ig. Sperim. Univ. Roma N. S. 1: 399-438, PI. 8, 1891. Oospoi'a Boriae Sauvageau & Radais, Ann. Inst. Pasteur 6: 242-273, 1892. Actinomyces hovis albus Gasperini, Centralbl. Bakt. I, 15: 684, 1894. Gladotlirix alba Mace, Traite Pratique Bact. ed. 3, 1897. Nocai'dia alba Froilano de Mello & St. Antonio Fernandes, Mem. Asiatic Soc. Bengal 7: 105, 1919. Reported by Massaglia from a case of tuberculosis, 1904; probably also case I of Chatschaturjan et al. (1933). Animal inoculations gave lesions of pseudotuberculosis. Not acid-fast. Aerobe, growing only slightly under anaerobic conditions. 726 MEDICAL MYCOLOGY On maltose peptone agar, colony verrucose, wrinkled, waxy, grayish white, light amber yellow margin with a chalky efflorescence, but no pigmentation of the medium. On potato, colony wrinkled, grayish to yellow or brown with a chalky efflorescence, diastatic action and brown pigmentation of the medium. On gelatin, a yellowish white flocculent mass on surface with center brown. Fir tree arborescence in the medium. In peptone beef broth, there is a sedi- ment of discrete opaque, white, flocculent colonies. No pigmentation of the medium. Litmus milk at first becomes pink, then a clear red brown, and alkaline. Horse serum is completely liquefied with the formation of a brown pigment. Gelatin is also liquefied. Actinomyces Garteni Brumpt, Precis Parasitol. ed. 4, 1191, 1927. Cladothrix liquefaciens No. 2. Garten, Deutsch, Zeitschr. Chirurg. 41: 257- 285, PI. 6, 1895. Discomyces Garteni Brumpt, Precis Parasitol. ed. 1, 1910. Oospora Oarteni Sartory, Champ. Parasit. Homme Anim. 778, 779, 1923. Nocardia Garteni Gougerot, Gaz. des Hop. 86: 199, 1913. Isolated from abscesses. Pathogenic to rabbits, guinea pigs, and pigeons. Colonies on agar much wrinkled, gray white, shining, with a chalky ef- florescence; later penetrating and darkening the medium. On glycerol agar, growth is slower, moist, and shining, with no efflorescence. Organism grows on potato, better at 37° C. than at 22° C, with a white efflorescence formed and potato discolored on the surface. Small grayish white adherent colonies on gelatin. In broth or blood serum, there is a grayish white deposit of flocculi, surface colonies showing the efflorescence. Solid blood serum is liquefied in 6 days, the liquid remaining clear. Gelatin also liquefied. Actinomyces Nicollei (Delanoe) Nannizzi, Tratt. Micopat. Umana [Pol- laeci] 4: 36, 1934. Nocardia Nicollei Delanoe, Arch. Inst. Pasteur Tunis 17: 257-274, 3 figs., 1928. Isolated from a voluminous mycetoma of the thigh in the tribe Oulad Bou Zerrara, Morocco. It began with inguinal adenitis and developed slowly, with no pain. Eight leg larger than left, lesions on the upper two-thirds of the thigh violet in color, producing pus with small yellow grains. Medication with KI helped somewhat, but patient not hospitalized long enough for a cure to be effected. Patient died six months later. Reported as not pathogenic to animals, but pathogenicity tested only after organism had been cultivated four years on potato. Grains yellowish white, 0.5-0.8, rarely 1 mm. in diam- eter, easily crushed, becoming ochraceous on drying. Hyphae branched, not over 1/x in diameter, nonseptate, no nuclei observed. Terminal, sporiferous portion of hypha is abjointed from the rest. Spores spherical, 0.3-0.7/a (also reported I/a) in diameter, not easily staining. Chlamy- dospores deeply staining. Gram-positive, not acid-fast. Optimum tempera- ture 25°-30° C, growth between 15° and 37° C. ACTINOMYCETEAE 727 Colonies, on ISabouraud glucose agar, small, round, elevated, uneven, mam- millate, snoAv white, dry, chalky, hard, crushing with difficulty. Subcultures gradually die out on this medium. On Langeron 8% sugar agar, very old colonies show a slight rose color. On simple agar, colony whitish, granular, humid, adherent, older colonies becoming folded and eerebriform, covering the surface of the agar and being somewhat sunken in the agar at the mar- gins. An efflorescence appears on the third day, with greater elevation. On potato, with or without glycerol, colonies become confluent, forming whitish cordons, sometimes sinuous, the chalky surface mammillate and uneven covering the whole surface after several subcultures, turning yellowish in age with the substrate becoming greenish. On carrot, colonies creamy, slightly yellowish, later becoming chalky and wiiite with carrot browning. In potato juice* a fine white powder appears at the surface with white floating crateri- form colonies. These are dry, 0.2-0.25, up to 0.5 mm. in diameter. Later, center of crater is elevated, yellowish, waxy, finally confluent, forming a waxy, thick, yellowish folded pellicle with a fine white efflorescence. In ear- rot juice, prepared as the first method for potato juice,* there is growth both in depths and at the surface. Medium remains clear, colonies grayish Avhite, not dry. Ring prominent, 6-7 mm. above the liquid. Pellicle is thick, folded, and gray with a white efflorescence, lower surface being black and liquid amber. In broth, growth is somewhat as in carrot juice, but no pellicle forms, and ring is not very adherent. Starch medium is liquefied. No fermentation of glucose, fructose, maltose, sucrose, or lactose. Serum is completely lique- fied. Gelatin also liquefied with the formation of a golden yellow pellicle. Odor of cultures moldy. Actinomyces Dassonvillei (Liegard & Landrieu) Brumpt, Precis Parasitol. ed. 4, 1191, 1927. Nocardia Dassonvillei Liegard & Landrieu, Ann. d 'Occulistique 46: 418- 426, 1911. Discomyces Dassonvillei Neveu-Lemaire, Precis Parasitol. Hum. 42, 1921. ISfreptothrix Foersteri Gasperini, Ann. Microgr. 2: 449-474, Pis. 5-7, 1890, non aliorum. Originally isolated from a case of conjunctivitis. Potron (1913) states that this species is pathogenic for man and rabbits. Since I have not been able to locate the original description, the following is taken from the paper by Potron & Thiry (1913). Hyphae 0.5-0.7/a in diameter, branched, septa not seen. Arthrospores ovoid, in chains, not staining, highly refringent. On solid media, colonies acuminate, white, dull, chalky, very difficult to separate from contaminating bacteria. On potato, colonies white, soon chalky, covering the whole surface of the medium, which is colored violaceous at first. Colonies become deep green {vert foncc), with guttation of greenish drops *Piit 250 pm. grratc-d potato in 300 c.c. water, triturate, decant, and filter. Sterilize at H5°-120° C. Alternate method; cover pulp with water, autoclave 15 minutes at 115*-iaO* C. decant, filter, and .^iterilize as above. 728 MEDICAL MYCOLOGY in old cultures. When the drops are removed, the colony is grayish green with a moldy odor. On carrot, growth slow, chalky, pure white and very thin. No growth on turnip. On broth, also asparagus and maltose broth, grayish fiocci at bottom of tube, liquid clear, some flocci adhering to the walls, or if they reach the surface they become chalky and float ; color of broth deepens; odor is slightly moldy. On coagulated egg albumen, colony chalky, medium becomes translucent, then brown, and is partially liquefied. On serum, growth slow with partial liquefaction of medium. Milk slowly coagulated then the eoagulum is digested, leaving a clear brown liquid in which the colonies sink. Most recent authors have referred Streptothrix Foersteri Gasperini to this species, but since it differs in some respects, I shall give the description below : Streptothrix Foersteri Gasperini, Ann. Micrographie 2: 449-474, Pis. 5-7, 1890 non aliorum. The cultures appeared as contaminants from the air. Mycelium l/x in diameter, spores 1-1. 5/a in slightly curved terminal chains. Colony on gelatin at room temperature, round, elevated, moist, slightly yellowish at first, becoming more elevated, white, powdery, medium liquefied late and very slowly. Agar colonies round, hemispheric with zones of sporu- lation, otherwise similar to gelatin colonies. Serum slowly liquefied. Little growth on cooked potato. On broth, a thick pellicle is produced. Probably the organism isolated from vaccine from heifers by Sabrazes & Joly, C. R. Soc. Biol. 50: 134, 135, 1898, also belongs here. Actinomyces minimus (LeCalve & Malherbe) Dodge, n. comb. Oospora forme de Microsporum [Audouini var. equinum] Bodin, Arch, de Parasitol. 2: 606-609, 1899. Trichophyton minimum LeCalve & Malherbe, Arch, de Parasitol. 2: 218- 250, 489-503, 1899; Ibid. 3: 108-110, 1900. Microsporum minimum Castellani & Chalmers, Man. Trop. Med. ed. 8, 993, 1919. Isolated from ringworm of horse and dog. Sabouraud suggests that this was a saprophyte, as Bodin was unable to repeat the results of LeCalve and Malherbe. On Sabouraud agar, surface becomes covered with yellowish gray eleva- tions, 6 X 4 X 1.5-2 mm. high. Colonies on beef agar circular, little ele- vated, with sinuous rays. Center elevated, grayish, powdery. On oatmeal agar, growth shows colony with irregular margins, eroded, granular, yellow- ish. Colonies on potato are eerebriform, sinking into the medium, white. On serum gelatin, there appears a grayish white efflorescence. Growth on horse- hair is in the form of irregular plaques, 10-15 mm. in diameter, yellowish gray, slightly elevated, chalky. On straw, colony is irregular and grayish. In turnip broth, there are floating powdery islets and spongy, mucoid sediment. In oatmeal broth, growth is more rapid, with a sort of grayish pellicle form- ing. In carrot broth, growth still better; in potato broth, still better. Pellicle ACTINOMYCETEAE 729 is grayish, umbilicate with the central elevation surrounded by a moat and a flat marginal zone. Reverse brownish yellow. In Raulin's solution con- taining 2% peptone, mucoid masses, suspended or settling, are slowly formed. Starch is hydrolyzed. Milk is peptonized without coagulation. Gelatin is liquefied. Actinomyces radiatus Namyslowski Bull. Internat. Acad. Sci. Cracovie CI. Sci. Math. Nat. 1909: 418-426, PJ. 21, 1910. Found in infection of the cornea. Hyphae about 1/x thick, branched, producing spherical to ellipsoid spores. On ordinary agar, colony gray, with an elevated center and round margin, central elevation more or less cruciform or star-shaped. On glycerol agar, growth is flat, round, crateriform, with concentric growth zones, white. On coagulated blood serum, colonies round, confluent with white efflorescence. Serum liquefied. On egg white, colonies white. Gelatin is liquefied, with a sediment of gray white colonies and a white pellicle. In broth, a gray white sediment, medium clear. Milk digested, finally becoming brown with thin white pellicle. No growth on sugar agar, pear, potato, bread, or carrot. Actinomyces rubidaureus Lachner-Sandoval, tJber Strahlenpilze 66, 1898. Cladothrix Thiry, Arch. Physiol. Norm. Path. 8: 283-288, 1897. Actinomyces mordore Thiry, These, Nancy 82-93, 1900. Nocardia modore Chalmers & Christopherson, Ann. Trop. Med. Parasitol. 10: 265, 1916. Nocardid Thiryei Froilano de Mello & Pais, Arq. Ilig. Pat. Exot. 6: 193, 1918. Actinomyces Thiryi Sartory & Bailly, Mycoses Pulmonaires 252, 1923. Oospora mordore Sartory, Champ. Paras. Homme Anim. 824, 1923. Isolated from a case showing anginous exudate with edema. Hyphae branched, spores on erect branches, hyaline. This organism is a strict aerobe ; gram-positive. On Sabouraud conservation agar, colonies circular, elevated, isolated, grayish or violaceous, with a white efflorescence and surrounded by reddish brown aureole, shining. This metallic lustre is produced by lamellar crystals with a clear amethyst violet and ruby red center. The medium browns some- times. Oil glycerol peptone agar, colony yellow, ochraceous or brownish, violet or white, without the metallic lustre. On potato, colonies are rose, grayish verrucose ; the medium browns and has an eroded (literally "worm- eaten") appearance. On gelatin, there forms a pellicle, yellow gray at the surface, Avith efflorescence and crystals, less colored than on agar. On peptone and peptone brotli, a ring or pellicle with floccose sediment. Serum and egg albumen are liquefied with a strong odor of trimethylamine. Gelatin is. rapidly liquefied. Milk is coagulated and rapidly peptonized. Actinomyces Avadi Dodge, n. sp. Streptofhrix Madurae Koch & Stutzger, Zeitschr. Hyg. 69: 17-24, 1911. nor. Vincent, 1894. 730 MEDICAL MYCOLOGY Isolated from a Madura foot in Egypt, case of Dr. Avad. Nonpathogenic to experimental mammals, pathogenic to frogs. Mycelium 0.3-0.5//, in diameter, branched at right angles. In young colo- nies hj'phae break up into ovoid or spherical spores. Gram-positive. Growtli aerobic, optimum temperature being 22° C, growth poor at 37° C. Organism grows on coagulated horse serum, glucose agar, simple agar, abundance of growth decreasing in that order. Colonies white, later brown- ish, adherent, confluent with a wavy, shining surface resulting, which is dull and brown in old cultures. On ascitic agar, colonies are thin and growth is slow. Growth also slow on potato. In peptone solution, colonies 3-4 mm. in diameter, white, filling bottom of the tube and adhering to the Avails. In broth, to which three parts horse serum has been added, there is a delicate white pellicle which sinks easily, a slight ring, and sediment similar to that in peptone solution. No growth at 37° C. In simple broth, no pellicle, growth as in peptone solution but less abundant. The addition of glucose does not alter this. Glucose and lactose unchanged. In milk, there is bottom growth, no color change in litmus. ]\Iilk coagulated in 2-3 days, curd digested in 4-6 weeks. Gelatin slowly liquefied, colonies gradually sink to the still solid substrate and eventually to the bottom of the tube. Coagulated serum digested Actinomyces Levyi Dodge, n. sp. Acthiomyces sp. Levy. Oospora sp. Sartory, Champ. Paras. 827, 1923, with incorrect reference. Sartory states : Isolated from pus. Not pathogenic for guinea pigs or white rats. Shining orange colonies on all media. On agar, colonies thick, adherent, without chalky efflorescence. Gelatin and serum liquefied. On broth, isolated colonies along the Avails of the tube. No groAvth on potato. On litmus milk, an orange color at the surface, no change of acidity, gradu- ally digested. Starch completely hydrolyzed. Actinomyces bovis Harz, Centralbl. Med. Wiss. 15: 485, 1877; Jahresber. Miinchen. Zentral Thierarzeneischule, 781, 1877. Discomyces hovis Rivolta, Clin. Veterinaria 1: 208, 1878. Bacierinm aciinocladothrix AfanassicA', St. Petersb. Med. Woch. 13: 84, 1888. Nocardia actmomyces Trevisan, I Genere e le specie delle Batteriacee 9, 1889. Cladothrix hovis Mace, Traite Practique Bact. ed. 2, 666, 1891. Streptothrix aciinomyces Rossi-Doria, Ann. 1st. Ig. Sperim. UniA\ Roma 1: 405, 1892. Oospora hovis Sauvageau & Radais, Ann. Inst. Pasteur 6: 271, 1892. Actinomyces hovis sulpliureus Gasperini, Centralbl. Bakt. 15: 684, 1894. Nocardia hovis Blanchard in Bouchard, Traite Path. Gen. 2: 857-868, 1895. Cladothrix aciinomyces Mace, Traite Practique Bact. ed. 3, 1038, 1897. Streptothrix actinom,ycotica Foulerton, Lancet 2: 780, 1899. Streptothrix hovis communis Foulerton. Jour. Comp. Path. Therap. 14: 50, 1901. ACTINOMYCETEAE 731 Streptothrix hovis Chester, Man. Determinative Baet. 361, 1901. Sphaerotilus hovis Engler, Syllabus Pflanzenfam. ed. 5, 5, 1907. This organism is the common cause of actinomycosis or lumpy jaw in cattle. Many strains, which probably belong elsewhere, have been referred here. In view of this confusion, Puntoni (1931) would discard the name alto- gether, in favor of the later more carefully described species of Gasperini and others. Apparently current usage tends strongly toward the identifica- tion of A. hovis with A. sulfureus of Gasperini. The following species descrip- tion is taken from Froilano de Mello & Pais (1918), from Bergey (1923), and from Puntoni 's description of A. sulfureus. Slender branching hyphae, 0.4-0.6/^ in diameter. Large club forms are seen in animal tissues ; gram-positive. Colonies on glycerol agar, waxy, smooth, yellowish, umbilicate cartilagi- nous, adherent to the medium. In age, they tend to become brown and color the medium. They are covered with a sulphur-colored efflorescence. On synthetic agar, growth restricted, yellowish aerial mycelium appears late, becoming light sulphur yellow, powdery. On starch agar, dirty yellow. On plain agar, colony abundant, cream colored, becoming fawn colored, brown or almost black. On gelatin stab, inverted fir tree, colonies on surface opaque, white, punctif orm. On coagulated serum, colonies similar but larger, adherent, no pigmentation. On potato, growth abundant, wrinkled, gray to canary yel- low; starch hydrolyzed. On carrot, whitish gelatinous colony not adherent. On glycerol broth and other liquid media, no pellicle, sediment of punctiform colonies. (Bergey reports thin yellowish pellicle.) Milk slowly coagulated and digested, sugars not fermented. Gelatin and serum digested. Acid in glucose, lactose, sucrose, maltose, and glycerol. Optimum temperature 37° C. Actinomyces Lanfranchii Sani, 1916. Nocardia Lanfranchii Froilano de Mello & Pais, Arq. Hig. Pat. Exot. 6: 178, 1918. Isolated by Finzi from glandular and ganglionar actinomycoses of ox. Gram-negative. Otherwise very close to A. hovis [original description not seen]. Actinomyces liquefaciens (Hesse) Brumpt, Precis Parasitol. ed. 4, 1192, 1927. Cladothrix liquefaciens Hesse, Deutsche Zeitschr. Chirurg. 34: 275-307, Pis. 11, 12, 1892. Nocardia liquefaciens Castellani & Chalmers, Man. Trop. Med. ed. 2, 818, 1913. Discomyces liquefaciens Neveu-Leniaire, Precis Parasitol. Hum. 42, 1921. Oospora liquefaciens Sartory, Champ. Paras. Homme Anim. 778, 1923. Streptothrix huccalis Goadby, 1903, non Roger, Bory & Sarto^^^ Isolated from a mycosis of the inguinal region, showing light yellow grains. Spores short, coccoid. Organism is a strict aerobe, gram-positive. Cultivable with difficulty. On glycerol agar, colonies confluent, giving a thick, snow-white covering, reverse dark yellow after two months, strong 732 MEDICAL MYCOLOGY odor, suggesting Camembert cheese, adherent. GroAvth on potato, straw yel- low, not coloring the medium. In broth, small flocei app«iar at the bottom of the tube, each being white above and yellowish white below. Medium remains clear, darkening in 7-8 months. On coagulated serum, colonies small, light yellow, becoming snow white after 5 days. There is some liquefaction, and the colony sinks in two months, showing hard yellow grains, partly as a result of incrustation by salt crystals. Gelatin liquefies in one week and the colony sinks, unless it happens to stick to the walls of the tube, when it forms a white partial pellicle. Liquid is clear, darkening after 7-8 months. Actinomyces aureus (DuBois Saint-Severin) Lachner-Sandoval, Uber Strahlenpilze 66, 1898. Streptothrix aurea DuBois Saint-Severin, Ann. Med. Nav. Colon. 63: 253- 260, 1895; Semaine Med. 15: 202, 1895. Nocardia aurea Castellani & Chalmers, Man. Trop. Med. ed. 2, 818, 1913. Oospora aurea Sartory, Champ. Paras. Homme Anim. 812, 1923. Isolated from case of conjunctivitis. Not pathogenic to laboratory animals. Hyphae 1/i in diameter, dichotomous, with helices which break up into ovoid arthrospores. Gram-positive. On agar, colonies thick, dry, acuminate, verrucose, convoluted, covered with a white powder. Reverse yellow. On potato, colony dry, verrucose, thick and yellow (color like that of gold chloride). Spores white. On coagu- lated serum, colonies gray, moist, horny, liquefying substrate. On human serum, colony flat, thin, circular, with alternating zones gray depressed and white powdery ; reverse yellowish. On gelatin, there is a thick, contorted, superficial colony, the gelatin becoming liquefied in a few days, with the center depressed and darkening, the white powderj^ portion submerged and yelloAv- ish. In broth, small spherical masses form on the surface and settle to the bottom. Gelatin and coagulated serum liquefied. Actinomyces luteolus (Foulerton & Jones), Brumpt, Precis Parasitol. ed. 4, 1192, 1927. Streptothrix luteola Foulerton & Jones, Trans. Path. Soc. London 53: 75, 1902. Oospora luteola Sartory, Champ. Paras. Homme Anim. 812, 1923. Nocardia luteola Castellani & Chalmers, Man. Trop. Med. ed. 2, 818, 1913. Discomyces luteolus Verdun & Mandoul, Precis Parasitol. 750, 1924. Isolated from a ease of purulent conjunctivitis wtih sloughing of cornea. Not pathogenic to ordinary laboratory animals. The organism is an aerobe, showing only scanty growth under anaerobic conditions. Optimum temperature 37° C. Gram-positive, not acid-fast. On peptone maltose agar, growth is a faint drab or whitish, later becom- ing faintly yellow. Colony on potato, cafe-au-lait in color, with no pigmenta- tion of the medium. Efflorescence only at 37° C. GroAvth on horse serum, dry and wrinkled, with a drab growth sunk in the medium. On gelatin, colony opaque white faintly tinged with yellow, sinking into substrate as it liquefies. ACTINOMYCETEAE 733 No pigmentation of substrate. Litmus milk is slightly acidified, but there is no coagulation. Litmus is slowly decolorized and the milk clears. Gelatin and serum somewhat liquefied. On peptone beef broth, sediment of discoid colonies. Slightly feculent odor in old cultures. The following two species belong in this group but are too briefly de- scribed to place more definitely. Actinomyces appendicis (Chalmers & Christopherson) Brumpt, Precis Parasitol. ed. 4, 1189, 1927. Nocardia appendicis Chalmers & Christopherson, Ann. Trop. Med. Para- sitol. 10: 256, 1916. Streptothrix hominis III, Foulerton 1906, 1910. Isolated from a case of appendicitis and right iliac abscess. Colonies on agar are yellow, on potato, brown, aero-anaerobic. Gelatin and coagulated serum liquefied. Actinomyces Salvati Langeron, Bull. Soc. Path. Exot. 15: 526-528, 1922; Fontoynont & Salvat, Bull. Soc. Path. Exot. 15: 596-607, 1922. Isolated from generalized nodular lesions in the Madagascar rat. Mycelial elements rare in pus. In culture, mycelium filamentous, branched, fragmenting into coccoid or bacilloid arthrospores. Hyphae swell terminally. Sometimes chlamydospores form from terminal thickenings. Gram-negative, not acid-fast. Cultures at 37°-38° C. brownish, more or less dark, finally covered with a chalky white efflorescence. Odor mouldy. Cultures on agar with glycerol, glucose, or maltose very adherent to the substratum. Organism grows well on carrot, turnip, potato. Coagulated serum is slowly liquefied with a repulsive odor. Actinomyces gedanensis (Scheele & Petruschky) Brumpt, Precis Para- sitol. ed. 4, 1196, 1927. Sfreptothrix sp. Scheele & Petruschky, Verh. Kong. Innere Med. 550, 1897. Streptothrix gedanensis I Scheele & Petruschky. Nocardia gedanensis Chalmers & Christopherson. Ann. Trop. Med. Parasitol. 10: 255, 1916. Isolated from sputum and pus. On agar, colony completely white with strong odor adherent to substrate. On gelatin, colony white. No growth on potato or glucose agar. Actinomyces catarrhalis (Sartor^^ & Bailly) Brumpt Precis Parasitol. ed. 4, 1195. 1927. Oospora catarrhalis Sartory & Bailly in Bailly, Contr. a I'Etude des Mycoses Pulmonaires. These Univ. Strasbourg Fac. Pharm. 4: 57-82, 1921. Isolated from sputum in a case of cough. Pathogenic to guinea pigs. H^'phae 0.4-0.5/x in diameter, long, branched, curved. Sporiferous branches close together. Spore chains coiled, but older hyphae not helical. Spores 0.5- 0.6/i in diameter. Optimum temperature between 32°-35°, but growth very good at 37° C. 734 MEDICAL MYCOLOGY No growth on carrot, potato, banana starch, turnip, artichoke or Raulin agar. Grows on gelatin, agar, sugar agars, prune decoction with agar or gelatin. Sabouraud, beef broth, and maltose media are best. On gelatin, small punctiform grayish colony. On carrot, small punctiform colonies, grayish white, then after a week becoming cacoa brown, discrete. Colonies similar on banana, but remain white. In beef broth, a thick mucous deposit, no pellicle. On other liquid media colonies are similar. Milk is coagulated on the second day and then the coagulum digested. Gelatin not liquefied. Actinomyces Matruchoti (Mendel) Nannizzi, Tratt. Micopat. Umana [Pol- lacci] 4: 51, 1934. Cladothrix Matruchoti Mendel, C. R. See. Biol. 82: 583-586, 2 figs., 1919. Isolated from the roots of a decaying tooth with tumefaction. Slightly pathogenic to rabbit on intramuscular injection. Hyphae straight or sinuous, variable in length, 100 x 0.3-0.5/i. approxi- mately, nonseptate, generally not branched. Branching false (?). In old cul- tures, hyphae fragment into segments 4-5/x long. Organism a strict aerobe. Gram-positive, not acid-fast. On ascitic agar, colonies gray, opaque, irregularly circular, adherent, not over 1 mm. in diameter, central zone elevated, yellowish, slightly umbilicate. Later, periphery becomes transparent and radiate. On potato and carrot, growth poor except in liquid of condensation. No growth on gelatin. In broth and peptone solution, small yellow grains are suspended in the liquid, then fall to the bottom, leaving the liquid clear. Litmus milk becomes red, with lower layer decolorized. Acid formation with glucose, lactose, fructose, su- crose. Milk coagulated, gelatin not liquefied. Actinomyces Rodellae Dodge, n. sp. Streptothrix sp. Rodella, Centralbl. Bakt. I, 84: 450-461, 1920. Isolated from abscesses of the tooth and jaw. Hyphae short, branched. Gram-positive and negative. On agar, colonies small, irregular, dry, efflorescent. No growth on potato or gelatin. Colonies on serum depressed, wavy, compact, almost leathery, dirty gray, adherent, though whole colony may be removed without breaking. In broth, growth slow with deposit settling and medium clear. In sugar broth as in simple broth, but growth more abundant, granules up to the size of petit pois. Milk is digested and cleared. The author identifies his organism with Microm,yces Hofmanni Gruber. Actinomyces Chalmersi (Froilano de Mello & St. Antonio Fernandes) Dodge, n. comb. Nocardia Chalmersi Froilano de Mello & St. Antonio Fernandes, Mem. Asiatic Soc. Bengal 7: 130, 131, 1919. Isolated from saliva of horse. Cells 3-5 X 0.5-0.7/x, gram-positive, acid-fast. Colonies on agar, dry, granular transparent at first, becoming white with white efflorescence. Similar on glycerol, maltose, and Sabouraud agar. On ACTINOMYCETEAE 735 potato, colony dry, membranous, folded, dirty white, growing along walls of tube, finally reticulate, brownish. On carrot, colony similar, folds thinner and disappearing in age ; yellowish white becoming brownish yellow. On broth, pellicle white, folded, adherent, medium remaining clear. On sugar broths similar, but folds more highly developed. On vegetable broth, pellicle grayish white, sediment sandy. Milk coagulated and digested, with yellowish white pellicle. Litmus milk decolorized. No diastatic action. Actinomyces Tarozzii (Miescher) Dodge, n. comb. Streptothrix Tarozzii, Miescher, Arch. Derm. Syphilis 124; 297-442, Pis. 20-25, 1917. Actinomyces alhus Tarozzi, Archivio Sci. Med. 33: 553-632, 1909. Isolated from a case of Madura foot with yellowish white granules. Path- ogenic to laboratory animals. Hyphae 0.5-1.5//. in diameter, repeatedly dichotomously branched, occa- sionally septate. Chlamydospores terminal, spherical, single, 4-5/x in diam- eter. This organism is a strict aerobe, grows at 15°-20°, better at 37° C. Gram-positive, not acid-fast. GroAvth on agar opaque, glassy, smooth, adherent with, sometimes, a white efflorescence. Colonies on glycerol agar velvety, snow white, soon confluent. On potato or potato glycerol, colonies hemispheric, concave below, adherent to the substrate, finally, confluent into a mammillate white, velvety colony, turning brown and black below, remaining white above. Dark color not formed on new potatoes. Velvety white colonies on gelatin. On coagulated blood serum, development is slow, with the small hemispheric colonies adherent to the sub- strate and becoming dry without the formation of aerial hyphae. In broth cultures, both beef and potato, the deposit is composed of small gelatinous colonies which are finally confluent into a mucoid mass. Surface colonies de- velop a white aerial mycelium with liquid remaining clear, moldy odor. No growth on hay infusion. Gelatin slowly liquefied. Actinomyces Ribeyro, Dodge, n. sp. Hongo artrosporado Ribeyro, Ann. Fac. Med. Lima 3: 1-5, 1 pi., 1919. Oospora sp. Sartoiy, Champ. Paras. Homme Anim. 810, 1923. Isolated from a generalized infection on the arms, legs, and chest in a patient from Cotabambas, Apurimac, Peru. There were from 30 to 40 dis- crete pustules in various stages of development. These start as small red papules, become acuminate, 1-2 cm. in diameter, yellowing with the central eminence and serous content darkening. On rupture, a dark crust forms and a lively burning pain is provoked for a few days. Finally the crust disap- pears, without leaving a sear. Not pathogenic to rabbit except on subcutane- ous inoculation, when typical nodules are produced. In tissues, coccobacilli, 1-2 x 1/x, straight or slightly curved. Mycelium fine, branched, 0.5/a in diameter, not septate. Sporulation in about 20 days. Colonies slightly elevated, creamy in consistency and color, with a white, powdery efflorescence. Very little or no growth on Sabouraud agar. In broth, 736 MEDICAL MYCOLOGY there is a fragile pellicle and sediment. Growth on gelatin creamy, then pow- dery white. Substrate not liquefied. Actinomyces keratolyticus Acton & MeGuire, Indian Med. Gaz. 66: 65-70, 3 pis., 1931. Produces cracked heels among the ryots of India. The disease known to the medical profession as keratolysis plantare sulcatum or keratodermia plan- tare sulcatum is known in Bengali as chaluni (literally, a sieve) from the pitted condition of the thick skin of the feet, or haja from the sodden condition of the skin between the toes which often splits, giving rise to the deep type of mango foe or phata, from the cracked and fissured condition of the heel. In Urdu it is called panki from its association with mud {pank). The thick homy skin of the plantar surface is dissolved in grooves. The thick sodden skin between the toes is suggestive of conditions produced by Epi- dermophyton in such positions, see p. 438. The epidermis on the sides of both toes at the interdigital cleft is thickened, white, and sodden in appearance. On separating the toes a deep fissure extends through the corium into the subcutane- ous tissues which is extremely painful and is likely to be infected by Streptococci. At other times, the infection produced an intertrigo between the web of the fingers or toes and extends to the thick palmar or plantar surface as a gyrate area of keratolysis. The cracked or split heel during the monsoon is usually due to Actinomyces keratolytica, while during cold weather it is produced by other agents. Ulcus interdigitale (Castellani 1907. Breinl 1915, Martinez & Lopez 1918) seems to be produced by the same organism. Pruritus between the toes is fol- lowed by deep fissure which gradually develops into a large oval ulcer. The margins consist of heaped up, sodden epithelium, and the base is a dull dark red color. There is little or no discharge, and these ulcers are very painful. The ulcers may also appear on the sole near the tread of the great toe and heel. Sometimes these lesions extend very deeply into the interdigital cleft and may even necessitate amputation of the toe by secondary sepsis. Breinl (1915) described an extreme form (ulcus interdigitale destruens). The lesion starts as a small fissure which gradually forms a painful ulcer and then spreads quite rapidly upward toward the toe and down to the sole. The ulcer is deep and has irregular edges, and the granulation tissue is covered by an irregular dirty gray scab. The floor of the ulcer is reddish and uneven and discharges much thick velloAv pus. The ulceration may spread upward betAveen the toes and gradually lead to complete loss of the affected toe. When healing occurs with- out amputation, the adjoining surfaces of the toes may grow together. The ulcers are very chronic and may cause considerable deformity. The disease is usually contracted by walking about barefooted on damp soil, contaminated by horse or cow manure. The ryots and maid servants who are continually walking on damp soil are prone to the disease of the feet, while the malis (gardeners) more frequently develop paronychia with greatly thickened skin margins, or onychomycosis with the base of the nails becoming I ACTINOMYCETEAB 737 brittle and moth-eaten in appearance. The lesions usually occur during the monsoon months (August to October). The disease has been twice observed in European officers of coastal vessels who were accustomed to walk about the decks with bare feet in the early morning while supervising washing the decks. Not pathogenic to experimental animals, but the disease was pro- duced on a human volunteer, and the organism recovered. The lesions clear up on treatment with a formalin lotion. Cultures inhibited by gentian violet 1 :400,000 and 5% solution used in treatment. Tissue stained by Ponder 's method shows the very slender mycelium, either nonseptate or closely septate, suggesting a string of Streptococci. The organism is best isolated on Norris' medium. The hyphae are very slender, 0.8/x, in diameter, differentiated slightly into aerial hyphae, surface runners and rooting hyphae extending into the agar. The terminal organs are of two kinds : terminal spindles, slightly curved, com- posed of 2-3 cells, probably chlamydospores (intercalary chlamydospores also occur) and conidia, 1.5/* in diameter, either singly or in groups. The radial surface runners spread centripetally, and are apparently important in deter- mining the shape of the pits and gyrate lesions, while the deep rooting hyphae penetrate as far as the prickle cell layer, open up the lymphatic spaces, and give rise to vesicles. The fungus appears to have a marked lytic action on the homy layer of the epidermis. On Norris agar, colonies in 4 days are small, pink, raised, gradually be- coming deeper in color; finally flat, dark, and moist. Aerobic. No acid or gas on any sugar tried, profuse growth on glucose and glycerol. Litmus milk was turned slightly acid, with the appearance of a pink ring at the surface ; slightly clotted with a musty odor. Actinomyces plurichromogenus (Caminiti) Dodge, n. comb. Streptothrix poly chromo gene Vallee, Ann. Inst. Pasteur 17: 288-292, 1903. Streptothrix pluricromogena Caminiti, Centralbl. Bakt. I, 44: 198, 1907. Isolated from the blood of a horse, dead of acute Pasteurella infection. Not pathogenic to laboratory animals. Hyphae branched. Organism a strict aerobe, growing at temperatures between 18° and 47° C, best at about 38° C. Gram-positive. Colony on peptone agar salmon red, shining, irregularly folded, central button with paler radiating periphery. Similar on gelatin. Growth on potato, a pale rosy gray which becomes yellowish red or even vermilion if the tube is removed from the incubator after the colony is formed. Growth folded, granu- lar, verrucose, and drs^ In peptone broth, there forms a pellicle rose salmon or deeper in color, breaking on about the eighth day and settling as a viscous ball; medium clear, but slightly colored by pigment dissolved from the pel- licle. Growth similar on milk, no coagulation. On sucrose with potassium phosphate, growth colorless, and a variety of abnormal forms appear. Glycerol added to media produced yellow to orange colonies. Gelatin not liquefied. 738 MEDICAL MYCOLOGY Actinomyces micetomae Greco, Origine des Tumeurs . . . 759-771, 1916. Streptothrix micetomae Argentinae fS, Greco apiid Durante, Segunda Obser- vacion de Pie de Madura o Micetoma en la Republica Argentina. Tesis, Buenos Aires, 1911 [reprinted by Greco, 1916]. Oospora micetomae Sartory, Champ. Paras. Homme Anim. 783, 1923. Isolated from a case of mycetoma pedis. Organism not pathogenic to rab- bits and guinea pigs. Mycelium 0.5-l^t, not septate, branched. Gram-positive, not acid-fast. Obligate aerobe, growth good at 30°-36° C. On 4% glucose agar, colonies grayish white, adherent, after a month becom- ing brick red, cerebriform, margins of radiating hyphae. On Sabouraud agar, colony as in glucose agar but color brighter. On simple agar, colonies less well developed, less cerebriform, yellowish grayish. On potato and carrot, little growth, moist, color as on simple agar finally with white efflorescence. In broth no growth ; on a decoction from 100 gm. meat, 4 gm. glycerol, water 100 gm., and potatoes at the rate of two large potatoes per liter, sterilized and filtered, or a similar broth with glucose replacing the glycerol, many small colonies appear, with cottony peripheries. No growth on milk. No growth on gelatin. Actinomyces verrucosus (Miescher) Adler, 1904 fide Nannizzi, Tratt. Micopat. Umana [Pollacci] 4: 46, 1934. Streptothrix verrucosa Miescher, Arch. Denn. Syphilis 124: 314, 1917. Isolated from mycetoma pedis with yellow grains. Medication with iodine proved ineffective, healed after application of pyrogallol — resorcin — gelante. Pathogenic to rabbit. Mycelium 0.3-0.5/i,, coiled, much branched, early fragmenting. Aerobic, growth good on sugar media, colonies easily separable, granular, verrucose, salmon red ; aerial hyphae not constant. Milk not coagulated or digested. Optimum temperature 37° C. Hiibschmann (1921) referred here a species which he had isolated from yellow grain mycetoma of the face, which was not pathogenic to experimental animals. Perhaps his organism should be referred to the group of A. Rodellae, since there are several discrepancies between the descriptions of Miescher and of Hiibschmann. The latter described his organism as follows : Streptothrix verrucosa? Hiibschmann, Arch. Derm. Syphilis 130: 341-352, 1921. Mycelium in tissues 0.2-0.5/a, in cultures 0.5-0. 7/t. Optimum temperature 37° C, scarcely any growth at 25° C. Growth equally good under aerobic and anaerobic conditions. On mal- tose agar, colonies hemispheric, smooth, shining, transparent at first, later whitish, adherent. Red brown pigment on coagulated serum. No growth on potato or gelatin, slight growth on milk and egg. On liquid media, granular floccose sediment, liquid clear, no pellicle. ACTINOMYCETEAE 739 Actinomyces Sommeri Greco, Origine des Tumeurs . . . 726-758, 1916. Streptothrix Madurae Greco, Primer Caso de Pie de Madura o Micetoma en la Republica Argentina, Tesis, Buenos Aires, 1904. Streptothrix micetomae Argentiiiae oc Greco Apud Durante, Segunda Ob- servacion de Pie de Madura o Micetoma en la Republica Argentina, Tesis, Buenos Aires, 1911. Oospora Sommeri Sartory, Champ. Paras. Homme Anim. 783, 784, 1923. Isolated from a case of mycetoma pedis in Argentina. Pathogenic to guinea pigs and rabbits. Hyphae about 1/^ in diameter, occasionally up to 1.2/x; spores 2-3/a long; branching, dichotomous; spore chains helical. Gram-positive, acid-fast, aerobic. On simple agar, colonies pale rose, hard; dry, finally light grayish rose, colonies rugose or craterif orm ; medium mahogany or port wine in color. On glycerol agar, colonies elevated 1-2 mm., umbilicate, rose, yellow, or orange, rose efflorescence, 3-4 times as large as on simple agar. On potato, colonies small, brick red, dry, medium not colored ; on carrot, colony similar but better growth, more rose color. On serum, colonies small, grayish white, few; on gelatin, colonies small, white, poor, no growth in stab. On coagulated egg white, colonies grayish rose with white margin ; on e^^ yolk, similar but more rose color, with a light gray granular efflorescence. In broth with peptone, pellicle light yellowish rose or gray, smooth ; colonies from the lower side set- tling, no cloudiness, becoming mahogany colored. In lactose broth, similar, pellicle yellowish straw color. In glycerol broth, pellicle folded and granular rose color, little or no sediment. Growth in milk good, pellicle orange, milk not coagidated but slowly digested. In liver infusion, pellicle dry, more or less thick, light yellowish rose, then brick red, sediment grayish white, granu- lar, medium not changed in color. In potato decoction, pellicle thin, fragile, white, sediment as above. In yeast infusion, no pellicle; medium remains clear but darkens. It is possible that the following unnamed organism belongs to this species. Streptothrix sp. Bloch, Cor. Bl. Schweiz. Aerzte 46: 270, 271, 1916. Isolated from a white grain mycetoma pedis. Pathogenic to rabbits, pro- ducing lesions similar to those on man. Growth aerobic, abundant, brick red color. Actinomyces Madurae (Vincent) Lachner-Sandoval, tJber Strahlenpilze 64, 1898. fOospora indica Kanthack, Jour. Path. Bact. 1: 140-162, Pis. 10-12, 1892 [organism seen in tissues but not cultivated]. Streptothrix Madurae Vincent, Ann. Inst. Pasteur 8: 129-151, 1894. Nocardia Madurae Blanchard in Bouchard, Traite Path. Gen. 2: 868, 1896. Discomyces Madurae Gedoelst, Champ. Paras. Homme Anim. 169-173, 1902. Nocardia indica Chalmers & Christopherson, Ann. Trop. Med. Parasitol. 10 : 255, 1916. Discomyces indicus Neveu-Lemaire, Precis Parasitol. Hum. 42, 1921. 740 MEDICAL MYCOLOGY Oospora Madurae Lehmann & Neumann, Atlas Grundriss Bakt. 2: 388, 389, 1896. Isolated from white grained Madura foot. Animal inoculations negative. Mycelium 1-1. 5/* in diameter. Arthrospores in the form of rods or spheres. Organism is aero-anaerobe but grows best anaerobically. Gram-positive, not acid-fast. On glycerol agar, colonies very adherent to the substrate, longitudinally folded, discoid, crateriform, the periphery yellowish at first, later becoming rose or even red, the crater remaining white. Colonies on maltose peptone agar are small, discrete, yellowish white, waxy, wrinkled, becoming pinkish; the medium brownish. Colonies on potato are spherical, becoming confluent in a heaped-up, irregular, mammillate mass, creamy yellowish white to red- dish, depending on the acidity of the medium, with a white efflorescence. The potato is eroded. According to Vincent there is no growth on serum. Mace claims growth similar to that on agar. No growth on egg albumen. Opaque, whitish, raised, discrete colonies on gelatin. In broth, opaque, white, floccose spheres appear at the bottom ; liquid remains clear. In vegetable decoctions (15 gm. potato, hay, or straw per liter), spherical flocci on sides or bottom, liquid clear but turns somewhat brown and becomes alkaline. Milk not coagulated but peptonized after 20 days. Gelatin not liquefied. Actinomyces bahiensis (Piraja da Silva) Brumpt, Precis Parasitol. ed. 4, 1195, 1927. Discomyces bahiensis Piraja da Silva, Brasil Med. 33: 81-83, 1919; Mem. Inst. Butantan 1: 187-207, Pis. 34-38, 1918-1919. Oospora hahiensis Sartory, Champ. Paras. Homme Anim. 784, 1923. Isolated from yellow grain maduromycoses. Mycelium 1-1.4/* in diameter, arthrospores observed, also deeply staining granules which the author thinks may be endospores. Colonies on potato with whitish bloom, at first rose color then deeper red, becoming very dark in old cultures, shape of a mulberry. On yam, growth very slow, colonies rose colored, humid; on carrot, whitish, more humid, be- coming light rose color and the medium finally assuming the same color. On liquid media, forming white granules which collect on the walls and bottom of the tube. No growth on maltose, glycerol, or crude sugar agar. Actinomyces Leishmani (Chalmers «fc Christopherson) Sartory & Bailly, Mycoses pulmonaires 253, 1923. Streptothrix sp. Birt & Leishman, Jour. Hyg. 2: 120-128, PI. 1, 1902. Nocardia Leishmani Chalmers & Christopherson, Ann. Trop. Med. Para- sitol. 10: 255, 1916. Isolated from empyema of the lungs. Pathogenic to guinea pigs, rat, and rabbit. Ultimate branches of hyphae break up into ovoid arthrospores. Organism a strict aerobe, growing between 22° and 46.5° C, best at 37° C. Growth ACTINOMYCETEAE 741 ceases at 50°, although five minutes at 75° C. did not kill it. Acid-fast, losing this property in old cultures, all save the arthrospores, which remain gram- positive and acid-fast. On agar, a snow white, dry efflorescence forms, becomes delicate pale coral pink, is not adherent. On potato, with or without glycerol, there is copious dry, chalky growth, circumscribed, granular, verrucose in age ; sur- face never wrinkled. On the third day, it assumes a coral tint which is espe- cially evident under the efflorescence. On gelatin, small, white, circular colo- nies at the surface. In broth, spherical white floating colonies, the emerged portion turning a powdery pink, the remainder white and woolly. No pellicle but a slight ring which settles to the bottom. Medium clear and odorless. Milk not coagulated but digested, becoming alkaline. No growth on serum. Gelatin not liquefied. Actinomyces carneus (Rossi-Doria) Gasperini, Centralbl. Bakt. 15: 684, 1894. StreptotJirix carnea Rossi-Doria, Ann. Inst. Ig. Sperim. Univ. Roma 1: 399-438, PI. 8, 1891. Nocardia carnea Castellani & Chalmers, Man. Trop. Med. ed. 2, 818, 1913. Discomyces carneus Neveu-Lemaire, Precis Parasitol. Hum. 42, 1921. Osopora carnea Sartory, Champ. Paras. Homme Anim. 810, 1923. Originally isolated from the air. Reported by Boldoni from a case of chronic bronchitis. Pathogenic to rabbit and guinea pig, producing pseudo- tuberculosis. Gram-positive. On agar, colonies small, globular, flesh or orange red in color, with rose white efflorescence. On potato, growth granular, color as on agar, but efflorescence reddish. In broth, scales at the surface, flocci at the bot- tom of the tube. In milk, flesh-colored growth on surface. No coagulation. Colonies on gelatin radiate with pink efflorescence, no liquefaction. Actinomyces minutissimus (Burchard) Brumpt, Precis Parasitol. ed, 4, 1199, 1927. Microsporon minutissimum Burchard & Barensprung in Uhle & Wagn. Pat. Gen. 1859; Balzer, Ann. Derm. Syphiligr. II, 4: 681-688, 1883. Trichothecinm sp. Neumann, 1868. Microsporon gracile Balzer, Ann. Derm. Syphiligr. II, 4: 681, 1883. [Balzer refers to this name as previously used (cf. Besnier) in France. No reference to publication.] Sporotrichum minutissimum Saccardo, Syll. Fung. 4: 100, 1886. Microsporioidcs minutissimus Neveu-Lemaire, Precis Parasitol. 1906. Discomyces minutissimus Verdun, Precis Parasitol. 1907. Oospora minutissima Ridet, Oospora et Oosporoses 68, 1911. Nocardia minutissima Verdun, Precis Parasitol. 1912 ; Castellani & Chal- mei-s, Man. Trop. Med. ed. 2, 819, 1913. Reported as the etiologic agent of erythrasma. 742 MEDICAL MYCOLOGY Mycelium 0.6-1.3/a, rarely branched, easily dissociating into bacilliform arthrospores. Aero-anaerobe. Micheli reports brownish growth on gelatin, wine red on agar. Ducrey reports brownish red on agar. Actinomyces Jollyi (Vuillemin) Brumpt, Precis Parasitol. ed. 4, 1196, 1927. Nocardia Jollyi Vuillemin apud Jolly, Rev. Med. de I'Est 48: 42, 43, 1920. Oospora Jollyi Sartory, Champ. Paras. Homme Anim. 770, 1923. Isolated from a case clinically resembling bubonic plague in which the author was unable to demonstrate the plague bacillus. Hyphae 0.2-0 A/x in diameter. No spores seen. Organism aerobic, optimum temperature 37° C. Gram-negative, acid-fast. Colonies on Sabouraud agar, potato glycerol, and carrot are white, deli- cate, velvety with deeper growth a compact, fleshy mass v/hich erupts smooth, salmon-colored mounds through the velvet. Growth on gelatin very slow. No liquefaction. Actinomyces Freeri (Musgrave & Clegg) Bergey, Man. Det. Bacteriol. 346, 1923. Streptothrix Freeri Musgrave & Clegg, Philippine Jour. Sci. B. 2: 477- 511, Pis. 1-4, 1907. Discomyces Freeri Neveu-Lemaire, Precis Parasitol. Hum. 42, 1921. [By Clegg & Hobdj^, 1915, said to be a synonym of Streptothrix Eppingeri Claypole?] Oospora Freeri Sartory, Champ. Paras. Homme Anim. 785, 786, 1923 [Castellani & Chalmers, Man. Trop. Med. ed. 3, 1058, 1919, give this as a synonym of Nocardia Asteroides, also Pollacci & Nannizzi, I Miceti Pat. 6: No. 52, 1927]. Isolated from a case of mycetoma. Pathogenic to monkeys, dogs, and guinea pigs. Hyphae branching, club forms in tissues ; acid-fast, gram-posi- tive. Optimum temperature 37° C. Colonies on agar are smooth, white, glistening ; on glycerol agar, elevated, sometimes umbilicate, yellowish with pink periphery. On potato, growth good, elevated, delicate pink with yellow center. Starch not hydrolyzed, milk and gelatin not digested. Actinomyces Hofmanni Gruber, Centralbl. Bakt. 16: 363, 1894. Oospora Hofmanni Sauvageau & Radais, Ann. Inst. Pasteur 6: 251, 1892. Micromyces Hofmanni Gruber, Trans. Int. Cong. Hyg. Derm. VI, 2: 65, 1891 (1892) ; Arch. Hyg. 16: 35-52, PI. 1, 1892. Streptothrix Hoffmanni Petri, N. Giom. Bot. Ital. n. s. 10: 592, 1903. Nocardia Hoffmanni Chalmers & Christopherson, Ann. Trop. Med. Para- sitol. 10: 268, 1916. Isolated from a sample of vaccine. Pathogenic to mice and rabbits. Hyphae less than Ifx in diameter, variously curved and thickened, branched, not septate. Only secondary hyphal branches break up into spores. ACTINOMYCETEAE 743 Occasionally tips of hypliae are swollen to five or six times liyphal diameter, probably as a degeneration phenomenon rather than for spore formation. Optimum temperature 37° C, no growth at 17°-22°, nor at 40° C. Organism aerobic. Gram-positive. On solid media, e.g., nutrient agar with sugar, colony shows an irregularly indented contour, gray Avhite, brownish in age — light brown by transmitted light, dark brown when opaque. Surface dull, uneven, folded regularly in radiating lines in old colonies, adherent, circumscribed, hemispheric or ir- regularly verrucose in appearance. No growth on gelatin or potato. Coagu- lated blood serum is an unsuitable medium unless sugar is added. A Avhite, powdery deposit appears in liquid media, the medium remaining clear. Alco- hol somewhat oxidized to acetic acid. Actinomyces niger (Rossi-Doria) Brumpt, Precis Parasitol. ed. 4, 1206, 1927. Streptothriic chromogena Gasperini (fide Rossi-Doria). Streptothrix nigra Rossi-Doria, Ann. 1st. Ig. Sperim. Univ. Roma N. S. 1 : 399-438, PL 8, 1891. [cf. Foulerton & Jones 1902.] Nocardia nigra Castellani & Chalmers, Man. Trop. Med. ed. 2, 1913. Isolated from air by Rossi-Doria. Colony white, substrate blackened on sugar media. Colonies on egg white remain humid, not becoming powdery. Milk is coagulated and digested in about 10 days, becoming yellowish red and then brown. No acid produced. Actinomyces lingnalis (Weibel) A. Sartory & A. Bailly, Mycoses Pulmo- naires 252, 1923. Vibrio lingualis Weibel, 1888. Spirosoma lingualis Migula, 1892. Streptothrix lingualis Bajardi, Centralbl. Bakt. I, 36: 129-137, 1 pi., 1900. Found in mouth. Of doubtful pathogenicity to guinea pig. On a gelatin plate, after 12-24 hours at 18° C, small punctiform colonies appear throughout the whole medium. On magnification, these colonies ap- pear yellowish with irregular margins from which project branched hyphae. On agar plate, growth similar except that individual isolated branches rarely project from the colonies into the medium. On gelatin stab, surface growth elevated, round, with center depressed, not spreading far from point of in- oculation. Margins wavy. Along stab, short dendritic projections with clavate ends perpendicular to line of stab, the length of these projections decreasing with distance from the surface, giving appearance of a test tube brush. Growth on agar stab somewhat similar except that colony is more extended, with fine wavy margins and small elevations on the surface which appear to be due to overlapping colonies. Color dirty white, gradually be- coming yellowish. Along the stab a band with finely lobate margins. Streak culture on agar slant shows a dirty white covering with surface at first moist, later dry, not developing far from the line of inoculation. In older cultures, appearance finely verrucose and canary yellow in color. Warts like pinheads 744 MEDICAL MYCOLOGY with rounded, projecting points. If the streak is so made that the colonies do not become confluent, they give the typical mammillate appearance. On potato, growth is slow and limited, growth only detectable by dampness of the surface of the potato. Later a dry, linear growth with even margins. In broth, at first a slight cloudiness with small amount of pulverulent deposit, medium later clearing with a granular sediment, granules becoming flocculent in old cultures. In 10-day-old broth cultures, there is a tendency to pellicle formation with a fragmentary surface covering. In any case, a ring forms with a dirty white color, later turning yellowish, easily broken up. No indol forma- tion. No fermentation with any sugars. Milk not coagulated, gelatin gradually liquefied. Organism grows between 18° and 42° C, optimum temperature being 37° C. Actinomyces phenotolerans C. H. Werkman in Gammel, Arch. Derm. Syphilol. 29: 286-297, 1934. Isolated from granuloma in man. Pathogenic for guinea pigs when in- oculated into the lungs. Mycelium straight with lateral branching, less than 1^ in diameter, chains of conidia abundant, prostrate, racemose type without helices, very acid-fast, hyphae gram-positive, spores gram-negative. On Krainsky's glucose agar, colonies small, round, adherent to medium; surface powdery, white to grayish white, becoming orange in old cultures; no odor, medium not discolored. On potato, colonies small, chalky, confluent into orange membrane with odor of market garden soil. On gelatin, white surface growth with very slow liquefaction. On serum, cream colored surface growth with arborization. In Dunham peptone broth, an abundant flaky sedi- ment. Good growth in broth with 0.5 per cent phenol. On sugar broths, growth good with most sugars, sediment flaky with solution becoming- alkaline. Litmus broth becomes alkaline with surface growth. No amylase on starch agar. Nitrates reduced to nitrites, no tyrosinase, no soluble pig- ment. Aerobic, optimum temperature 37° C. Actinomyces Bellisari, Dodge, n. nom. Streptothrix alba Bellisari, Ann. Ig. Sperim. 14: 467-483, 1904. Oospora alba Sartory, Champ. Paras. Homme Anim. 819, 1923. Isolated in a warehouse in Naples from the dust of cereal coming from California. Pathogenic to rabbits. On agar, colonies amber yellow, very small, not confluent, spores produc- ing a whitish efflorescence which seems limited to the periphery. On potato, growth thick and folded, white, after a week with brownish fissures. No diffusion of color into the medium. On gelatin stab, small white granules ap- pear. These are larger near the surface where the growth expands gradually into a folded white pellicle covering the surface. In broth, colonies small, spherical, isolated, settling and leaving the medium clear. Gelatin slowly liquefied, liquefaction beginning about the twentieth day. k ACTINOMYCETEAE 745 Actinomyces convolutus (Chalmers & Christopherson) Brumpt, Precis Parasitol. ed. 4, 1195, 1927. Nocardia convoluta Chalmers & Christopherson, Ann. Trop. Med. Parasitol. 10: 223-282, PZs. 8-ii, 1916. Discomyces convolutus Neveu-Lemaire, Precis Parasitol. Hum. 44, 1921. Oospora convoluta Sartory, Champ. Paras. Homme Anim. 769, 1923. Isolated from a yellow grain mycetoma in the Sudan. Not pathogenic for monkeys or other laboratory animals. Mycelium less than l/m in diameter. Spores in chains, spherical, 1/i, in di- ameter. No claviform formations. Organism an aero-anaerobe. Gram- positive, not acid-fast. Optimum temperature 30° C, fair growth anywhere between 22° and 37° C. Colonies warm buff (Ridgway) before the white efflorescence appears, convoluted, not pigmenting agar media. Growth best on alkaline media. Colony on Sabouraud maltose agar more radiate than convoluted. On potato, good development, convoluted, moist, warm buff with an efflorescence. Colo- nies on gelatin lig'ht buff, small, round ; medium not pigmented. Similar on serum. White efflorescence on milk ; medium not coagulated or digested but is made slightly more alkaline. In peptone broth with glucose, white, non- cohering flocci are deposited, with liquid remaining clear. No acid or gas with glucose, maltose, lactose, sucrose, raffinose, mannitol, or salicin. Serum liquefied. Gelatin not liquefied. Actinomyces serratus (Sartory, Meyer & Meyer) Dodge, n. comb. Actinomyces asteroides var. serratus Sartory, Meyer & Meyer, Ann. Inst. Pasteur 44: 298-309, 1930. (See also C. R. Acad. Sci. 188: 745-747, 1929.) Isolated from a case of actinomycosis of bones with yellow grains. Patho- genic for guinea pigs, but no bone lesions obtained, not even with dog. Filaments fragile, branched, segmenting easily, 0.3-0.6/x in diameter. Ar- throspores Ifx in diameter. Gram-positive, spores acid-fast. On agar, colonies whitish, verrucose, becoming ochraceous in age, con- fluent into a folded pellicle. On potato, colonies mammillate, whitish, lumpy, becoming powdery, yellowish gray. Serum not liquefied, milk not coagulated but slowly digested. Gelatin not liquefied, no growth in depths. Cultural characters suggest A. convolutus Chalmers & Christopherson, but this organism differs in action on serum. Actinomyces Donnae Dodge, n. sp. Streptothrix sp. Donna, Ann. Ig. Sperim. 14: 449-459, PI. 3, 1904. Isolated from sputum in a pulmonary infection. Pathogenic to rabbits. Organism aerobic. Colonies small, 2 mm. in diameter, opaque, yellowish white. Growth less on gelatin. In beef broth, there is turbidity with floccose sediment and some evolution of gas. Liquid finally clears. At lower tempera- ture there is no turbidity, and broth remains alkaline. Gelatin not liquefied. 746 MEDICAL MYCOLOGY Actinomyces deBerardinis Namslowsky, Centralbl. Bakt. I, 62: 566, 1912. Streptothrix sp. Berardinis, Ann. Ottalniol. Lavori Clin. Oculist. Napoli 33: 914-924, 1 pi., 1904. Isolated from a case of conjunctivitis. Following accidental inoculation of the eye, corneal ulcers were produced. Organism pathogenic to rabbits. Long spore chains formed. On agar, there is a delicate yellow growth. On potato, colony is elevated in the center with margin undulate and reddish. Small, yellowish white colonies on gelatin. Broth becomes uniformly turbid and of a delicate yellow color, eventually tvirning brownish. Only moderate growth in glycerol broth, better with glucose or lactose added. Growth good on rabbit serum, poor on milk. Gelatin not liquefied. This organism is close to A. Gruheri or A. asteroides (S. Eppingeri). Actinomyces farcinicus (Trevisan) Gasperini, Ann. 1st. Ig. Sperim. Univ. Roma 2: 2, 1892. Bacillus du farcin Nocard, Ann. Inst. Pasteur 2: 293-302, Pis. 7, 8, 1888. Nocardia farcinica Trevisan, I Genere e Species delle Batteriacee 9, 1889. Streptothrix farcinica Rossi-Doria, Ann. 1st. Ig. Sperim. Univ. Roma. N. S. 1: 404, 1891. Oospora farcinica Sauvageau & Radais, Ann. Inst. Pasteur 6: 248, 1892. Actinomyces hovis farcinicus Gasperini, Centralbl. Bakt. 15 : 684, 1894. Streptothrix Nocardi/i Foulerton & Jones, Trans. Path. Soc. London 53: 57, 97, 98, 1902. Discomyces farcinicus Gedoelst, Champ. Paras. Homme Anim. 167-169, 1902. Nocardia alhida Chalmers & Christopherson, Ann. Trop. Med. Parasitol. 10: 271, 1916. Found in farcy of oxen. Pathogenic for guinea pig, ox, and sheep, not very pathogenic for rabbit, goat, dog, cat, horse, or ass. Hyphae long, branched, 0.25/a in diameter. Arthrospores cylindrical, 2/* long. Organism a strict aerobe. Gram-positive, acid-fast. On maltose peptone agar, colonies opaque, dull, granular, circular, flat, becoming confluent and mammillate, yellowish white with a white efflorescence of arthrospores. Colonies on potato dry, grayish buff to yellow, granular, confluent into a folded or verrucose pellicle with white efflorescence, margins elevated. On serum, growth is slower and humid, otherwise as on agar. On gelatin, growth slow, slightly elevated, medium brownish when acid. In pep- tone beef broth, whitish irregular flocculent masses appear at the bottom with scales on the surface, resembling drops of fat (especially on glycerol broth). Similar in litmus milk. No alteration in medium or coagulation. Serum not liquefied. Gelatin liquefied. Actinomyces japonicus Caminiti, Centralbl. Bakt. I, 44: 198, 1907. Streptothrix sp. Aoyama & Miyamoto, Mitteil, Med. Fak. K. Jap. Univ. Tokio 4: 231-276, Pis. 16-18, 1901. k ACTINOMYCETEAE 747 Cultivated from the pus in a fatal infection of the lungs. The sputum was too contaminated with bacteria to permit of easy isolation of the fungus. Pathogenic to guinea pigs. Authors finally decided the infection was a mix- ture of tuberculosis and actinomycosis. Terminal portions of the hyphae coiled. Aerobic. Gram-positive. Colonies star-shaped, white at first, yellowing and elevated in age, size of lentil, surface rough, center darker, opaque, yellowish, dry. On potato, chalky white granules, confluent, center of colony brownish yellow with chalky white efflorescence, finally brownish. Also grow Avell on Colocasia antiquorum, Neluml)ium setosum, carrot, and Dioscorea japonica, on the first and last sub- strate not adherent. On blood serum, growth not rapid, colonies yellowish and flat. On broth gelatin, growth poor. On broth and Piorkowsky urine-peptone, small gray white or chalk white nodules, often confluent, settling to the bot- tom, no turbidity, never yellowish in color. Surface of milk yellows in 2 days, milk not coagulated or digested. Actinomyces pretorianus (Pijper & Pullinger) Nannizzi, Tratt. Micopat. Umana [Pollacci] 4: 38, 1934. Nocardia pretoriana Pijper & Pullinger, Jour. Trop. Med. Hyg. 30: 153- 156, 2 pis., 1927. Isolated from a mycetoma of the shoulder showing yellow grains, 0.5 mm. in diameter, possessing clavate bodies. Pathogenic to guinea pig. Hyphae long, O.dfi in diameter, showing marked branching. Organism a strict aerobe, growth best at 37° C, but possible at 22° C. Growth circumscribed, in general tending toward orange in pigmentation, no efflorescence, no pigmentation of the medium, adherent. No diastase present. No growth on Sabouraud agar. On potato and simple agar, growth largely buff and elevated, in parts convoluted and orange. Better growth when glyc- erol is added. Growth on carrot convoluted, orange ; on blood agar, buff. On gelatin or blood serum, growth poor, white to orange. In simple broth, glyc- erol broth, or sugar broth, thick, convoluted, bright orange pellicle forms. In hay extract, gray surface films showing small round tufts and a radial struc- ture. Surface of litmus milk is orange, convoluted. No acid or fermentation with any of the usual sugars. Milk coagulated after 10 days. No liquefaction of gelatin or coagulated serum. Actinomyces miacrodipodidarum (Fox) Dodge, n. comb. Nocardia macrodipodidarum Fox, Disease in Captive Wild Mammals and Birds, 570-595, 1923. Found in lumpy jaw with septicemia and gasteroenteritis of kangaroos. Inoculations into experimental animals negative. Hyphae up to Ifi in diameter. Gram-positive. On blood serum, colonies opaque, pale yellow, circular, discrete, slightly depressed, irregular; no umbilication in center; smooth and slightly glisten- ing for several days, then wrinkled and twisted. Growth on potato is dirty yellow, much wrinkled, friable, adherent. A tough wrinkled scum forms on 748 MEDICAL MYCOLOGY the surface of gelatin. Litmus milk shows no change in 6 days, then slight alkalinity with thin pellicle on surface; caseinogen digested. In broth, sur- face growth only as a wrinkled, pale yellow pellicle with faint turbidity in medium. No action on sugars. Actinomyces canis (Rabe) Gasperini, Centralbl. Bakt. 15: 684, 1894. Cladothrix canis Rabe, Berlin. Tierarztl. Woch. 1888. Oospora canis iSartory, Champ. Paras. Homme Anim. 821, 1923. Streptothrix caprae Silberschmidt (1899), referred here by Chalmers & Christopherson. Pathogenic for rabbit and guinea pig (subcutaneous and intravenous in- jections produce tubercles with giant cells, rapidly becoming cheesy). Acid-fast, aero-anaerobic. Agar colony dry, verrucose, yellowish white with white efflorescence; some colonies crateriform. On potato, small whitish colonies which become rose brown after a while; isolated colonies round, dry, crateriform. Colony on gelatin, brownish at the center, clear at the periphery, formed of very slender radiating filaments. In broth, concave dry discs on surface, whitish, small, farinose; slight deposit, liquid clear. On milk, pink pellicle at the sur- face, no coagulation. Actinomyces caprae (Silberschmidt) Nannizzi, Tratt. Micopat. Umana [Pollacci] 4: 50, 1934. Strepiothrix caprae Silberschmidt, Ann. Inst. Pasteur 13: 841-853, 1899. Discomyces caprae Gedoelst, Champ. Paras. Homme Anim. 174-176, 1902. Oospora caprae Sartory, Champ. Paras. Homme Anim. 813, 814, 1923. Referred to A. canis by Chalmers & Christopherson, Ann. Trop. Med. Para- sitol. 10: 255, 1916. Found in the lungs of a goat in Switzerland. Pathogenic to rabbit and guinea pig. Aerobic. Optimum temperature 33°-37° C. On 4% glycerol agar, colony dry, verrucose, shrivelled, surface brownish white with a raised, irregular, brownish white farinose center, flattened to subcrateriform. On Loeffler's serum, growth good, similar. On maltose pep- tone agar, at 37°, spreading drab color with dull efflorescence; medium dark- ening in age. On potato, colony small, whitish then rose brown, round, dry, sometimes crateriform, with a white efflorescence, finely granular, medium browned. On gelatin, colonies deep brown at center, lighter at margin, be- coming chalky white (spore formation) ; reverse pink, wrinkled. In peptone beef broth or sugared broth (2%), colonies are concave discs on surface, dry, whitish, powdery, forming pellicle by coalescing, not readily sinking. Slight grumous deposit. Medium clear. On milk, rose white pellicle, no coagula- tion, no change in pH (litmus). Neither gelatin nor serum liquefied. Actinomyces bicolor Trolldenier, Zeitschr. Tiermedizin 7: 81-109, 9 figs., 1903. ACTINOMYCETEAE 749 Nocardia hicolor Froilauu de Mello & St. Antonio Fernandes, Mem. Asiatic See. Bengal 7: lOG, 1919. Found in eerebromeningitis, bronchitis, and lymphadenitis in dog. Patho- genic to white mice, guinea pigs, rabbits, dog, with infection easy. Horse, calf, and birds infected with difficulty, cats not at all. On agar, glycerol peptone, or albumose, colonies white at first, then yel- low at the center with Avhite margins. Whole surface wavy and crateriform, crater elevated 3 mm. On potato, growth is in the form of white granules, either discrete or confluent, showing small, grayish white heads on rough spinulose surface; center finally becomes yellow gray; sweet odor developed. On gelatin, colonies white, becoming dirty yellow after several weeks, with the upper layer of the gelatin yellowed. Colonies smaller on coagulated horse serum. In liquid horse serum, small white colonies on the surface, forming a granular pellicle, and at the bottom, woolly, discrete balls. In alkaline broth, the medium remains clear, bottom growth as in serum. Surface growth shows discrete colonies with center yellow and margin white, suggesting a floating composite flower with white ray flowers. Neither coagulated horse serum nor gelatin liquefied. It is possible that the following unnamed species belongs here. mreptothrix sp. Horst, Zeitschr. Heilk. Abt. Prakt. Anat. 24: 157-176, Ph. 13, 14, 1903. Isolated from patient with pyemia and brain and bronchial abscesses. Path- ogenic to mice, guinea pigs, and rabbits. Helices on potato. No clavate forms in tissues. Organism gram-positive and acid-fast. Colonies white, powdery, becoming gray, slightly shining, 2-3 mm. in di- ameter. On glycerol agar, growth is thick, folded, moist; the efflorescence whitish, becoming yellowish, and then reddish with white margin. On serum agar, small granular colonies appear. Colony on potato thick, white, dry, fine-grained, becoming thick, wrinkled, and finally brown. Growth on beet simi- lar. On gelatin, thin, fine, granular, dry colonies. In broth, small white gran- ules from a very thin, fragile pellicle, also small floating colonies in the depths, but no turbidity. Growth much less in sugar broth, and poor in peptone solution. Organism grows on milk as a thick sulphur yellow, later orange, pellicle, without altering the medium. Gelatin not liquefied. Actinomyces brasiliensis (Lindenberg) Gomes, Ann. Palistas Med. Cirurg. 14: 150-156, 1923. Discomyces hrasiliensis Lindenberg, Rev. Med. Sao Paulo Sept. 30, 1909 ; Arch, de Parasitol. 13: 265-282, 3 figs., 1909. Nocardia hrasiliensis, Castellani & Chalmers, Man. Trop. Med. ed. 2, 816, 1913. Streptofhrix hrasiliensis Greco. Origine des Tumeurs . . . 724, 1916. Oospora hrasiliensis Sartory, Champ. Paras. Homme Anim. 786, 1923. 750 MEDICAL MYCOLOGY Isolated from a mycetoma of the thigh and leg (10 cm. below inguinal fold to upper portion of the foot with movement of the knee inhibited), Brazil. Not inoculable to ordinary laboratory animals. Hyphae O.dfi in diameter, spores 1-2/x, in diameter, spherical to ovoid. Op- timum growth at room temperature [tropical]. Aerobic. Colonies on ordinary agar opaque, chalky white to yellow, being white at the center and orange ochraceous at the margin, showing concentric zoning. Colonies on Sabouraud agar at 37° C, rose violet. On Sabouraud glucose, growth poorer than on ordinary agar, white at room temperature, rose violet at 37° C. On potato as on agar, orange yellow with radiating furrows; me- dium colored brown. Colony on serum, whitish at 37°, yellowish at room temperature. Good growth on egg. Milk shows an orange yellow growth, pellicle on surface, no coagulation. Slight growth on broth at 37°, hay in- fusion shows filamentous flocci. Gelatin not liquefied. Chalmers & Christopherson relegate this species to Actinomyces asteroides, but it differs in the nature of the grain (soft in A. hrasiliensis) and the colony on Sabouraud agar here is wholly rose -vdolet, while that of A. asteroides is yellow at the center and red at the periphery on this medium. Castellani & Chalmers, Man. Trop. Med. ed. 3, 1058, 1919, also give this as synonym of Nocardia asteroides, as do also Neveu-Lemaire, and Pollacei & Nan- nizzi. Actinomyces asteroides (Eppinger) Gasperini, Centralbl. Bakt. I, 15: 84, 1894. Cladothrix asteroides Eppinger, Wiener Klin. Woch. 3: 321-323, 1890; Beitr. Path. Anat. [Ziegler] 9: 287-328, Pis. 13, 14, 1891. Streptothrix asteroides Gasperini, 1891; Petri. N. Giom. Bot. Ital. n. s. 10: 591, 1903. Streptothrix Eppi^igeri Rossi-Doria, Ann. 1st. Ig. Sperim. Univ. Roma. N. S. 1: 399-438, PI. 8, 1891. Oospora asteroides Sauvageau & Radais, Ann. Inst. Pasteur 6: 250, 1892. Nocardia asteroides Blanchard, in Bouchard, Traite Path. Gen. 2: 811-926, 1895. Diseomyces asteroides Gedoelst, Champ. Paras. Homme Anim. 173, 174, 1902. Streptothrix Freeri Musgrave & Clegg, Philippine Jour. Sci. B. 2: 447-511, 1907, was referred here by Chalmers & Christopherson, Ann. Trop. Med. Para- sitol. 10: 255, 1916. This organism is an aerobe (Poulerton-Jones), aero-anaerobe (Chalmers & Christopherson), or anaerobe (Castellani) found in mycetoma (Castellani) in brain abscess from a case of meningitis, and in diffuse peritonitis (Mac- Callum). The original case was pseudotuberculosis with cerebrospinal menin- gitis. Pathogenic to rabbits, guinea pigs, and monkeys with pseudotuberculo- sis following intraperitoneal inoculation. Cultures soon lose their virulence. Rossi-Doria found them nonpathogenic. ACTINOMYCETEAE 751 Mycelium 0.2/x. in diameter, spores nearly spherical and in chains. On agar, colonies verrucose, becoming confluent and erateriform or ir- regularly folded, pale yellow becoming ochraceous in age, 3-4 mm. in diameter, adherent. Colonies similar on glucose or glycerol agar, but growth is more rapid. On potato, colonies are elevated and verrncose, white at first, then brick red, color darker at the center, then covered with a white conidial efflo- rescence. On gelatin and serum, culture yellow, raised and wrinkled, no pig- mentation of the medium. In peptone beef broth, liquid remains clear with a sediment of small, very thin scales at the bottom of the tube, and a thin waxy, fragile pellicle. Litmus milk is at first acidified, then becomes alkaline with the formation of a yellowish granular sediment. Litmus whey decolorized, with the formation of a thick pellicle and sediment. In a mixture of agar and hydrocele fluid, growth is thick, homogeneous, with a yellowish white film over the whole surface. Milk not coagulated. Gelatin and coagulated serum not liquefied. Actinomyces Sanfelicei (Redaelli) Nannizzi, Tratt. Micopat. Umana [Pol- lacci] 4: 51, 1934. Nocardia Sanfelicei Redaelli, 1st. Sieroterapico Milanese 7: 75-101, 121-135, Pis. 1-8, 1928. StreptothtHx acido-resistente Sanfelice, Boll. 1st. Sieroterapico Milanese 2: 327-331, 1922. Isolated from fatal lesions in rat ; source unknown. Pathogenic to guinea pig, rabbit, rat, etc. Growth at temperatures between 15° and 40° C, optimum 37° C. Aerobic, gram-positive, usually acid-fast. Growth on solid media best in those containing glycerol, colony semitrans- parent, margins irregular, surface minutely verrucose, becoming thick rugose, yellowish, not adherent. On carrot agar, colonies round, elevated, folded to cerebriform. On Petragni & Petroff medium, development is more abundant, with thick, yellowish, finely mammillate colonies, margins thin, smooth, semi- transparent, festooned. On potato as on agar. Colonies on potato glycerol yellowish red, thick, irregularly rugose, very adherent. On liquid media, thin grayish pellicle becoming thick, yellowish, and rugose; small, hard, granular colonies in sediment. No fermentation of sugars, no liquefaction of gelatin, no coagulation of milk. Slight acidity on sugar broths (except lactose). Actinomyces spumalis (Sartory) Dodge, n. comb. Oospora spumalis Sartory in Sartory & Bailly, Mycoses pulmonaires 318- 321, 1923. Isolated from sputum. Patient recovered under iodide treatment. Path- ogenic to guinea pig. Growth at 37°. Mycelium 0.4-0.5// in diameter, irregularly branched. Terminal ehlamydospores, often in chains, 0.8//, in diameter. Helices of 4-5 turns abundant in older cultures. 752 MEDICAL MYCOLOGY Growth good on carrot and maltose. Gelatin not liquefied, but a red pig- ment rapidly diffuses into the gelatin. No growth on Raulin's neutral me- dium, coagulated serum, egg albumen, potato, or potato-glycerol. Perhaps the following variety of Sartory is a synonym. Oospora pulmo7ialis var. chromogene Sartory, C. R. Soc. Biol. 74: 166-168t 1913. Isolated from sputum of a patient suspected of having pulmonary tuber- culosis. Pathogenic to guinea pig. Mycelium branched, spores O.S/x in diameter. On maltose agar, colonies 1 cm. in diameter, white to cream white. On maltose gelatin, colony punctiform, red pigment diffusing into the medium, no liquefaction. No growth on Raulin's neutral medium, coagulated serum, egg albumen, or potato. Actinomyces flavus (Chester) Dodge, n. comb. Actinomyces sp. Brans, Centralbl. Bakt. 26: 11-15, 1899. Streptothrix flava Chester, Man. Det. Bact. 362, 1901. Nocardia Brum Chalmers & Christopherson, Ann. Trop. Med. Parasitol. 10: 256, 1916. Actinomyces Bruni Bnimpt, Precis Parasitol. ed. 4, 1204, 1927. Isolated from pus from a case of actinomycosis of the abdominal wall. Nonpathogenic to laboratory animals. Hyphae l-3ju, in diameter, up to 100/a or more long, with swollen tips. Gram-positive, aero-anaerobic ; optimum temperature 35°-38° C, no growth below 25° C. Colonies on agar yellowish, surface irregular, subcerebriform, adherent. Very slight development on potato. In broth, fragile, yellowish white scales in sediment, medium clear, no pellicle. Growth much slower under aerobic conditions. Gelatin not liquefied. Actinomyces Spitzi (Lignieres & Spitz) Dodge, n. comb. Streptothrix Spitzi Lignieres & Spitz, Arch, de Parasitol. 7: 428, 1903. Oospora Spitzi Sartory, Champ. Paras. Homme Anim. 775, 776, 1923. Found in mycosis of the upper jaw in oxen in Cordoba Province, Argen- tina. Pathogenic to oxen and sheep ; less so to horse and pig ; not pathogenic to the usual laboratory animals. In pus, small grains, 40-50/t in diameter, composed of young, radiating hyphae. Medium-sized grains 50-100/a in diameter. Hyphae more branched, with pyriform swellings up to 2/x, in diameter and with mucous secretion. The very large grains, up to 2 mm. in diameter, are formed of agglomerations of medium-sized granules and are firm and grayish or yellowish in color. These degenerate with deposit of calcium. The outer layer is composed of clavate forms, 3-4 x 15-20yu,. This organism is a facultative aerobe, with growth at 37° C. only when first isolated. In young cultures, hyphae bacilliform, straight, or slightly curved. Gram-positive. ACTINOMYCETEAE 753 Growth under aerobic conditions as follows: On agar, growth slow. Large colonies, white, opaque, circular, with the center elevated in the shape of a hemisphere and the marginal zone flat. The small colonies are more ir- regular, not adherent but penetrating the substrate, some becoming crateri- form; aerobic subcultures rapidly dying. Growth similar on coagulated serum. Usually no growth on potato, but occasionally small, grayish white colonies of gram-positive filaments. No growth on potato-glycerol. In broth, there ap- pear grayish white, fine grains, slightly adherent to the walls of the tube, and an abundant floceose deposit ; the medium remains clear. In serum broth, growth better but otherwise the same. No growth on gelatin at 22° C. At 37° C, groAvth as in broth. In hay infusion, growth as in broth. Milk is coagulated slowly, with the whey becoming acid. Curd not digested. Neither coagulated serum nor gelatin liquefied. No indol formation. Anaerobic cultures show some variations. On agar, colonies round, ele- vated, shining, moist, whitish, 2-3 mm. in diameter, not adlierent, no efflores- cence, odor of hydrogen sulphide. No growth on potato. Growth on gelatin and broth as in aerobic, except that with the latter there is also produced the odor of hydrogen sulphide. Actinomyces cruoris (Macfie & Ingram) Brumpt, Precis Parasitol. ed. 4, 1195, 1927. Nocardia cruoris Macfie & Ingram, Ann. Trop. Med. Parasitol. 15: 283-285, 1921. Oospora cruoris Sartory, Champ. Paras. Homme Anim. 809, 1923. Isolated from the heart blood of a patient, both of whose lungs were af- fected with a bronchopneumonia and whose case had otherwise presented the clinical picture of encephalitis lethargica. Necropsy findings given in detail. Pathogenicity not proved. Guinea pigs not affected after 7 months. Hyphae freely branching, nonseptate, Ifx or less in diameter, in old cul- tures fragmenting with ends of branches clavate. No growth on glucose or maltose agar. Growth on blood agar or ''nas agar" at 26° or 37° C, slowly spreading, smooth and dome-shaped, later opaque and puckered in the middle, radially striate and semitransparent at the periph- ery. In early cultures, the efflorescence is abundant and blue. In later sub- cultures, it is scanty and gray or white, medium not pigmented. Gro\vth slow and feeble under anaerobic conditions. Cultures have no distinctive odor. On potato, growth is abundant, whitish, developing a brown or gray brown efflores- cence and staining medium dark brown. Growth also good on coagulated blood serum with substrate also being pigmented dark broAvn. No growth on gel- atin, broth, or peptone water. No change in arabinose, rhamnose (isodulcite), galactose, glucose, fructose, mannose, lactose, maltose, sucrose, raffinose. starch, dextrin, glycogen, inulin, amygdalin, helicin, phlorhizin, salicin, glyc- erol, erythritol, adonitol. dulcitol, inosite, mannitol. sorbitol. Litmus milk un- changed. Coagulated serum not liquefied. 754 MEDICAL MYCOLOGY Actinomyces africanus (Pijper & PuUinger) Naimizzi, Tratt. Micopat. Umana [Pollacci] 4: 8, 1934. Nocardia africana Pijper & Pullinger, Jour. Trop. Med. Hyg. 30: 153-156, 2 ph., 1927. Isolated in a case of mycetoma pedis involving bones. Grains carmine red, 1/4 to 1/^ mm. [micron in original description. Error?] in diameter. No clubs, no sheaths. Foot was amputated. Pathogenic to guinea pig. This organism appears to be a strict aerobe, is gram-positive, but not acid-fast. No odor. No growth at 21° C. No growth on Sabouraud agar, broth agar, potato, glycerol potato, coagu- lated serum, gelatin, glycerol broth, carrot sugar broth, hay infusion. Or- ganism is hemolytic, growing on blood agar with the formation of red colo- nies. In milk, bright red grains similar to those in lesions and soft curd formed. No change in reaction of the milk. Some colonies 2-3 mm. iu diam- eter. When Avhej' is separated from curd, organism grows with white colonies in the whey and red on tlie clot. There is good growth, bright orange in color, on transverse sections of maize cobs. Appearance on solid egg medium is the same. No growth on tyrosine-free media, the red pigment being depend- ent on the presence of tyrosine in acid solution, optimum pH being 5. No de- colorization of pigment once formed nor transition to melanin. The color may be partiallj^ extracted with ether, leaving colorless colonies which again be- come red on the addition of acid. Actinomyces Berestneflfi (Chalmers & Christopherson) Sartory & Bailly, Mycoses pulmonaires 256, 1923. Nocardia Berestneffi Chalmers & Christopherson, Ann. Trop. Med. Para- sitol. 10: 1916. Streptothrix cas I, II Berestnev, Inaug. Diss. Moskva, 1897. Organism is both aerobic and anaerobic, grows on serum, not on gelatin or potato. Actinomyces Ponceti Brumpt, Precis Parasitol. ed. 4, 1205, 1927. Nocardia Ponceti Verdun, Precis Parasitol. 1912 ; Castellani & Chalmers, Man. Trop. Med. ed. 3, 1060, 1919. Discomyces Ponceti Neveu-Lemaire, Precis Parasitol. Hum. 43, 1921. Oospora Po7iceti Sartory, Champ. Paras. Homme Anim. 778. 1923. Said to be a synonym of A. Krausei Chester (1901) by Chalmers & Archi- bald, N. Orleans Med. Surg. Jour. 70: 465, 1918; recognized by Froilano de Mello & Pais, Arq. Hig. Pat. Exot. 6: 133-206, 1918, and by Castellani & Chal- mers, Man. Trop. Med. ed. 3, 1060, 1919. Isolated by Morhof , Door & Poncet from a muscular pseudo-actinomycosis. Bacilliform elements appear on serum in 24 hours. No growth on agar, gelatin, carrot, potato, or oatmeal. Broth turbid with fine flakes at the bottom of the tube. ACTINOMYCETEAE 755 Actinomyces buccalis (Roger, Boiy & Sartory) A. iSartory & A. Bailly, Mycoses pulmonaires 256, 1923. Oospora huccalis Roger, Bory & Sartory, Bull. Mem. Soc. Med. Hop. Paris 27: 319, 326, 1909; C. R. Soc. Biol. 66: 301-303, 1909. Discomyces huccalis Brumpt, Precis Parasitol. 1910, not Streptothrix huc- calis Goadby, 1903. Nocardia huccalis Castellani & Chalmers, Man. Trop. Med. ed. 2, 819, 1913. Found in a case of creamy stomatitis with tonsillar abscess. Caused gen- eralized pseudotuberculosis in guinea pigs. Hyphae 0.7-0. 8/a in diameter, branching, forming irregular spores in long undulating chains. Spores at first barrel-shaped, then ovoid. Growth good on maltose broth, or broth with glycerol or glucose. On carrot, small white protuberances form, 0.8-2 mm. in diameter. No growth on potato or Jerusalem artichoke. Growth very poor on gelatin or maltose agar, with small, punctiform colonies, 0.5 mm. in diameter, slightly depressed in the center. Perhaps the following unnamed species is closely related. Actinomyces sp. A. Sartory & R. Sartory, Bull. Acad. Med. Ill, 94: 893, 894, 1925. Isolated from patient with bronchitis. Pathogenic to rabbit and guinea pig. Hyphae 0.4-0.5/x in diameter, variable in length, some branches coiling about three turns, other branches giving rise to chains of arthrospores, 0.5- 0.6/i. in diameter. No growth on potato, Jerusalem artichoke, agar, or gelatin. Good growth on carrot and banana. Further publication promised. Actinomyces Krausei (Chester) Brumpt, Precis Parasitol. ed. 4, 1204, 1927. Streptothrix sp. Krause, Centralbl. Bact. 26: 209, 1899. Streptotkrix Krausei Chester, Man. Det. Bact. 364, 1901. Nocardia Krausei Chalmers & Christopherson, Ann. Trop. Med. Parasitol. 10: 263, 1916. Isolated from the pus of an abscess on the lower jaw. Pathogenic to guinea pig, rabbit, and mouse. Hyphae composed of long and short cells suggesting Corynehacterium. Organism and aero-anaerobe, gram-positive. Optimum temperature 37° C, no growth at 22° C. On glycerol agar, light yellowish colonies, 2-3 mm. in diameter, with deep, toothed margins; slow in growth, adherent, not confluent. In broth, growth rapid, no pellicle at the surface, no turbidity, spherical colonies at bottom. Claviform formations on serum. No growth on gelatin or potato. No fermen- tation. No formation of hydrogen sulphide or indol. Actinomyces Pijperi (Castellani & Chalmers) A. Sartory & A. Bailly, Mycoses pulmonaires 256, 1923. k 756 MEDICAL MYCOLOGY Nocardia sp. Pijper, Folia Microbiol. 5: 50-53, 1 pi., 1917; Med. Jour. S. Africa 12: 141, 142, 1917. Nocardia Pijperi Castellani & Chalmers, Man. Trop. Med. ed. 3, 1060, 1919. Discomyces Pijperi Neveu-Lemaire, Precis Parasitol. Hum. 44, 1921. Oospora sp. Sartory, Champ. Paras. Homme Anim. 811, 1923. Pound in the mucopurulent sputum of a patient with chronic bronchitis that had lasted for eleven years. Pathogenic to guinea pigs on intraperitoneal injection. Hyphae 0.2-0.3/^ in diameter, nonseptate, ends swollen, often containing granules; gram-positive, not acid-fast. On agar, small whitish colonies which increase in size and become hard, cartilaginous crusts, yellowish, and adherent. Surface corrugated. Occasion- ally a small whitish growth on potato. On serum and serum agar, growth same as on agar. No growth on gelatin at 22° C. Litmus milk discolored. Broth clear, only bottom growth. Anaerobic cultures never reach a great size. Actinomyces Sartoryi Dodge, n. sp. Oospora pulmonalis var. acido-resistant Sartory, Arch. Med. Pharm. Milit. 70: 605-609, 1916. Isolated from a patient showing symptoms of pulmonary tuberculosis. Hyphae straight or slightly curved, not tortuous, little branched, 0.4-0. 5/x X 1.5 mm., no spirals seen. Chlamydospores and arthrospores rare except on potato glycerol. Conidia rare. No growth on carrot, potato, acid potato, banana, starch gel, ordinary agar, fruit decoctions with agar on gelatin, turnip, artichoke, or Kaulin's gel- atin. Growth slow on potato glycerol, colonies 2 mm. in diameter, with a eon- tour irregular and festooned. Surface folded, yelloAvish white. On Sabou- raud agar, growth a little better, colonies 3-4 mm. in diameter, otherwise the same as on potato glycerol. In broth with maltose or glycerol ; long, little branched filaments appear in 48 hours. In liquid serum no growth. Maltose and lactose fermented. Actinomyces transvalensis (Pijper & Pullinger) Nannizzi, Tratt. Micopat. Umana [Pollacci] 4: 46, 1934. Nocardia transvalensis Pijper & Pullinger, Jour. Trop. Med. Hyg. 30: 153- 156, 2 ph., 1927. Found in mycetoma pedis in Pretoria, S. Africa. Lesions produced oily pus with whitish grains, 0.5 mm. in diameter, Avhich showed club forms. Path- ogenic to guinea pigs, grains with clubs being reproduced. Hyphae 0.5 fx in diameter, beaded, breaking up into cocciform and bacilli- form fragments. Organism a strict aerobe, growing very slowly at 22° C, very well at 37° C. Gram-positive, not acid-fast. In general, colonies circumscribed, adherent, with a white chalky efflores- cence. Old cultures rarely show a yellowish tinge. Diastase present. No growth on gelatin, blood serum, Sabouraud agar, broth ; very little growth on I ACTINOMYCETEAE 757 glycerol broth. On ordinary agar, blood agar, or potato, a thin, white chalky growth appears. On glycerol agar or glycerol potato, growth rich, white, chalky, becoming yellowish, potato turning a dirty gray. White chalky growth on carrot. On glucose or maltose broth, there is a lacy white pellicle. Hay in- fusion is the best liquid medium. There forms a thin gray pellicle composed of small grains, 360/i. in diameter, showing a radial structure but no clubs. No acid or fermentation with any of the usual carbohydrates. Litmus milk shows good surface growth but no clotting or digestion. Scarcely any growth on whey. Actinomyces fuscus (Karwacki) Sartory & Bailly, Mycoses Pulmonaires 256, 1923. Streptothrix fusca Karwacki, C. R. Soc. Biol. 66: 180, 181, 1911, non Corda. Nocardia fusca Castellani & Chalmers, Man. Trop. Med. ed. 2, 818, 1913. Oospora fusca Sartory, Champ. Paras. Homme Anim. 809, 1923. Found in sputum of a tuberculous patient. Brownish yellow pellicle on liquid media, browning in time, pigment dif- fusing into medium, making it red brown, then dark brown. Otherwise very close to A. pulmonalis Roger [which Karwacki characterizes as follows: opti- mum temperature 32°-37° C. Deposit of granules in liquid media. No growth on potato and glycerol, coagulated serum and glycerol, or on gelatin. Glycerol agar colonies isolated, circular, white, slightly domed, center adherent. Not pathogenic to laboratory animals. Gram-positive, not acid-fast]. Actinomyces valvulae (Froilano de Mello) Nannizzi, Tratt. Micopat. Umana [Pollacci] 4: 51, 1934. Streptothrix valvulas destruens hovis Luginger, Monatsh. Prakt. Thierheilk. 15: 289-336, 6 figs., 1904. Nocardia valvulae Froilano de Mello & Pais, Arq. Hig. Pat. Exot. 6: 194, 1918. Oospora valvulas destruens hovis Sartory, Champ. Paras. Homme Anim. 788, 1923. Found in endocarditis of cattle. Pathogenic to laboratory animals. Organism a facultative aerobe but grows in depth of agar stab where sur- face has melted, sealing the stab. Gram-positive. Colonies on peptone medium hemispheric, small, later becoming more flat- tened with center elevated and a thin, smooth margin. Growth on milk gray to white. Along agar stab there are clavate projections into the medium, giving an effect suggestive of a row of poplar trees and their reflections in a stream. In broth, there is a slight tubidity, with a mealy sediment, later becoming flocculent. A pellicle is formed. No growth on potato, gelatin, or coagulated ox serum. Actinomyces bronchialis Sartory, Bull. Sci. Pharm. 23: 12-19, 1916. Discomyces hronchialis Neveu-Lemaire, Precis Parasitol. Hum. 43, 1921. Oospora hronchialis Sartory, Champ. Paras. Homme Anim. 802-805, 1923. Found in the sputum of a patient suffering with cough, loss of weight, putrid breath. Pathogenic to guinea pigs and rabbits. 758 MEDICAL MYCOLOGY Hyphae 0.4-0.5/i, iu diameter and up to 2 mm. Jong, regularly branched. Arthrospores and chlamydospores formed. Gram-positive. No growth on the usual bacteriologic media, either solid or liquid, except maltose broth, peptoglycerol, glucose, maltose and Sabouraud maltose agar, and malt extract. Slight growth on maltose broth. Colonies on Sabouraud (maltose) agar Avhite, shining, with irregular margins, 1 mm. iu diameter, in 6 days. In 2 weeks, slightly mammillate and convex, white, finally becoming cream color. Slight decolorization of neutral red. Slight fermentation of glucose, lactose, and maltose. Actinomyces pulmonalis (Roger, Sartory & Bory) A. Sartory & A. Bailly, Mycoses Pulmonaires 256, 1923. Oospora pulmonalis Roger, Sartory & Bory, C. R. Soc. Biol. 66: 150-152, 1909. Discomyces pulmonalis Brumpt, Precis Parasitol. 1910; Froilano de Mello & Pais, Arq. Hig. Pat. Exot. 6: 186, 1918. Nocardia pulmonalis Castellani & Chalmers, Man. Trop. Med. ed. 2, 817, 1913. Found in the whitish grains in the expectoration of a patient with pul- monary mycosis. Also found in otitis. Pathogenic to rabbit and guinea pig, intrapulmonary inoculations producing purulent pleurisy, other inoculations producing local and metastatic abscesses. Hyphae 0.4-0.5 x 1,500/i, branching with terminal cells, claviform. Spores in chains of 8-10, ovoid or spherical, chlamydospores present. Organism a strict aerobe, growing best at 33°-35° C, not at all at 22° C. Gram-positive, not acid-fast. Slight development on simple agar. Colonies round, white, odorless on maltose agar. In maltose broth, white flocci at the bottom of the tube. No fermentation. Actinomyces equi (Chalmers & Christopherson) A, Sartory & A. Bailly, Mycoses pulmonaires 256, 1923; Sartory, Champ. Paras. Homme Anim. 753, 1923. Streptothrix sp. Dean, Trans. Path. Soc. London 51: 26-46, 1900. Nocardia equi Chalmers & Christopherson, Ann. Trop. Med. Parasitol. 10: 263, 1916. Oospora sp. Sartory, Champ. Paras. Homme Anim. 822, 1923. Found in nodule on jaw of horse with pale yellow pus, no granules. Pathogenic to laboratory animals. Hyphae 3-5 x 0.3-0.4/a, curved, club-shaped. Cells occasionally, suggest- ing Corynebacterium, ovoid or ellipsoid cells 0.8-1/* in diameter (chlamydo- spores?). Hyphae best on ascitic fluid or serum broth. Staining irregular, not acid-fast. Optimum temperature 35° -37° C. Aero-anaerobe. Growth poor on solid media, dying out after a few generations. On agar or glycerol agar, colonies irregular with angular projections at surface and ACTINOMYCETEAE 759 into depths, suggesting coralloid growth. Depth growth greater than sur- face growth, maximum 0.5 cm. in diameter, opaque, white, waxy. Stab growth good on upper two centimeters, growth poorer and colonies smaller below. In broth, growth at bottom, liquid clear, once a mass the size of pea, with cauliflower appearance. Maltose broth is favorable to development. Growth in straw, oats, or hay infusion poor. No growth on gelatin, potato, serum, glucose agar, milk, or egg albumen. Actinomyces Taraxeri-cepapi (Sehottmiiller) Dodge, n. comb. lSt)'cptot]iri.r Taraxeri-cepapi Schottmiiller. Derm. Woch. 58: Erganzungs- heft: 89-103, 1914. Pound in fever simulating rat-bite fever, following bite by Taraxerus cepapi, the African squirrel. In pus, straight or curved cells, branching not seen, very slender. Organ- ism gram-negative. No growth on common media, slight growth in broth, to which 5 c.c. of patient's blood was added. Actinomyces Putorii (Dick & Tunnicliff) Dodge, n. comb. Streptothrix Putorii Dick & Tunnicliff, Jour. Infect. Dis. 23: 183-187, 1 fig., 1918. Isolated from blood in case of fever following bite by weasel. Not patho- genic to laboratorj^ animals. Hyphae slender, curved, branching. Ball and club-shaped swellings. Gram-negative, not acid-fast. A facultative anaerobe, growth better under aerobic conditions. Growth on blood agar or Loeffler's blood serum as discrete, colorless or grayish white, pinpoint colonies with elevated, sharp margins, dull at first then glistening. Medium not changed. Growth in glucose blood agar more profuse, grayish yellow. In ascitic broth, slight, whitish, flocculent growth. Growth also in inulin, salicin, maltose, mannite, sucrose, raffinose, lactose, or glucose broths to which ascitic fluid is added in the proportion of 1 :4. No change in reaction. Only slight growth in plain or dextrose broth or milk; in the former a yellowish white, glistening growth appears. Actinomyces Muris-ratti (Schottmiiller) Nannizzi, Tratt. Micopat. Umana [Pollacci] 4: 51, 1934. Streptothrix Muris-ratti Schottmiiller, Derm. Woch. 58: Erganzungsheft, 77-103, 1914; Blake, Jour. Exp. Med. 23: 39-60, Pis. 8-14, 1916. Nocardia Mnris Froilano de Mello & Pais, Arq. Hig. Pat. Exot. 6: 183, 1918. Isolated from sodoku or rat-bite fever, also in bronchopneumonia of rats. Reported by Tunnicliff as pathogenic to white rats, not to guinea pigs or rab- bits. Reported by Blake (1916) from an old woman with sodoku. He reports it pathogenic to rabbits. Hyphae slender, tangled, branching, curved in waves or straight, staining homogeneously. Spore chains, of varying length and staining capacities, break 760 MEDICAL MYCOLOGY up into rods of varying lengths and curvature. Organism is a facultative anaerobe, shows no capsule. Optimum temperature 37° C, little or no growth at lower temperatures. Reported both gram-positive and negative, not acid- fast. This fungus is difficult to cultivate, giving no growth on potato, plain or glucose agar, ascitic glucose agar, plain or glucose broth, milk or gelatin. On blood agar, colonies fine, grayish white, discrete, elevated, dull becoming glistening. On ascitic agar, growth grayish white. On Loeffler's blood serum, colonies colorless (Schottmiiller) ; colonies whitish, circular, elevated, sharply margined, smooth, glistening, moist, fine (Blake). Similar on human blood agar. Actinomyces londinensis Brumpt, Precis Parasitol. ed. 4, 1204, 1927. mreptothrix hominis II Foulerton, Lancet 1 : 970, 971, 1906 ; Ibid. 1 : 627, 1910. Nocardia londinensis Chalmers & Christopherson, Ann. Trop. Med. Para- sitol. 10: 256, 1916. Found in an abscess of the tongue. Pathogenic to guinea pig and rabbit. Claviform formations in broth and whole tufts of hyphae incrusted. On glycerol agar, slight growth, colonies small and white. No growth on potato. Organism a strict aerobe, not acid-fast. Actinomyces Foulertoni (Chalmers & Christopherson) A. Sartory & A. Bailly, Mycose pulmonaires, 256, 1926. Streptothrix hominis Foulerton & Jones, Trans. Path. Soc. London 53: 56- 127, 1902. Nocardia hominis Castellani & Chalmers, Man. Trop. Med. ed. 2, 819, 1913. Nocardia Foulertoni Chalmers & Christopherson, Ann. Trop. Med. Para- sitol. 10: 256, 1916. Found in abscesses of chest and in sputum. Not pathogenic to rabbits. No claviform formations. Organism a strict aerobe, growing best at 37° C. Gram-positive, not acid-fast. On glycerol agar, colonies small and whitish. No growth on potato. On peptone agar, growth slow, colonies small, whitish, heaped up, dry. In inspis- sated horse serum, growth scanty, colonies sinking into medium. In peptone broth, small, spherical, whitish colonies cohering in flocculent masses. Actinomyces decussatus (Langeron & Chevallier) Brumpt, Precis Para- sitol. 1206, 1927. Discomyces decussatus Langeron & Chevallier, C. R. Soc. Biol. 72: 1030, 1031, 1912. Nocardia decussata Castellani & Chalmers, Man. Trop. Med. ed. 2, 817, 1913. Oospora decussata Sartory, Champ. Paras. Ilomme Anim. 825, 1928. Found in whitish, dry, scaly lesions suggesting pityriasis circinata or marginata. Pathogenicity doubtful. Hyphae 0.3-0.5/^ in diameter, cells 1 x 3-4/x in old hyphae. Arthrospores 1-1.5/x in diameter. ACTINOMYCETEAE 761 Growth slow on all media. Ou ordinary agar, at room temperature, colo- nies milk white, with a prominent button in the center surrounded by a circular moat and four cruciform furrows. No growth on Sabouraud agar. Best growth on beef broth neutralized with calcium carbonate or on peptone agar. Inadequately Described Species Actinomyces Genesii (Froes) Dodge, n. comb. Nocardia Genesii Froes, Do mycetoma pedis no Brasil 50, 1930. Case of Genesio Salles from Bahia. Pathology given in detail by Froes, Ibid. 181-191, also Rev. Sud-Amer. Med. Chirurg. 3: 495-501, 1932. Isolated from red grain mycetoma pedis in Brazil. Author unable to in- fect laboratory animals. Grains 150-300/a in diameter, very hard, red; very abundant in tissues. Gram-positive. Cultures difficult and slow on Sabouraud 's agar and potato, red at first, becoming yellowish orange. No growth on liver infusion, broth, milk, egg albumen, or Besredka medium. Actinomyces putridog'enes (Vezspremi) Nannizzi, Tratt. Micopat. Umana [Pollacci] 4: 41, 1934. Cladothrix putridogenes Vezspremi, Centralbl. Bakt. I, 44: 408-415, 515- 523, 1907; Ibid. 45: 15-33, 1 pi, 1907. Nocardia putridogenes Froilano de Mello & Pais, Arq. Hig. Pat. Exot. 6: 187, 1918. Oospora putridogenes Sartory, Champ. Paras. Homme Anim. 823, 1923. Isolated from greenish pus from gingival ulcers and abscess of the jaw, with periostitis and metastases in lung. Pus contained Bacillus fusiformis and Spirochaeta gracilis as well as Actinomyces putridogenes. Of doubtful pathogenicity. Hyphae curved, of varying lengths up to several hundred microns. Grown at 37° C. ; at first only bacilli appear. After 3 days, hyphal threads irregu- larly swollen. Involution forms in old cultures. Organism an aero-anaerobe. Gram-positive and negative. Streptotlirix alba I Sanfelice, Centralbl. Bakt. 36: 355-367, 1904. Oospora alba I, Sartorj^ Champ. Paras. Homme Anim. 817, 818, 1923. Isolated from the air. Pathogenic to rabbit but not to guinea pig. Strain II is similar, but pathogenic to guinea pig and not to rabbit. Colonies on agar are shining and white at first, later becoming chalky, easily separable from the substrate. On potato, colonies look like lime. Anaeromyces bronchitica Chalmers, Douglas & Thomson, Jour. Trop. Med. & Hyg. 24: 151, 152, Pis. 1, 2, 1921. Cohnistreptothrix bronchitica Verdun & Mandoul, Precis Parasitol. 754. 1924. This genus is intermediate between Mycobacterium and Actinomyces. 762 MEDICAL MYCOLOGY Actinomyces cameli (Mason) A. Sartoiy & A. Bailly, Mycoses Pulmonaires 253, 1923. Streptothrix cameli Mason, Agr. Jour. Egypt. 9: 7-13, 1919 (1920). fOospora cameli Mason in Sartory, Champ. Paras. Homme Anim. 822, 1923. Found in pseudotuberculous lesions of lungs, ganglia, and kidneys of a camel. Actinomyces cati Gasperini, Centralbl. Bakt. 15: 684, 1894. Disconiyces cati Rivolta. Actinomyces Dori (Beurmann & Gougerot) Brumpt, Precis Parasitol. ed. 4, 1206, 1927. " Sporotrichum" Dor, Presse Med. 14: 234, 1906. Sporotrichum Dori Beurmann & Gougerot, Ann. Derm. Syphiligr. IV, 7: 996-998, 1906. Disconiyces Dori Beurmann & Gougerot, Les Nouvelles Mycoses 59, 1909. Rhi7iocladium Dori Neveu-Lemaire, Precis Parasitol. 84, 1921. Oospora Dori Sartory, Champ. Paras. Homme Anim. 770, 771, 1923. Found in subcutaneous abscesses resembling sporotrichosis. Not seriously pathogenic to guinea pig or rabbit. Mycelium 0.5-1/a in diameter, cells 6-8/a long, hyphae dichotomous, spores 1-1.5^ in diameter. In broth, a fine deposit of hyphae. Growth slow but much more rapid after adding a drop of acetic acid. Medium remains clear, cultures early lost. Actinomyces enteritidis (Pottien) Brumpt, Precis Parasitol. ed. 4, 1191, 1927. Streptothrix eyiteriticlis Pottien, 1902. Nocardia enteritidis Castellani & Chalmers, Man. Trop. Med. ed. 2, 819, 1913. Disconiyces enteritidis Verdun & Mandoul. Precis Parasitol. 752, 1924. Aerobe-anaerobe. Brownish colonies on potato. Actinomyces erysipeloides (Neumann & Lehmann) Lachner-Sandoval, Ueber Strahlenpilze 64, 1898. Cladothrix des Erythema migrans Rosenbach, Arch. Klin. Chirurg. 36: 2, 1887. Streptothrix Rosenhachii Kruse, Flugge Mikroorganismen ed. 3, 1896. Oospora erysipeloidis Neumann & Lehmann, Atlas und Grundriss der Bakt. 392, 393, 1896. Disconiyces Bosenhachi Gedoelst, Champ. Paras. Homme 177. 1902. Streptothrix erysipeloides Caminiti, Centralbl. Bakt. I, 44: 198, 1907. Nocardia Bosenhachi Gougerot, Gaz. Hop. 86: 202, 1913; Castellani & Chalmers, Man. Trop. Med. ed. 2, 815, 1913. Found in erysipeloid dermatitis. Mycelium slender with claviform formations, aero-anaerobe. Growth on gelatin white when young, brownish gray when old, no lique- faction. On agar, growth brown. ACTINOMYCETEAE 763 Actinomyces Gruberi Terni, Centralbl. Bakt. 16: 362, 363, 1894. INocardia pluricolor Terni, 1894 [fide Chalmers & Christopherson, Ann. Trop. Med. Parasitol. 10: 268, 1916]. f Actinomyces pluricolor Gasperini, Centralbl. Bakt. 15: 684, 1894. Nocardia gruberi Blanchard in Bouchard, Traite Path. Gen. 2: 855, 1896. Nocardia phiricolor Froilano de Mello & St. Antonio Fernandes. Mem. Asiatic Soc. Bengal 7: 110, 1919. Brillant red on gelatin. Glycerol necessary for pigment production at 20°-30° C, slight metabolism. Pathogenic to guinea pig. Streptothrix hominis IV Foulerton, 1906, 1910. Isolated in a case of renal infection secondary to pulmonary infection. Colonies on agar grayish, gradually darkening. Serum not liquefied. Actinomyces Lasserrei (Verdun) Brumpt, Precis. Parasitol. ed. 4, 1206, 1927. Nocardia sj). Lasserre, Contr. Etude Genre Nocardia These Toulouse, 1904. Nocardia Lasserrei Verdun, Precis. Parasitol. 1912 ; Castellani & Chalmers, Man. Trop. Med. ed. 2, 819, 1913. Discomyces Lasserei Neveu-Lemaire, Precis Parasitol. Hum. 43, 1921. Oospora Lasserei Sartory, Champ. Paras. Homme Anim. 798, 799, 1923. Found in ulcer of the pharynx and of upper lip. Pathogenic for rabbit and guinea pig in intracerebral injection. Hyphae 0.5-0.75/* in diameter. Claviform formations present. Colonies on agar discrete, yellowish, surrounded by a layer of white. On potato, yellow nodules with white snowy efflorescence. Actinomyces luteus Brumpt, Precis Parasitol. ed. 4, 1206, 1927. Nocardia lutea Christopherson & Archibald, Lancet 2: 847, 1918. Found in daeryocystes at Khartoum. Streptotlirix madurae Solari, Semana Med. 24: 573-582, 7 iigs., 1917 (non Vincent). Isolated from Madura foot with small yellow grains superficial in skin and muscle, not attacking bone, anesthetizing. Mycelium septate, without constrictions, brownish drab (similar to fila- ments killed with osmic acid), branched, perpendicular to main hypha. Chlamydospores present, spherical, terminal, often several in a chain. On Sabouraud or Sabouraud glucose agar, colonies black, rounded, ele- vated, smooth, not confluent. Surface not umbilicate. Agar blackened. Actinomyces odorifer (Rullmann) Lachner-Sandoval, Tiber Strahlenpilze 65, 1898. Cladothrix odorifera Rullmann, Chem. Bakt. Unters. . . . Inaug. Diss. Miinehen. 47 p.. 1 ph, 1895. Streptothrix odorifera Foulerton & Jones, Trans. Path. Soc. London 53: 112, 113, 1902. Nocardia odorifera Castellani & Chalmers, Man. Trop. Med. ed. 2, 818, 1913. Oospora odorifera Sartory, Champ. Paras. Homme Anim. 809, 1923. 764 MEDICAL MYCOLOGY Actinomyces Pelletieri (Laveran) Briimpt, Precis Parasitol. ed. 4, 1204, 1927. Micrococcus Pelletieri Laveran, C. K See. Biol. 61: 340, 341, 1906. Oospora Pelletieri Thiroux & Pelletier, Bull. Soc. Path. Exot. 5: 588, 1912. Nocardia Pelletieri Pinoy, Bull. Soc. Path, Exot. 5: 589, 1912 [said by Chalmers & Christopherson, Ann. Trop. Med. Parasitol. 10: 255, 1916, to be a synonym of Nocardia indica (Kanthack) Castellani & Chalmers {A. madurae) incorrect] . Discomyces Pelletieri Neveu-Lemaire, Precis Parasitol. 42, 1921. Found by Laveran in tissue of tumor (not a mycetoma) in a negress in Senegal. Not cultivated. Thiroux & Pelletier report a case of mycetoma in the thorax with evidently some of the lung near the lesion infected and an area in the side of thorax. Usual type of mycetoma with red grains similar to A. madurae, but unusual location. Treated externally with iodine, inter- nally with KI, etc. Native thought he was not getting well fast enough and disappeared, balking knowledge of necropsy or cure. Cells 0.7/A in diameter. On Sabouraud agar, growth slow ; colonies small, cerebriform, coral red, not adhering to the substratum. Mycelium not seen in the tissues, but rather cocciform bodies. Differs from A. madurae in the redder color of colonies, growth only on Sabouraud agar, colonies not penetrating the agar and easily detached from it; colonies mucilaginous instead of scaly and dry, in tissues only in micro- coccus form. — Thiroux & Pelletier. Actinomyces pseudotuberculosae Brumpt, Precis Parasitol. ed. 4, 1206, 1927. Case of Flexner, Jour. Exp. Med. 3: 435-450, PI. 41, 1898. Actinomyces pulmonalis Burnett, Rept. N. Y. State, Vet. Coll. 10: 167, 1909. Actinomyces pseudotuberculosis Keller. Nocardia pseudotuberculosis Froilano de Mello & Femandes, Mem. Asiatic Soc. Bengal 7: 110, 1919. Actinomyces purpureus Cavara, Micosi Occ. 99, 1928. Orloff, Vestnik Ofth. 29: 653, 1912 [rev. Klin. Monatsbl. Augenheilk. 51: 529, 1913]. Isolated from lesions of the cornea. Slight virulence for cornea of rabbit, less for dog. Hyphae branched, sometimes with terminal swellings. Aerobic. When grown on agar or gelatin, this organism imparts purple color to medium. Large spherical colonies in fish broth. When grown on coagulated serum, purple pigment is produced. Same on potato. Actinomyces Rog-ersii Brumpt, Precis Parasitol. ed. 4, 1206, 1927. Nocardia (Cohnistreptothrix) Rogersi Froilano de Mello, A Med. Contemp. 1919. ACTINOMYCETEAE 765 Original reference not seen. Froilano de Mello, in a personal letter dated July 7, 1932, states: "A partially acid-fast Nocardia, which further studies have shown to be a contaminating agent which can easily be taken for B. of Koch when present in short rods." Nocardia rubea Chalmers & Christopherson, Ann. Trop. Med. Parasitol. 10: 271, 1916, published without description or synonymy. Oospora rubra Wilbert, Recueil, Hyg. Med. Vet. Militaire, 1908. Actinomyces Somaliensis Brumpt, Precis Parasitol. ed. 4, 1201, 1927. Indiella Somaliensis Brumpt, Arch, de Parasitol. 10: 555-562, Fig. 12, 1906. Indielopsis Somaliensis Brumpt, Precis Parasitol. ed. 2: 1913. Nocardia Somaliensis Chalmers & Christopherson, Ann. Trop. Med. Para- sitol. 10: 255, 1916. Streptothrix Somaliensis Miescher, Arch. Derm. Syphilis 124: 297, Pis. 20-25, 1917. Discomyces Somaliensis Neveu-Lemaire, Precis Parasitol. Hum. 42, 1921. Parasitic on man in Somaliland (Bouffard). Mycelium very slender when young, 0.5/* in diameter, white with lateral branches with septa rare. Older filaments remain white, but their form is more irregular, often moniliform. Chlamydospores (?) intercalary, 1.5-2.5/i in diameter. Fungus grows radially and the young peripheral parts are separated from the inflamed tissue by a hyaline zone, taking the colors of the background, and probably protoplasmic. In this zone, an Actinomyces is fre- quently found in association. Grains hard, softened with difficulty by the usual reagents. Material cementing hyphae together takes methylene blue as readily as cells, Actinomyces urethritidis (Rocek) Brumpt, Precis Parasitol. ed. 4, 1206, 1207, 1927. Isolated by Rocek in 1920, later by Schwartz & Cancik (1922) from sev- eral cases of prostatitis. Actinomyces zur-Neddeni Namyslowski, Centralbl. Bakt. I 62: 564-568, 1912. Streptothrix sp. zur-Nedden, Klin. Monatsbl. Augenheilk. [N. F. 3:] 45: 152-158, 1 fig., 1907. Isolated from eyelid and tear duct. Not pathogenic to guinea pig or rabbit. Gram-positive. Aerobic. No growth on milk, potato, or gelatin. On agar, growth suggests B. xerosis. Streptothrix sp. Gauducheau, Bull. Soc. Path. Exot. 3: 488-490, 1910. Oospora S2^- Sartory, Champ. Paras. Homme Anim. 807, 808, 1928. Found in greenish yellow grains in pseudotuberculosis which was im- proved by medication with KI. Rods 0.3 X 4-5^. curved slightly. Gram-positive, not acid-fast. Hyphae present. 766 MEDICAL MYCOLOGY Authors unable to cultivate. Discomyces sp. De Senez, Bol. del. Lab. de Bact. de Tucuman (Argentina) 1 : 243-247, 2 figs., 1918. Oospo7-a sp. Sartory, Champ. Paras. Homme Anim. 810, 1923. Original description not seen, Sartory describes this organism as follows : Isolated from a case of sup- purating otitis. Pathogenic to laboratory animals, especially by intraperi- toneal injection. Hyphae short, little branched. Optimum temperature 37° C. Growth good on sugar media. On solid media, colony becomes cafe-au- lait in color, with a white efflorescence. Actinomyces Bolognesii-Chiurcoi (Vuillemin) Dodge, n. comb. Malhranchea Bolognesii-Chiurcoi Vuillemin apud Bolognesi & Chiurco, Archivi di Biol. 1: 13 figs., 1925; Pollacci & Nannizzi, I Miceti Pat. Uomo Anim. 5: No. 46, 5 figs., 1926; Bolognesi & Chiurco, Micosi Chirurgiche, Trat- tato di Micopatologia Umana 2 : 566-574, 10 figs., 1927. Isolated from ulcers on the thorax ; grayish seropurulent discharge. Finally metastasis to the lungs. Pathogenic to white rats and guinea pigs. Mycelium hyaline, branched, 0.7-1.5/* in diameter, septate; fertile hyphae more or less prostrate, branched in compound thyrses, branches arcuate or spiral, producing a few ovoid or cylindric spores, 2.5-5.2/t long, hyaline at first, then reddish. Colonies on Pollacci medium round, planoconvex, floccose velvety, with a prominent mammillate center, snow white, then with a flesh tint. Growth rapid, after 5 months showing irregular elevations and depressions in the center and concentric zones at the margin. On blood agar and Petragnani medium, the colonies are smaller but similar to those on Pollacci medium. Actinomyces sp. Ping-Ting-Huang, Derm. Woch. 97: 1679-1685, 2 figs., 1933. Isolated from actinomycosis dyshidrosiformis in Japan. Dysidrosis of the sago grain type and erosio interdigitalis of the fingers and toes but not keratolysis plantare sulcatum of Acton & McGuire. Not definitely pathogenic for white mouse. Mycelium 0.5/x in diameter, with long spore chains, spores 0.7-1.0/* in diameter. On Sabouraud glucose agar, pinhead colonies with chalky surface, con- fluent into greenish brown colony with white spores, earthy odor. On malt and koji agar, thick brownish colonies with white spore covering. On Petragnani medium, yellowish folded nodules with malachite green margins. On peptone agar, white punctiform colonies with reddish reverse. On potato, thick tumid colonies with granular uneven surface covered by a powder of white spores. Litmus broth becomes blue, milk coagulated in 10 days, hemolysis present. ACTINOMYCETEAE 767 BIBLIOGRAPHY Aars, C. G. 1930. Madura foot; its histology and mycology, Arch. Derm. Syphilol. 21: 571- 574, S figs. Abbott, A. C. & N. Gildersleeve. 1902. On the actinomycosc-like development of some of the acid resisting bacilli, Centralhl. Baht. I, 31: 547-550, 1 pi. Acosta, E. 1894. Nueva propriedad del Cladothrix invulnerabilis, Cron. Med. Quirurg. Edbana 20: 479-481. 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Chemisch- bakteriologische Unter&uchungen von Zwischendeckenfiil- lungen mit besondere Beriicksichtigung von Cladothrix odorifera, Inaug. Diss. Miinchen, 47 pp., J pi. — . 1898-1902. Uber eine aus Sputum isolierte pathogene Streptothrix, Miinchen. Med. Woch. 45: 919-922, 1898. II. [with Fr. Perutz] Hid. 46: 407-409, 1 fig., 1899. III. Jbid. 49: 925, 926, 2 figs., 1902. *Sabrazes, Jean. 1896. Presence d'un Streptothrix dans trois cas de pyorrhee alveolo- dentaire, Bev. Mens. Stomatologic. :269. Sabrazes, Jean & P. R. Joly. 1898. Sur un nouveau Streptothrix frequemment isole du vaccine de genisse, C. B. Soc. Biol. 50: 134, 135. Sabrazes, Jean & P. Riviere. 1894. Sur un Streptothrix rencontre dans un cas d'abc^s du cerveau et d 'infarctus suppure du rein, Presse Med. 2: 302-304. — . 1895. Les parasites du genre Streptothrix dans la pathologic humaine, Semaine Med. 15: 383. Sanfelice, Francesco. 1895. tJber einige Infektionskranklieiten der Haustiere in Sardinien. Zoopathologische Untersuchungen, Zeitschr. Eyg. 20: 1-30, Pis. 1-3. — . 1896. Beitriige zur Kenntnis der Aktinomykose der Leber bei den Rindern, Arch. Wi^ss. PraU. Thierheilk. 22: 153-170, Pis. 2, 3. — . 1904. uber die pathogene Wirkung einiger Streptothrix (Actinomyces) Arten, Centralbl. BaJct. 36: 355-367. — . 1905. Streptothrix-Pseudotuberkulose, Centralbl. Bakt. 38: 30-41, 1 pi. — . 1921. Intorno alle mutazione che presenta Streptothrix acido resistente nell 'organismo animale. Boll. 1st. Sieroterapico Milanese 2: 1-15. — . 1922. Streptothrix Actinomyces cuniculi. Una streptotricea che negli animali de esperimento si presenta con colonic raggiatc, Boll. 1st. Sieroterapico. Milanese 2: 327-333. — . 1933. Ueber einige pathogene Variationen der Streptothrix carnea, Centralbl. Bakt. I, 130: 228-234, 6 figs. 782 MEDICAL MYCOLOGY Sanford, Arthur H. & Thomas B. Magrath. 1922. Etiology and laboratory diagnosis of actinomycosis, Minn. Med. 5: 71-80, 4 jigs. Santillan, Prudencio & Gerardo Palacios. 1927. Discomicosis de la pierna, Bol. iThSt. Clin. Qwirurg. Univ. Buenos Aires 3: 825-82S [Sra Bennion Soc. Argentina Patol. Begional del Norte, Tucumdn 1927]. Sartory, Antoine. 1910. Sur les caracteristiques du genre Oospora et son extension dans I'etat actuel de nos connaissances, Presse Med. 577-579. — . 1910, 1911. Contribution a I'otude de quelques Oospora pathogenes. Bull. Soc. Myc. France 26: 394-403, 1910; 27: 160-171, 1911. — . 1916. fitude d'un Oospora pathogene nouveau "Oospora bronchialis n. sp.," Bull. Sci. Fharmacol. 23: 12-19, 1 pi. — . 1916. fitude d'un oospora acido resistant Oospora pumonalis variete acido-resistant. Arch. Med. Pharm. Milit. 65: 604-609. — . 1920. Oosporoses renales, Bev. Med. de I 'Est. 48: 20-25. Sartory, Antoine, M. Meyer & J. Meyer. 1930. Contribution a 1 'etude des mycoses osseuses; trois cas d 'osteites dues d'une part a 1 'Actinomyces asteroides (Eppinger) et d 'autre part a I'Hemispora stellata, Ann. Inst. Pasteur 44: 298-329, 8 figs. Sartory, Antoine & E. Sartory. 1925. Deux cas d 'oosporoses pulmonaires gurries, provoquees par un champignon du genre "Actinomyces" (Oospora), Bull. Acad. Med. [Paris] III, 94: 893, 894. Sartory, Antoine, E. Sartory & Jacques Meyer. 1930. Contribution a 1 'etude des mycoses osseuses primitives: Un nouveau cas d 'actinoniycose osseuse a grains jaunes sans massues, Bull. Acad. Med. [Paris] III, 104: 243-246. — . 1931. fitude du parasite isole d'un cas d 'actinomycose tibiotarsienne a grains jaunes, C. B. Soc. Biol. 106: 212-214. — . 1931. Une microsiphonee a pigment rouge isolee de glaudes sous-maxilaires, C. B. Soc. Biol. 108: 1186-1188. — . 1931. fitude d'un Actinomyces isole d 'expectorations chez une femme suspecte de tuber- culose pulmonaire, C. B. Soc. Biol. 1U8: 1188-1190. — . 1933. Le cycle evolutif des Actinomyces dans les cultures apres passage a travers I'ultrafiltre de collodion, C. B. Acad. Sci. 197: 1465-1467. Sauvageau, Camille & M. Eadais. 1892. Sur les genres Cladothrix, Streptothrix, Actinomyces et description de deux Streptothrix nouveaux, Ann. Inst. Pasteur 6: 242-273, PI. 9. Schabad, J. A. 1904. Actinomycosis atypica pseudotuberculosa, Zeitschr. Hyg. 47: 41-80, PI S. Scheele & J. Petruschky. 1897. Kulturen und Priiparate einer Menschenpathogenen Strepto- thrixart, Verhandl. Kongr. Inn. Med. 15: 551-553. Schlange, H. 1892. Zur Prognose der Aktinomykose, Arch. Klin. Chirurg. 44: 863-875. Schlegel, M. 1913. Aktinomykose, Handb. Path. Mikroorg. [Kolle und Wassermann] 5: 301- 364, illus. — . 1927. Strahlenpilzkrankheit. Aktinomykose, Eandh. Path. Mikroorg. [Kolle & Wasser- mann, ed. 3] 5: 41-112, 20 figs. Schmorl, Georg. 1891. tJber ein pathogenes Fadenbacterium (Streptothrix cuniculi), Deutsch. Zeitschr. Thiermed. Vergleich. Path. 17: 375-408, Pis. 7, 8. Schottmiiller, H. 1914. Zur Atiologie und Klinik der Bisskrankheit (Eatten-Katzen-Eich- hornchen-Bisskrankheit), Derm. Woch. 58: Erganzungsheft :77-103, 2 pis., 4 figs. Scott, W. M. 1922. Bovine actinomycosis, its pathogenesis and treatment by vaccines, Brit. Med. Jour. 2: 1163, 1164. Serra, Giovanni. 1926. Infezioni sperimentali da Actinomyces carneus, Sperim. 70: 251-278, £ pis. *Setti, Carlo. 1929. Discomycosi di Eivolta (actinomycosi di Harz) Studio storico, mor- fologico fisiologico, Giorn. Batter. Immunol. 4: 1-96, 48 figs. ACTINOMYCETEAB 783 Shiota, H. 1909. Beitrag zur Kenntnis der menschlichen Aktinomykose (Klinisclie und bakteriologische Studie), Deutsch. Zeit.tchr. Chirurg. 101: 289-401, 11 figs. Silberschniidt, W. 1899. Sur un nouveau Streptothrix pathogene (Streptothrix caprae), Ann. Inst. Pasteur 13: 841-853. — . 1900. Ueber zwei Falle von Pilzinassen im inneren Thianenkanalchen, Centralbl. Balct. 27: 486-493, 1 pi. —. 1901. Ueber Actinomykose, Zeitschr. Hyg. 37: 345-380, Pis. S, 4. Silva Araujo. 1928. Estudo anatomo-piithologico de um caso de actinoniycose liumana, Sciencia Med. 6: 125-131, 7 figs. Silva, Flaviano. 1929. Contribuicao para o estudo do mycetoma podal na Baliia, Sciencia Med. 7: 153-164, 6 figs. [rev. Brasil Med. 43: 635, 636.] Smith, Theobald. 1918. A pleomorphic bacillus from pneumonic lungs of calves simulating Actinomyces, Jour. Exp. Med. 28: 333-344, Pis. 25-28. — . 1921. The etiological relation of Bacillus actinoides to bronchopneumonia in calves, Jour. Exp. Med. 33: 441-469, Pis. 42-50. Solari, Emilio F. 1917. Primer caso de inicetoma o pie de madura observado en Kosario de Santa Fe, Semana Med. 24: 573-582, 7 figs. *Sommer, Baldomero & Nicolas V. Greco. 1904. Primer caso de micetoma o pie de madura en la republica Argentina, Argentina Med. [Segundo congreso medico latino-amer- icano, Buenos Aires]. Souza-Araujo, H. C. de. 1929. Essais de culture du Mycobacterium leprae (Coccothrix leprae Lutz 1886). Isolement a partir d'un leprome d'un Actinomyces, Actinomyces lepromatis n. sp., C. R. Sac. Biol. 100: 937-939. Speroni, David and Joaquin Llambias. 1930. Actinomicosis generalizada, Bol. Inst. Clin. Quirurg. Univ. Buenos Aires 5: 357-361, 1 fig. [5ta Reunion Soc. Argentina Patol. Regional del Norte, Jujvy, 1929.] Steele, Albert. 1924. A Streptothrix organism from a brain abscess. Jour. Med. Res. 44: 305-310. Sternberg, Carl. 1900. Zur Kenntnis des Aktinomycespilzes, Wien. Klin. Woch. 13: 548-551. — . 1931. Zur Kenntnis der Aktinomykose des Gehirns, Zeitschr. Hyg. 113: 151-159. Steyn, D. G. 1931. Investigations into the cause and transmission of lumpy wool affecting merino sheep and its treatment, Rept. Dir. Vet. Serv. and Animal Industry, Union of South Africa 17: 1:205-213, 6 figs. Stokes, William Koyal. 1904. A study of the group Actinomyces with the report of a pathogenic species for man. Am. Jour. Med. Sci. 128: 861-875, 4 figs. Sutton, Kichard L. 1913. Mycetoma in America, Jour. Amer. Med. Ass. 60: 1339-1342. Talice, Eodolfo V. 1927. Actinomicosis en el Uruguay, Bol. Inst. Clin. Quirurg. Univ. Buenos Aires 3: 819-824, 1 pi. [Sra Reunion Soc. Argentina Patol. Regional del Norte, Tucumdn 1927.] — . 1932. Sur quelques champignons parasites du genre Actinomyces Harz isoles a Montivideo, C. R. Soc. Biol. 109: 144-146. Tarozzi, Giulio. 1909. Ricerche anatomopatologiche, batteriologiche e sperimentali sopra un caso di actinomicosi del piede, Archivio per le Sci. Med. 33: 553-632, 15 figs. [* Thesis Univ. Modena.] Tempsky, Arthur V. 1927. Resultate der Eontgentherapie bei der Strahlenpilzerkrankung, Beitr. Klin. Chirnrg. [Bruns] 139: 207-216. Terni, C. 1894. Eine neue Art von Actinomyces (Actinomyces Gruberi), Centralbl. Bakt. 16: 362, 363. Thiriet, L. 1919. Contribution a 1 'etude des Oospora et des Oosporoses, These Doci. Pharm. Nancy, II, 56: 1-132. — . 1920. Oosporose buccopharyngee. Rev. Med. de I' Est 48: 26-30. Thiroux, A. 1906. Mycose de la jambe .du au Streptothrix Madurae, Ann. Hyg. Med. Colon. 9: 453-455. 784 MEDICAL MYCOLOOY Thiroux, A. & J. Pelletier. 1912. Mycetome a grains rouges de la parol tlioracique. Isolement et culture d'une nouvelle Oospora patliogene, Bull. Soc. Path. Exot. 5: 585-589. Tliiry, Georges. 1897. Contribution a 1 'etude du polychromisme baeterien- — bacille et Clado- thrix polychromes — cristaux colores, Arch. Physiol. Norm. Path. Y, 9: 283-288, 1 fig. — . 1900. Bacille polychrome et Actinomyces mordore. Eecherches biologiques sur les bacteries bleues et violettes. Polychromisme. Corps bacteriens et cristaux colores. Matiere colorante cristallisee, Trav. Lab. Hyg. Inst. Serotherapique Univ. Nancy, 153 pp., 7 pis. Thj0tta, T. & E. Gundersen. 1925. A Streptothrix isolated from the blood in a case of acute rheumatism, with lemarks upon the classification of ray fungi, Jour. Bad. 10: 1-12, 9 figs. Toole, H. 1931. Die Madurafusserkrankung in Grieclienland, Deutsch. Zeitschr. Chirurg. 232: 78-91, 4 figs. Torres, Octavio. 1916. Observacao de um case de actinomycose: 2" case observado na Baliia, Brasil Med. 30: 225-228. — . 1919. Um caso de actinomycose cervieo-facial, Brasil Med. 33: 361-363. TroUdenier. 1903. Ueber eine bei einem Hunde gefundenen Streptothrix, Zeitschr. Tiermed. 7: 81-109, 9 figs. Tsiklinsky. 1903. Sur la flore microbienne thermophilc du canal intestinal de I'homme, Ann. Inst. Pasteur 17: 217-240. Tunnicliff, Euth. 1916. Streptothrix in bronchopneumonia of rats similar to that in rat -bite fever, Jmtr. Infect. Bis. 19: 767-772, S pis. Tunnicliff, Euth and Leila Jackson. 1930. Vibriothrix tonsillaris n. s., the organism of actinomyces-like tonsillar granules, Jour. Infect. Bis. 46: 12-17, Pis. 1, 2. Tusini, Giuseppe. 1900. Ueber die Actinomykose des Fusses, Arcli. Klin. Chirurg. 62: 249- 287, Pis. 3-7. Unna, P. G. 1897. Ueber Aktinomykose und Madurafuss, Miinchen. Med. Woch. 44: 150 [Beutsch. Med. Zeitung 18: 49-51, 1897]. Unna, P. G. & Ernst Delbanco. 1900. Beitrage zur Anatomic des indischen Madurafusses (Mycetoma, Fungus disease of India), Monatsh. PraTct. Berm. 31: 545-567, Pis. 3, 4. Vallee, H. 1903. Sur un Tiouveau Streptothrix (Streptothrix polycliromogene), Ann. Inst. Pasteur 17: 288-292. Vera, Jose E., Donate Vivoli & Eemigio Consiglieri. 1932. Sifilis y actinomicosis pleuro- pulmonar, Meunion Soc. Argentina Pafol. Eegional del Norte en Tucumdn 7: 296- 315, 12 figs. Veszpremi, D. 1907. Zuchtungs- und Tierversuche mit Bacillus fusiformis, Spirochaete gracilis und Cladothrix putridogenes. Beitrage zur Bakteriologie und Histogenese der experimentellen gangranosen Entziindungen, Centralbl. Bakt. I, 44: 408-415, 515-523; 45: 15-33, 1 pi. * Vincent E. and H. Soulie. 1905. Quelques cas de maladies a Streptothrix chez I'homme en Algerie, Bull. Med. d' Alger. Vincent, H. 1894. 'fttude sur le parasite du pied de Madura, Ann. Inst. Pasteur 8: 129-151. Vuillemin, Paul. 1904. Necessite d'instituer un ordre des Siphomycetes et un ordre de Microsiphonees, paralleles a 1 'ordre des Hyphomycetes, C. R. Acad. Sci. 138: 219- 221. — . 1908. Sur I'utilite du groupe des Microsiphonees, C. R. Soc. Biol. 64: 1042, 1043. Waksman, Selman A. 1918. Studies in the metabolism of pathogenic Actinomyces (Strepto- thrices) I, Jour. Infect. Bis. 23: 547-554. Warthin, Aldred Scott & Herbert S. Olney. 1904. Pulmonary streptothricosis, Amer. Jour. Med. Sci. 128: 637-649, 3 figs. Waters, E. 1932. Mycotic dermatitis in sheep, N. Zealand Jour. Sci. Techn. 13: 309, 310, Ifig. AOTINOMYCETEAE 785 Weidman, Fred D. 1928. The affinities between black tongue and trichomycosis, Arch. Derm. Syphilnl. 18: 647-6()5. White, Paul A. 1923. Actinomycosis; diagnosis and treatment, Jo^^r. Iowa State Med. Soc. 13: 105-107. *Wilbert. 1908. Erythasma d'origine parasitaire chez le cheval, Eecueil d'hyg. Med. Vet. Militaire. Williamson, G. A. ^905. Interesting case of mycetoma in Cyprus, Jour. Trop. Med. Hyg. 8: 81. Wissraann, Wilhelm. 1931. Ueber Ergebnisse der Eontgenbestrahlung bei pyogenen Entziin- dungen und der Aktinomykose in der Knpf- und Halsgegend, Inaug. Diss. Med. Falc. Albertus Univ. Kocnigshcrg Pr., 27 pp. Witter, Paul. 1933. Beitrage zur Kenntnis der Aktinomyceten, Inaug. Diss. Math. Naturw. Falc. Univ. Gottingen, 20 pp. Woiil, Michael G. 1923. Fungous diseases of man in the State of Nebraska. Sporotrichosis, blastomycosis, actinomycosis, Trans. Sect. Path. Physiol. Am. Med. Ass. 74: 26-45, 10 figs, [condensed Jour. Am. Med. Ass. 81: 647-653, 10 figs.]. Wolff, Max & James Israel. 1891. Ueber Keinkulturen. des Aktinomyces und seine Ueber- tragbarkeit auf Thiere, Arch. Path. Anat. Physiol. [Virchow] 126: 11-59, Pis. 1-8. Wright, James Homer. 1905. The biology of the organism of actinomycosis, Joiir. Med. Res. 13: 349-404, Pis. 24-33 [reprinted Pull. Massachusetts Gen. Hosp. Boston, 1905]. Zeisler, Joseph. 1906. Notes on a case of Actinomycosis, Trans. Amer. Derm. Ass. 1906: 104-107. *Zschokke. 1888. Doppelfjirbung von Strahlenpilzen, Schwr,is. Arch. Thierheilk. 30: 81. Zur Nedden. 1907. Ueber Infektionen des Auges mit Streptotricheen, Klin. Monatsbl. Augenheilk. N. F. 3: 152-158, 1 fig. I CHAPTER XXI SPOROTRICHEAE Mycelium branched, septate ; spores unicellular, usually solitai-y, never in chains, borne either on very short and little differentiated sterigmata along the hyphae or sessile. Conidiophores wholly undifferentiated. The genera included here are often included in a larger group, the Botrytideae, in which the conidiophore is variously branched. These genera form a transition from the highly developed blastospores and arthrospores of the Eremascaceae Imperfectae to the better known genera of Hyphomycetes in which the conidia are borne on specialized conidiophores. Vuillemin sought to place in a separate group, those species in which there seems to be little or no provision for the separation of the conidium from the conidiophore, naming the group Aleurismeae from the genus AleHrHsma, and calling such spores aleurospores instead of conidia. For the most part, his Aleurismeae have already been discussed in the Gymnoascaceae Imperfectae. There are still a few forms, in which the mycelium is wholly uninucleate, which grow in the horny layer of the epidermis. It is possible that they represent a stage in the life cycle of some of the dermatophytes, but none of the species have been adequately studied. The other genera, Acremonium, Acremoniella, Sporo- trichum, and Trichosporivm, are all found in the subcutaneous tissues or in the internal organs. Most species in these genera are saprophytic on decaying vegetable matter. The whole group badly needs monographic study. Key to Genera Conidia or aleurospores sessile on the mycelium, depending on the death of the mycelium for liberation. Mycelium and spores white or light colored; found in the horny layer of the epidermis. A leurisma. Mycelium and spores black. Trichosporium. Conidia usually on .short sterigmata, not in the horny layer of the epidermis. Conidia terminal on sliort, simple, lateral branches. Conidia hyaline or bright colored. Acremonium. Conidia black or dark brown. Acremoniella. Conidia on long sterigmata, lateral on somewliat branched conidiophores. Bhinotrichum. Conidia on short, simple hyphae or lateral or terminal, on short branches. Sporotrichum. ALEURISMA AleuHsma Link, Mag. Ges. Naturf. Freunde Berlin 3: 18, 19, 1809. Blastomyces Costantin & Rolland, Bull. Soc. Myc. France 4: 153-157, 1888. Aleurophora Magalhaes, Brasil Med. 30: 369, 370, 1916. 786 SPOROTRICHEAE 787 The type species is Aleurisma sporulosum Link. This grenus was first described by Link as follows: 18 Almorisma. Thallus e floccis raniosis, septatis, dcnsissime intertextis. Sporidia in- spersa, minuta, glohosa. Thallus initium Boleti seu Poriae refert at sporidiis creberrimis immixtis genus declarat. E floccis valde contextis, persistentihus constat, ita tit pcmnulwm fere sistat. Sporidia intra thulium conglamerata. Unica species nondurrv descripta. Al. sporulosum, caespitibus indeterminatis crassis deusis albis. Caespitis ad 2-4 linneas latos, ad lin. fere crassos format in ramis dejectis Nudo oculo massa/m farinosam continere videtur. Et e Lusitania habemus. Iconem v. fig. 25. The figure does not help in the interpietation of the above, as it shows only flexuous tangled nonseptate hyphae and a mass of unattached spores. Nees von Esenbeck, Bas System der Pilse und Schioomme 51, 1816, described Aleurisma erubescens as new. Martius, Flora Cryptog. Erlungensis 335, 1817, described Aleurisma granulosum. as new. In 1818, Link reduced Aleurisma to Sporotrichutn and moved part of the original species of Sporotrichum, to Alytosporium. In 183(5, Clievallier in Ins Flore gcncrale des environs d> Farts, ed. l', 1: 51, reestablished Aleurisma, recognizing five species: A. erubescens, A. sporulosum, A. bulborum, A. saccharinum, and A. flavissimiim. Of these, only A. sporulosum was described by Link when he first estab- lished the genus and must be taken as the type. Other authors^ in general, ignored the work of Chevallier until the genus was taken up by Vuillemin in 1911. A careful study of the mechanism of spore development in A. fiavissimum led him to redefine the genus as recog- nized at present. Vuillemin also reduced Blastomyces luteus Costantin & Eolland, the type of Blastomyces, to synonymy with A. flavissirrmm. Hyphae repent, septate, branched ; aleurospores intermediate between chlamydospores and true conidia, not provided with a well-developed mecha- nism for spore discharge, but depending on the disintegration of the support- ing cells in the mycelium ; spores unicellular, fairly thick-walled. Saprophytic, often on dung, very rarely parasitic. Some are inclined to place the greater portion of the dermatophytes here, but it should be noted that the mycelial cells are wholly uninucleate in Aleurisma Avhile they are multinucleate in a relatively large portion of the life cycle in the dermato- phytes. Very few of the pathogenic species referred here have been well described, and still fewer have been carefully studied. It is possible that some of the species reported as growing in the homy layer of the epidermis belong in the dermatophytes. Aleurophora has also been referred here for want of information to place it elsewhere. Key to Species Aleurospores elongate, coremia present. Colonies salmon color. A. salmoneum. Colonies white, elevated. A. dermatitidis. Colonies brown. A. albiciscans. Aleurospores ovoid to spherical. From patclies of scaling witli slight pruritus. A. benigna. From vesico pustules. A. Ivgdunense. A. Vuillemvni. 788 MEDICAL MYCOLOGY Aleurisma salmoneum Vuillemin, C. R. Acad. Sci. 189: 407, 1929. A laborer in wine vaults showed on the anterior face of the right arm a pustule, about 1 cm. in diameter, which had been the seat of a slight but incessant suppuration for about a month. On pressure, lesion exuded a thin pus, whose granules showed neither spores, grains, nor filaments. Cultures salmon color. If inoculation is deep, the surface of the agar is smooth and mammillate; if superficial, the colonies are dull, rough, with a deeper color. Aleurospores terminal, 4 x 2.3 — 10.5 x 4/i.. Aleurisma albiciscans (Nieuwenhuis) Dodge, n. comb. Trichophyton aTbiciscans Nieuwenhuis, Geneesk. Tijdschr. Nederl. Indie 48: 35-51, Pis. 1-3, 1908; Arch. Derm. Syphilis 89: 3-30, Ph. 1-4, 1908. Atricho'phyton alhiciscans Castellani & Chalmers, Man. Trop. Med. ed. 3, 1008, 1919. Glenospora alhiciscans Ota, Ann. Parasitol. Hum. Comp. 3: 79-84, 1 fig., 1925. Glenosporella alhiciscans Nannizzi apud Agostini, Atti 1st. Bot. R. Univ. Pavia, IV, 2: 98, 1930 [1931]. Found in tinea albigena. On soles or palms, little itching nodules appear. These become vesicles 3-4 mm. in diameter, at first filled with an amber colored liquid, then purulent and drying. In time, these vesicles are more frequent and larger, followed by drying and keratinization. The epidermis becomes very thin and easily torn, so that secondary infections frequently occur. Finally, pigment is lost over the infected areas, which have expanded to in- clude the lower arms and legs. This disease is widespread in the interior of Java, Borneo, and Lombok, probably also in New Guinea and Sumatra. Mycelium of young cultures slender, 2/x in diameter, guttulate, cells 15-20/x long, spherical or ovoid or elongate. Occasional lateral spores with or without pedicel. In older cultures, spores more or less ovoid, 1-1. 5/x in diameter, lateral with or without pedicel. No thyrses present — Nieuwenhuis. Hyphae 2-3/u., rarely 3-5/*, in diameter, walls thin, covered with ver\^ fine rugosities. Inter- calary spores present, 7-10/i, in diameter, coremium branching either perpen- dicular or at an acute angle to the hyphae. Conidiophores simple or irregu- larly branched, no verticils present. Aleurospores terminal or lateral, usually sessile on conidiophore, usually solitary, occasionally in groups of two to three, 2-3 X 2-7/i,, thin-walled, not filled with granules. Very rarely there is a sug- gestion of thyrses, but these are not well developed as in Microsporum. Coremia bearing numerous aleurospores and chlamyclospores usually are present. Growth is slow, attaining a diameter of only 10 mm. in 21 days on the first isolation. On Sabouraud agar, there forms an irregularly rounded disc, 4-6 cm. in diameter, after 42 days, clear chamois color in malt agar, coarse velvety with numerous small coremia, 1-3 mm. long in center. As cultures dry, medium becomes homy, white, with aerial mycelium appearing. Develop- ment good on potato. SPOROTRICHEAE 789 Aleurisma dermatitidis (Agostini) Dodge, n. comb. Glenosporella dermatitidis Agostini, Atti 1st. Bot. R. Univ. Pavia, IV, 2: 93-101, 4 figs., 1930 [1931]. [Case of Mantarro?* Giorn. Ital. Derm. Sifilol. 72: 131-11:3, 1931.] Heads sparse, suborbieular, minute or confluent in larger colonies, irregu- larly subrotund and raised above the substrate, white without, yellow within. Hyphae hyaline, densely interwoven, simple or irregularly branched, continu- ous or septate, variously granulose 2-7ja in diameter. Aleurospores hyaline, spherical or subrotund, thick-walled 7-10 x 4-7/x, clinging to the branches of mycelium, acrogenous, solitary or irregularly disposed, sessile or on aleuro- phores of varying length. Arthrospores rounded or oblong or irregular, 4-15ju. in diameter, intercalary, catenulate. Aleurisma benig^a (Magalhaes) Vuillemin, Champ. Paras. Mycoses Homme 114, 1931. Aleurophora henigna Magalhaes, Brasil Med. 30: 369, 370, 1916. Isolated from circular patches of scaling with slight pruritus, infection scarcely noted by patient until attention was called to it by physician. Fungus most abundant in the stratum disjunctum of the epidermis. Mycelium curved, long or short, dichotomous, 2-3^ in diameter. Reproduc- tion mainly by arthrospores (thallospores), 2-3//, in diameter, or by aleurospores. Yeast cells spherical to ovoid, 2-5fx in diameter. Cultures mentioned but not described at all in this place. Genus not described. No figures. Aleurisma lug-dunense Vuillemin apud Massia & Grigorakis, C. R. Soc. Biol. 91: 1381-1383, 1924. Scales of erythremato-squamous lesion with peripheral vesicopustules on internal face of knee where skin was kept moist by prosthetic apparatus. Mycelium yellow, with some racquet cells, aleurospores as in Sporotrichum 3-4 X 2-2. 5/a; chlamydospores 5-6yu,. Aleurospores germinate with 1-3 germ tubes, chlamydospores Avith 1-4. All cells uninucleate except rapidly growing terminal cells, which may have as many as four nuclei. Colonies cafe-au-lait in color, smooth becoming powdery white. Baudet (1930) studied a lesion on a dromedary, in Avhicli the organism was identified by Vuillemin as this species along with Grubyella Langeroni. Baudet describes the cultural characters of his organism as follows: Aleuro- spores 1-2.5 X 0.8-1.5/1, ovoid, borne singly on short branches as in Monosporium. He failed to find the chains of aleurospores reported by IMassia. The chlamydo- spores were mistaken for racquet cells. Produced a diffusing pigment on glu- cose media, rose, then wine red, which disappeared in subcultures, apparently developed on acid media pH 4.4-4.6, while it was greenish yellow on more alkaline media. Reverse honey yellow (melleus of Saccardo Chromotaxia) . There seems to be little to differentiate the following species from this, but both the original descriptions are too poor for certain identification. 790 MEDICAL MYCOLOGY Aleurisma Vuillemini, Grigorakis, Fayet & Magrou, C. R. Soc. Biol. 95: 649-651, 1926. From lesions in dog similar to those caused by Microsporum canis. Aleurospores 2-4.5 x 2.3/*, spherical to ovoid, germinating with 1-3 germ tubes. Whole cycle uninucleate as far as observed. At first smooth, cafe-au-lait, then white. On liquid media, a ring and thick pellicle with white surface and cafe-au-lait color below. On malt gelatin, a red pigment in stripes or patches is produced and medium liquefied, the whole then becoming chocolate colored. Doubtful Position The following incompletely described species probably belong here but may belong in the imperfect Eremascaceae. They do not belong in Glenospora by any definition current in mycology or medical literature. Aleurisma metaeuropeum (Castellani) Dodge, n. comb. Glenospora metaeuropea Castellani, Med. Press Circular 136 : 440, 1933 ; Jour. Trop. Med. Hyg. 36: 309, 310, Figs. 30-33, 1933. Originally isolated from a case of dermal blastomycosis contracted in the Balkans. Later isolated from a woman in Italy following an injection of cam- phorated oil in the buttock, which produced ulcers that extended slowly, causing large cavities with granulomatous margins and finally affecting the sacrum and some vertebrae. Cured by prolonged treatment with massive doses of potassium iodide. Hyphae septate, 2-6//, in diameter, aleurospores terminal, subspheric, 6-10/i,, or ovoid, 8.4 x 3.5/i,. On glucose agar, colonies white and fluffy. No pigmentation of mannitol agar. No fermentation or acid production with carbohydrates, no action on milk. Gelatin liquefied. Aleurisma metamericanum (Castellani) Dodge, n. comb. Glenospora metamericana, Castellani, Jour. Trop. J\Ied. Hyg. 36: 310, 311. Fig. 34, 1933. * ' Blastomycosis. ' ' Hyphae septate, 1.7-2/x intercalary chlamydospores occasional ; lateral aleurospores spherical or ovoid, 3-6// in larger diameter. On glucose agar, growth sometimes moist, center vermiculate with fur- rows. No pigmentation on mannitol agar, gelatin liquefied promptly. Perhaps this species should be placed in Proteomyces, but it is too poorly described to place it definitely. AleurismBi breve (Castellani) Dodge, n. comb. Glenospora hrevis Castellani, Jour. Trop. Med. Hyg. 36: 311, Fig. 3.5, 1933. Isolated from a case of blastomj^cosis in America. Aleurospores lateral, spherical, 3.5-8/i, on short sterigmata. Growth good on ordinary media, no pigmentation on ordinary media, gel- atin not liquefied in the first two weeks. SPOROTRICHEAE 791 TRICHOSPORIUM Trichosporium Fries, Summa Veg. Scand. 492, 1849. This genus was first described by Fries in his Systema Orbis Vegetabilis 306, 1825, without adding species, hence was not valid at that date. He failed to recognize it in the Systema Mycologicivm in 1829, but did in the Summa. Since the description is most com- plete in the Systema Orbis Vegetabilis, it may be considered here. It was defined as Flocci varii, septati, sporidiis e floccis enatis nudis adspersi {Saxicola, truncicola, colore varia). Since it was reported as found on rocks, it is probable that some lichens were also included here. In no place does Fries carefully distinguish it from existing genera. In 1849 he characterized it as Sporae simplices, nee septatae, solitariae, reliqioa prioris Trichothecium. He recognizes twelve species, all previously described, but it is almost impossible to select a type species, as none is described here. Saccardo, Michelia 2: 125, 1880, redefined the group, taking his T. nigricans as the type. In this sense, it has been used as the black-spored analogue of Sporotrichum, or, as defined by Lindau 1906, as the black-spored analogue cf Aleurisma. In this sense, it is equivalent to the use of Glenospora of Vuillemin and later medical authors, not of Berkeley & Curtis which is an imperfect stage of Septobasidium (Couch 1933), a wholly unrelated fungus. Hyphae repent, irregularly branched, brown or pale; conidia (or aleuro- spores) terminal or lateral on the hyphae or ultimate branches spherical or ovoid, smooth or slightly rough, brown or occasionally almost hyaline. The whole group needs monographing before one can be certain of the proper name to apply to these species. Key to Species Gelatin liquefied ; from generalized infection. T. Ga/nimeli. Gelatin not liquefied; from local infections. Aleurospores 3 x 5m; from lesions of cornea. T. graphii. Aleurospores 5-6.5 x 6.5-9fi; from mycetoma with black grains. T. hhartoumensis. Aleurospores, 6-7/j. in diameter; from gummatous nodular lesions in thorax. T. Mantegazzae. Trichosporium Mantegazzae Pollacci, Riv. Biol. 4: 318-328, Pis. 1, 2, 1922; Bolognesi & Chiurco, Micosi Chirurg. 888-895, 1927. Isolated from gummatous nodules in swellings of the thorax. For case history see Mariani, 1923. Pathogenic for laboratory animals. Sterile hyphae repent, branched, subhyaline then fuscous, and finally black, 7-8/n in diameter. Vegetative hyphae moniliform, black. Chlamydo- spores numerous. Conidiophores ascending or repent, septate, simple or branched, roughened, brown, long, slender, bearing numerous lateral conidia which are spherical or ellipsoid, easily separable, more or less in whorls, fuliginous, black, somewhat papillate, 6-7/x in diameter. Optimum tempera- ture 15° -20° C. Colonies small, discrete, grayish, becoming greenish gray and finally- black, confluent, covering the whole surface of the medium. Surface of colony velvety, wrinkled, rugose, mammillate, not dry or shining, adherent. Ring formation in glucose broth, submersed mycelium not blackening, spores 6.5 x 792 MEDICAL MYCOLOGY 11/A. Growth good on blood agar or potato. JSmall colonies on serum, partially liquefying the substrate. Milk coagulated and partially digested. Gelatin not liquefied. Trichosporium Gammeli (Pollacci & Nannizzi) Dodge, n. comb. Glenospora Gammeli Pollacci & Nannizzi in Blankenhorn & Gammel, Jour. Clin. Invest. 4: 471-484, 9 figs., 1927. [Case history by Blankenhorn and Gam- mel, Trans. Assoc. Amer. Physicians 41: 268-282, 9 figs., 1926.] Isolated from a case of generalized mycosis involving lungs, digestive tract (?), and the epidermis, the latter with small furuncles and much necro- sis, the others without either inflammation or necrosis but showing larger and deeper nodules, which soften and ulcerate as in blastomycosis. Symptoms dis- appeared in 61 days on medication with KI and neoarsphenamine. Nt) recur- rence in 2 years. Organism not pathogenic to monkeys, guinea pigs, or rabbits. Sterile liyphae hyaline or subhyaline, often guttulate, continuous when young, becoming septate in age, 2-5. 3/a in diameter. Cells 15-100/t long, mono- podially branched, never dichotomous, larger ones frequently septate, com- posed of clavate cells as in Microsporinn Audoiiini, sometimes fasciculate, here and there anastomosing. Fertile hyphae concolorous, decumbent, a little more slender, with short branches and with aleurospores. Aleurospores sessile on aleurophores of variable length, spherical, smooth, 3.3-6/x. in diameter, at first hyaline and chlorine colored, later 10-12/a in diameter, thick-walled, gran- ulose, pale luteofuscus. Chlamydospores spherical to irregular, 10-12/t in di- ameter, brown. Nodular organs resembling perithecia formed, but develop- ment stops here. Growth strictly aerobic, optimum temperature being 18°- 22° C. Growth on Sabouraud glucose agar a dense, thick, and rather undifferen- tiated felt of delicate hyphae, sometimes with slight concentric zones, white to light brown and yellowish brown, 2.6-3.2 cm. across in 3 weeks. On Sabou- raud maltose agar, colony is white and growth slower. Best growth on Grlitz malt agar, with colony brown in color. Grayish white on peptone agar and white on most other media. Upper portions of carbohydrate media darken slightly. In liquid media, powder-puft'-like growths appear at the bottom and rise to the top, forming a thick pellicle. In milk, only surface growth, with alkaline reaction produced. In 8 weeks, milk finally clears with calcium oxalate and calcium acid phosphate crystals deposited. Solution is deep yel- low brown in color, with light brown growth in upper third and coarse, heavy, flocculent precipitate on the bottom appearing in old cultures. Litmus broth with galactose, glucose, or fructose very slowly decolorized. No fermentation. Gelatin is liquefied in 3-4 weeks, with dense yellow mycelium on surface and fine sediment. Var. lanug^nosus (Castellani) Dodge, n. comb. Blastomycoides lanuginosus Castellani, Brit. Jour. Derm. Syphilis 42: 365- 374, 1930. Glenospora lanuginosa Agostini, Jour. Trop. Med. Hyg. 34: 287, 288, 1931. SPOROTRKIIEAE 793 In tissues, cells l()-20fi in diameter, 1 hick-walled. On PoUacci agar, hyphae hyaline, sinii)le, septate, variously granular, 2-7/a in diameter, forming some racquet mycelium in old cultures. Aleurospores terminal or unbranelied, hyphae oblong or rounded, 7-11 x 3-6/x, apiculate with thick membrane, usually single, rarely in chains of two or three. On Sabouraud's and Raulin's media, arthro- spores up to IS^u, in length are formed. Colonies develop 24-30 hours after inocu- lation. Optimum temperature for growth 25° -30° C. Colonies on Pollacci agar spreading, languinous. rising above the surface 2-3 mm., white, then yellowish or brownisli. No pigment in mannitol agar. In blood agar, colonies punctiform, confluent, slightly lanuginous, light yel- low. No fermentation or acid production in carbohydrates. Gelatin and coagulated serum rapidly liquefied. Ota & Kawatsure (1933) reduce Glenuspum Gainmeli to synonymy with their Aleuiisma tulanense (see p. 841, Monospormm iulanense), but their de- scription of cultures fits Castellani's description of Blastomycoides lamiginofsvs better than that of his Blastomycoides tidanensis, so that one wonders if there has not been a confusion of culture labels. Trichosporium graphii (Harz & Bezold) Dodge, n. comb. AspergilUis glaucus Ilassenstein, Zeitschr. Parasitenk. 1: 111-113, 1869, non aliorum. VeriicilUnm graphii Harz & Bezold apud KSiebenmann, Die Schimmelmy- kosen des Menschlichen Ohres, 95, 1889. Glenospora graphii Vuillemin, C. R. Acad. Sci. 154: 141-143, 1912. Description of Vuillemin was based on a corneal organism isolated by Morax. Conidiophores verticillate, branched or solitary, ending in a single conid- ium ; at 18°-20° C, conidium ovoid, grayish or yellowish, 7.5 x 5.7/x, or sub- pyriform and only 1.4/x in diameter at the disjunctor. Coremia present on poor media, not on rich. On coremia, successive conidial production takes place, but conidia never appear in chains. Conidia may be cylindric, 6.5 x S/jl, spherical, o/j. in diameter, or ovoid, 5 x 3/a. At the optimum temperature, 37° C, spores are slightly smaller. On most media, colonies begin as small grayish tufts, then blacken, with brown mycelium penetrating the medium. No action on milk or gelatin. Trichosporium khartoumensis (Chalmers & Archibald) Dodge, n. comb. Glenospora khartoumensis Chalmers & Archibald, Ann. Trop. Med. Para- sitol. 10: 169-222, Pis. 4-7, 1916. Isolated from a black grained maduromycosis in the Soudan. On microscopic examination, the grains show a light-colored interior with branched, septate hyphae and chlamydospores. Hyphal cells 2.1-2.3 x 5.6-8/i,; chlamj^dospores 7 x 15.3 x 14/*. In cultures, aleurospores appear on branched hyphae. No sexual reproduction observed. Optimum temperature 30° C. On maltose agar, colony shows a central elevation surrounded by a groove separating it from a plateau which is grayish at first becoming darker (dusky 794 MEDICAL MYCOLOGY drab, Ridgway). On glucose peptone medium, hyphae are white with a dark, reddish-brown pigment diffusing into the medium. Milk neither acidified nor coagulated. Gelatin and serum not liquefied. There seems little to differentiate the followng species from this but, since it is so much more fully described, 1 hesitate to reduce it to synonymy without further evidence. Trichosporium Clapieri (Catanei) Dodge, n. comb. Glenospora Clapieri Catanei in Montpellier & Catanei, Arch. Inst. Pasteur Algerie 5: 489-508, 1927; Montpellier, Catanei & Clapier, Bull. Soc. Path. Exot. 20: 502-511, 1927. Isolated from a black grain tumor of the maxillary region. The grain in section suggested Madurella in appearance. Mycelium septate, 2.5-3//, in diameter, brownish with occasional coremium formation, aleurospores terminal or lateral and sessile, on branches, smooth, ovoid, truncate, 6.5-9 x 5-6. 5/a. Chlamydospores 10.5-15 x 9-15/^, intercalary or terminal, brownish black. Growth best at 37° C. On Sabouraud glucose agar, colony black, slightly shining, with little pointed black coremia, 1-2 mm. tall, finally covered with a gray velvet, some- times rough, with a black border, center elevated, colony deep in the gel and adherent. On ordinary agar, colony is deep brown, almost black, covered with coremia, 0.5 mm. tall, whole colony becoming 2-2.5 mm. thick with the medium brownish around the colony. On potato, colony deep gray with a velvet finer than that on Sabouraud glucose agar. On ageing it is slightly pulverulent, brown, with the color of the substrate not modified. On potato glycerol, sur- face is black, shining, irregular. On carrot, growth is about the same as on potato. On coagulated serum, colony small, elevated. On gelatin, both super- ficial and deep colonies. In peptone water, there appear small tufts, gray in color at first, then deepening, small ring, no turbidity. Milk is not coagulated, but turns alkaline after one week and continues to turn. Neither coagulated serum nor gelatin liquefied. Doubtful Position Glenospora g-andavensis Vuillemin, C. R. Acad. Sci. 173: 378, 1921. Isolated from sputum, Ghent, Belgium. Chlamydospores 6-10 x 5-8/a; hyphal cells 8-13 x 1.6-3.5;u,, branching in all directions. Hyphae hyaline, without chlamydospores in Veillon agar, in low oxygen tension. Growth good at 20°-37°, optimum 32°-35° C. On Veillon stab, surface growth only. On maltose agar, a central black dome with clear fringes. Growth on carrot blackens on the fourth day, shows chlamydospores on the fifth. Glenospora Semoni Chalmers & Archibald, New Orleans Med. & Surg. Jour. 70: 455-475, 1917 (nomen nudum) ; Semon, Brit. Jour. Derm. Syphilis 27: 299-303, PI. 8, 1915 (case history). SPOROTRICHEAE 795 This name is used for the organism in cases of maduromycosis with red grains first clinically described by Carter in India and first cultivated by Semon from the foot of an Indian soldier in the British army in France. Mycelium dichotomously branched with a "stippled appearance." No chlamydospores found. On solid media, a central black zone with gray periphery which becomes black in 10 days. In Kaulin's liquid, at the optimum temperature, 35° C, a delicate, translucent, grayish, feathery growth appears. Perhaps the following unnamed species may belong here, or it may be re- lated to Chalara. Oospora sp. Sartory, C. R. Soc. Biol. 84: 939-941, 1921. Mijcoderma sp. Sartory & A. Bailly, Mycoses pulmonaires 324, 1923. Isolated from sputum ; patient improved on medication with KI. Not pathogenic for laboratory- animals. Mycelium white, becoming brown or chocolate color, 4:fi in diameter, sep- tate, sinuous branched, walls slightly verrucose, conidiophores rigid, spores in sinuous chains, yellow and brownish, ellipsoid 5.5-8.8/^, with granular contents. On agar, colonies convoluted, irregular, powdery, white, browning after a week. On potato, brown, then mauve violet and reddish brown. On gelatin, irregTilar white colonies, brow]i, powdery, mammillate, with white pellicle. No growth on seiiim. Milk coagulated in 10 days, casein precipitated and digested. No indol. No action on starch. Sucrose hydrolyzed in fermentation of sugars. Gelatin liquefied. ACREMONIUM Acremonium Link, Mag. Ges. Naturf. Freunde Berlin 3: 15, 1809. The type species is Acremonium verticillatum Link. Hyphae branched, septate, repent; conidiophores simple, scarcely more than short lateral branches, more or less erect bearing single solitary terminal conidia which are hyaline or light colored, mostly ovoid and small (Fig. 119). This group has been isolated from deep-seated abscesses and from gener- alized infection resembling sporotrichosis, so far reported only from the Ivory Coast and once from France. Acremonium Muthuoni Fontoynont & Boucher, Ann. Derm. Syphiligr. VI, 4: 339-344, Figs. 9, 10, 1923. Found in abscess on the neck and arm which caused intense pruritus Scratching caused flow of blood but no pus. After various antiseptics had failed, the abscesses were excised. Mildly pathogenic to rat and guinea pig. Hyphae 1/i, in diameter. Conidiophores short, straight, or curved, 5-20/j in diameter, swollen in the middle. Conidia 2.5-3/t in diameter, elongate el- lipsoid. 796 MEDICAL MYCOLOGY Oil glucose agar, colonies grayish white, dry, powdex-y, reverse, slightly rose colored; agar turning brown. After 75 days, the reverse becomes brick red. On "poor" agar, growth much the same but slower and the brown deeper. Growth on maltose agar, creamy, gray, folded, suggesting the ex- creta of earthworms, tinted brick color. On sweet potato, colony is a white velvet finally becoming powdery; medium turns greenish. On potato glycerol, colony is of much the same aspect as the substrate and is, therefore, difficult to see. On glucose gelatin, growth is at first much as on glucose agar, then villous and shining with liquefaction on the eleventh day. In glucose broth, medium turbid, with floating white colonies which form a ring and tend to form a pellicle. Acremonium niveum Boucher, Bull. Soc. Path. Exot. 11: 324-327, 1918. Isolated from lesions on hands and feet. Two cases reported cured by medication with potassium iodide. Fatal to guinea pigs. Fig. 119. — Acremonium alternatum Link. (After Saccardo. ) Mycelium septate, 2fi in diameter, hymenium formed of conidiophores, branching at right angles with swelling in the median portion, each branch being terminated by a long, pointed extremity bearing a single fusiform spore, 4-5 X 2-2.5/A. Conidiophores 20-22/x long, sometimes united in a coremium. On ordinary glucose agar, growth slow, Avliite then rose color, slightly velvety, becoming dry and wrinkled. On Sabouraud conservation agar, colony white, velvety, dry. On potato glycerol, colony snow white, depressed. On sweet potato, colony velvety, white becoming rose color, with a snow white pellicle covering the water in the bottom of the tube and extending about 5 mm. up the side of the tube. In giant colony, center wrinkled, velvety, white, radiately striate, margins rose and velvety. In glucose broth, white flakes float in all depths, forming a discontinuous pellicle, finally settling out, with hyphae growing up the glass for 6 mm. SPOROTRICHEAE 797 Acremonium Potronii Vuillemin, Bull. Soc. ISci. Nat. Nancy III, 1: 144- 148 (15-19), PI. 2, 1910. [First case reported by Potron & Noisette, Rev. Med. del'Est43: 132-139, 1911.] Isolated from a generalized infection resembling sporotrichosis followed by hydrarthrosis. This was the only organism found. Medication with potassium iodide successful. Not pathogenic to guinea pigs or rabbits. Hyphae branched, septate, scarcelj^ l/x in diameter, sometimes in fascicles. Conidiophores simple, usually divaricate, rarely once branched, phialiform but not septate at the base, ultimate branches, 15-20^i long, inflated below (up to 1.75/a), with a subequal neck (O.S/i,). Conidia successively formed and expelled, rose, ovoid, smooth, with a short appendiculate base 4-5 x 2.2/a (Pig. 120). Optimum temperature 37°, scarcely any growth at 10° C. I Fig. 120. — Acremonium Potronii Vuillemin. (After Vuillemin 1910.) On agar, colony grayish white, smooth, covering the surface of the medium. On glucose gelatin, small white tufts, 2-4 mm. in diameter, later becoming confluent and rose color, adherent to the medium. On potato, colony finely granular, later becoming rose color. On carrot, finely radiate colonies in tufts, 3-4 mm. in diameter, white. As the tufts grow and become confluent, the colony becomes coralloid and rose color. On coagulated egg albumen, a white pellicle, becoming transparent and yellowish as the medium liquefies. On peptone broth, tufts of long, tangled hyphae on the surface and in the liquid which otherwise remains clear. In acid broth, growth poor; better in alkaline broth. In broth or peptone solution, to which nitrate has been added, pellicle white with white flocci in liquid. No indol formation. Milk coagu- lated and liquefied, yellowing, colonies white. 798 MEDICAL MYCOLOGY ACREMONIELLA Acremoniella Saccardo, Miehelia 1: 270, 1878. The type species is Acremonium airum Corda. Hyphae repent, simple or branched, hyaline or colored ; conidiophores simple, short, bearing solitary terminal eonidia which are spherical or ovoid, brown, unicellular. This genus is predominantly saprophytic, but it has been isolated twice from lesions, once from a small tumor and once from the lungs. It occasion- ally is found as a contaminant. Acremoniella Berti Pollacci, Atti 1st. Bot. R. Univ. Pa via 18: 126, 127, PI. 30, 1921. (Case histoiy and cultural characters by Berti, Policlinico Sez Chirurg. 29: 484-489, Figs. 6-9, 1922.) Isolated from a small tumor. Granuloma produced in a guinea pig. Not obtained pure ; sterile hyphae repent, branched, septate, hyaline, sparse, 3-4/* in diameter; conidiophores hyaline, erect or curved, not cuspidate, 15-25|U. long ; eonidia spherical, unicellular, 6-7/t in diameter, brown. Growing with Penicillium Burci Pollacci. Grows on Sabouraud glucose ; figures very poor. Acremoniella Perinii Pollacci, Rev. Biol. 5: 358-367, 3 figs., 1923. Found in sputum from a patient with pulmonary lesion. Pathogenic to guinea pig. Tufts at first white then fuscous, diffuse ; sterile hyphae repent, intricate, septate, hyaline or pale ; conidiophores erect, simple, short, 3.5-4/i, in diameter, 16-24/x long, septate or continuous, pale, apex obtuse or often inflated. Conidia spherical, granulose, hyaline at first, becoming avellaneous, echinulate, unicellu- lar, acrogenous, 7.8-9.7/i, in diameter. Colony grayish at first, then brown. Growth on potato, silky white, with small islands of green, then the surface becomes brown. No growth at 20° C, good at 37° C. At 45° C, a pellucid colony, white, rapidly turning brown. No growth at 55° C. On milk, after 4-5 days, coagulation followed by peptoniza- tion, mycelium brownish with violaceous tint. Acremoniella (Acremoniopsis) olivaespora Ciferri & Ashford, Mycologia 22: 62-68, 2 figs., 1930. A saprophyte isolated from human skin. SPOROTRICHUM Sporothrichum Link, Mag. Ges. Naturf. Freunde Berlin 3: 12, 13, 1809. The type species is Sporothrichum hadium Link. In the original description of the genus eleven species are described as new, and S. virescens and S. abietinum transferred here from Persoon's genus Dematium. Most of the species were bright colored and were found on decaying wood. Since S. hadium is the only one figured and the first to be described, it may be taken as the type of the genus. SPOROTRICHEAE 799 Link describes the genus as follows: 3. Sporothrichum. Thallus e flocvis vntricatis decumbentibus aut erectiusculis, ramosis, septatis, Sporidia ubique inspersa, rotumdata. Caespitibus late effusis truncos caesas aliaque vegetabilia putrescentia investiunt. Varios hdbent colores, saepe pulchtrriinos. Sporidia plerumque globosa, parva aut minuta, varia copia inspersa. Fleraeque species persistunt. Genus hinc affine Asporotricho et Dematio quibus vero sporidia nulla, inde Botryti, a quanam floccis decumbentibus differt a Geotricho, Trichothecio, Epochnio discrepat sporidiis multo minoribus nee truncatis, nee didymis nee appendiculatis. Fleraeque species nondum descriptae. Sp. badiutn caespitibus tcnmbus, floccis badiis decumbentibus, sporidiis parvis. Tomenti instar tenuis truncos emortiio^is hurmdos laic obducit, saepe Sphaeriis incumbens. Sporidia non ita crebra. Iconem V. fig. 14. A study of Link's Fig. 14 shows a repent, septate mycelium with scattered lateral, nearly spherical conidia, no erect conidiophores. In 1815, Link again treated the genus, adding his previous Asporothricum and Dematium. He divided his genus into two subgenera, Lysisporvwm, thallus septate, densely covered with spores, easily falling apart, and Alytosporiuvi, thallus either septate or not, conidia strongly adherent, rarely absent. His type species, S. badium, belongs in the latter. The first sub- genus contained thirteen species, the second twelve. The descriptions are extremely brief, based largely on the host. As a whole, the genus seemed to be saprophytic on deadwood, although fallen leaves and earth are mentioned. In 1818, Link again treated the genus, adding species which he had previously treated in Aleurisma and Collarvum and separating Alytosporium. He no longer recognized subgenera and arranged his key strictly on the basis of color. Thus, his genus Sporotrichum of 1818 becomes the same as his genus Aletirisma of 3S09, while his Sporothrichum becomes Alyto- sporium, in 1818. Martins, Flora Cryptogamica Erlangensis 335-337, 1817, treated the genus, including only species already described by Nees and Link. Nees, Das System der Pilse und Schiodmme 1816, described iSi. laxu/m and mentions sev- eral of Link's species. He would select the group of S. fulvum, S. badvumi, etc., as true Sporotrichum. In Gray's Natural Arrangement of British Flants 550-551, 1821, 5 species are described, all originally described in Link, 1809, including ;Si. badium; Aleurisma not mentioned. In his Sy sterna Mycologicum 3: 415-425, 1832, Fries treats 32 species, arranging them by color, as Link had done in 1818. Saccardo, Michelia 2: 16, 1882, considers S. rosewm or S. virescens as the type of the genus. In 1885, Saccardo & Marchal (Bull. Soc. Eoy. Bot, Belgique 24: 65) described Ehino- cladvum based on B. coprogenum from rabbit dung (Fig. 121) later adding Sporotrichum torulosum Bonorden. This genus was evidently erected to be analogous to Sporotrichum, but differed in having black spores. It is said to differ from Trichosporium, Fries by the presence of sterigmata. As the presence or absence of color is rather variable in this group, it has seemed best to retain Sporotrichum, although the commonest species, S. SchencJci, might with equal propriety be placed in Bhinocladium, as is frequently done by French workers. Hyphae septate, repent, irregularly branched. Conidiophores not differen- tiated, or at most, a terminal conidium on a short branch ; conidia lateral or terminal, often sessile or on a short sterigma, ovoid or spherical, hyaline or light colored, usually small. This is a large, rather poorly defined genus of saprophytes, which has been used as a place to put poorly defined mycelia with spores ever since it was first created. Some of the older mycologists, such as Saccardo and Lindau, placed most of the dermatophytes here without regard to their evi- 800 MEDICAL MYCOLOGY dent relationsliijjs. Tliere are few pathogenic species, but one of them, 8. Schencki and its \'ariety Beurmanni, is rather widespread and serious. The organism often enters through a wound, such as a prick of a thorn, and produces cutaneous lesions which may or may not later become systemic. The cutaneous types, in which multiple subcutaneous gummata develop with or without ulceration, appear to be the most frequent type in France, while lymphangitic sporotrichosis and adenitis seem to be more frequent in the American literature. AVhere the disease becomes systemic, various organs may be involved, such as the mucous membranes, muscles, bones, joints, very rarely the testicles, epididymis, lungs, and brain. For a full recent discussion of the clinical aspects of this disease, the reader is referred to the excellent account in Beurmann & Gougerot, Les SporotricJioses, 1912, and Jacobson, Fungous Diseases 128-145, 1932. Fig. 121. — Rhino claMum coprogenuni Saccardo & Marchal. ' (After MarchaL) The delimitation of species in this genus is very difficult. Color, which has often been used not only to separate species but even genera, has been shown by Davis (1915) to be variable. Practically all the strains that he tried formed pigment when first isolated; carrot, potato, 3% maltose agar, and 3% glucose agar are more suitable media for pigment production, as is also an abundance of oxygen, while light is apparently without effect. Albino strains frequently occur and remain fixed even after passage through an ex- perimental animal. Meyer & Aird (1915), after a study of 18 strains, found some differences in carbohydrate fermentation, but these were not sufficiently constant or correlated with other characters to warrant separation into spe- cies. Meyer (1915), while admitting that rarely is the equine sporotrichosis transmitted to man. w^as unable to find sufficient differeiiees in the strains to separate them. SPOROTRICHEAE 801 Benham and Kesten (1932), working with 8porotrichum Schencki pro- duced nodnlar sporotrichosis in a monkey similar to the lymphangitic type in man. Thej' found the human disease transmissible to carnations, in which it produced lesions similar to those produced by S. Poae, a common pathogen of these flowers. After living parasitically and saprophytically on plants, S. Schencki retained its virulence for animals while 8. Poae and *S^. pruinosum, from the soil, showed no pathogenicity for experimental animals at any time. Lesions produced by Sporotrichum respond readily to medication with po- tassium iodide, although this salt is not toxic to the fungus in cultures. Davis (1919) suggests that it may stimulate the host cells to proliferate and heal the lesions. Key to Species Colonies remaining white or pink on all merlia. Gelatin not liquefied. Colony becoming pink on potato. S. Grigshyi. Colony becoming yellowish on potato. S. Fonsecai. Gelatin liquefied. Milk not coagulated, producing a pellicle on liquid media, clavate bodies in lesions as in Actinomyces hovis ; from subcutaneous lesions. S. asteroides. Milk coagulated, producing islets which quickly settle, without forming pellicle on liquid media, no clavate bodies in lesions. Ulcers in mouth in man. S. cracoviense. Lesions on horse similar to those produced by Zymonema farciminosus ; Madagascar. S. equL Action on milk unknown, little or no growth on liquid media, no clavate bodies in lesions; ulcers. S. Carougecmi. Colonies often becoming black or brown on most media. Gelatin not liquefied. s. Jeanselmei. Gelatin slowly liquefied. S. SchencM. Sporotrichum Grigsbyi Dodge, n. sp. Sporotrichum Schenckii Grigsby and Moore, Southern Med. Jour. 15: 684- (?87, 1922, non aliorum. Isolated from lesions on chest, elbow, and neck. Morphology as described from smears unintelligible. Aerobe, sporogenous, nonliquefying. Pinkish colony after 6 days on po- tato or plain agar. Colony otherwise white, dry, heaped up, and wrinkled. No indol. No fermentation of glucose, sucrose, or lactose broth, no turbidity, sediment granular, pellicle(?) formed. Litmus milk unchanged. Sporotrichum Fonsecai Pereira Filho, Rev. ]Med. Chirurg. Brasil 37: 265, 26G. 1 pi., 1929 ; Ihid. 38 : [3-21 ] , P/.s. 1-24, 1930. Rhinocladium Fonsecai Grandinetti, Centr. Estudo Esporotricose Sao Paulo 52, 1934. Isolated from abscesses on the nose, Brazil. Pathogenic to rats and mice. Mycelium branched, often in large strands of parallel hyphae, conidia mostly terminal on these strands, 9-20 x l-2fi, 2-7 x 1-2/a; chlamydospores abun- dant in old cultures. 802 MEDICAL MYCOLOGY On plain agar, colony white, center granular, margin striate. On grape juice agar, growth more rapid and medium somewhat darkened. On Sabou- raud agar, colony white, creamy, cerebriform, very adherent, yellowish, finally becoming slightly brownish. Colony similar, but remaining white on conser- vation agar. On human blood agar, growth rapid, slightly brownish. Similar on glycerol agar. On potato and potato glycerol, colony white at first, becom- ing yellowish. On bile potato, colonies prominent, yellowish, surface granular, with depressions finally somewhat brownish. On carrot, beet, banana, sweet po- tato, turnip or manihot, colonies white, eventually becoming slightly brownish, adherent. On gelatin at 17° C, colony white, prominent, without liquefaction; medium becoming brownish. Glucose gelatin finally liquefied. On plain broth, floccose sediment, upper portion of the liquid clear. On glucose broth, a pel- licle is formed, Avhich soon settles to the bottom without breaking up. No fermentation of sugars; acid with glucose, fructose, and m.altose. Milk sup- ports rapid growth and is coagulated. Sporotrichum asteroides Splendore, Rev. Soc. Sci. Sao Paulo 3 : 62, 1908 ; Brasil Med. 33: 361-365, 2 pis., 1909. Rhino cladium Beurmanni var. asteroides Vuillemin, 1910. Sporotrichum Beurmanni var. asteroides, Beurmann & Gougerot, Arch, de Parasitol. 15: 40-45, 1911. Rhinotrichum asteroides Verdun, Precis Parasitol. 1912; Verdun & Man- doul, Precis Parasitol. 714, 715, 1924. Sporothrix asteroides Davis, Jour. Infect. Uis. 12: 453-458, 1913. Rhinocladium asteroides Grandinetti, Contr. Estudo Esporotricose Sao Paulo 53, 1934. Found in subcutaneous lesions, in general much as in the European S. Beur- manni and North American ;S'. Schencki (case history and pathology in full in Brasil Med. 33: 361-365, 2 pis., 1909). In experimental lesions, organism regu- larly produces ray-shaped bodies suggesting those seen in Actinomyces hovis, hence the name. At first budding forms, then septate hyphae, spores at first isolated and hyaline, lateral as well as terminal, later crowded, often verticillate and dark colored. Optimum temperature about 30° C, organism growing more slowly at lower temperatures. Grows on all the common media, but prefers media with glucose and slightly acid. Growth good on slightly acidified rye grains, slower on potato, without pigmentation on the latter. Slowly liquefies gelatin, does not coagu- late milk. Growth superficial on liquid media, leaving the liquid clear. Sporotrichum cracoviense Lipinski, Medycyna Doswiadczalna i Spoteczna 2: 153-169, 7 figs., 1924. Oospora sp. Griitz, Sporotrichosen und verwandte Krankheiten, Handb. Haut- Geschlechtskr. [Jadassohn] 11 : 797-799, 1928. SPOROTRICHEAE 803 Found in lesions on the tongue of an otherwise healthy ten-year-old girl. First ulcer on tongue, then on hard and soft palates, and on tonsils. Some lesions were small, hard, pinhead size, grayish white, reddish below, removed by platinum loop with difficulty but without causing pain. Other lesions, on tonsils and hard palate, apparently derived by softening of nodules, were flat, shallow, ovoid, with irregular edges. Glands not swollen, temperature nor- mal. The patient had returned from the Soviet Republic only a short while before. Patient seen only once, since mother refused to permit hospitalization or further treatment. Pathogenic to white rats. In the nodules, cells spherical or ovoid, 2-3 x 6-7/a. Hypliae short, branched, terminal cells swollen. Gram-positive, old hyphae not staining, acid-fast. Some are partly gram-positive and partly negative. In cultures, sterile and fertile cells grow along side of hyphae, sessile or in fine sterigmata, either singly or in groups. Chlamydospores present. This organism grows well on a variety of liquid or solid media either with or without sugar, aerobically or anaerobically, between 18° and 37° C. On Sabouraud agar, colony circular or slightly oval, silvery white, dull, slowly becoming convex, elevated above the surface of the agar with a delicate, vel- vety margin, growing at an irregular rate so that it finally becomes cerebri- form. Reverse concolorous even after 10 weeks. Growth on potato rather poorer than is the case with other species, but growth good in water of con- densation. On Loeffler's medium, growth good and characteristic, center colo- nies being surrounded by an aureole with rays like icicles. On gelatin, either with or without sugar, growth good, colony becoming crateriform with a wide rim, finally, as the medium liquefies, reaching the bottom. No characteristic growth on blood agar, no hemolysis. In common broth, tiny surface colonies appear after 24 hours. These settle to the bottom on shaking. Odor nauseat- ing on standing. Growth still good in broth covered with paraffin. Milk is coagulated in 6 days, with sediment and turbidity of the medium. Pleomorphisni. — At room temperature (more slowly at 35°-37° C), the center of colony becomes covered with rugose coremia, destroying the appear- ance of the colony. At thirty-fourth day, colony rounded, with or without development below the surface of the medium, colorless at first, later becom- ing pearly, retaining aureole, adherent. Appearance changes on subcultures. Yeast forms are produced on many different media when temperature is changed. Colonies consisting of yeast cells small, flat, oval, brownish white, slimy. Sporotrichum equi, Carougeau, Jour. Med. Vet. Zootechnie 60: 8-22; 75-90; 148-153, 1909. Epizootic on horses in Madagascar, symptoms similar to those caused by Zymonema farciminosum, with which it since has sometimes been confused. Thought by Hyde & Davis, Jour. Cutan. Dis. 28: 321-352, Pis. 36^5, 1910, to be identical with S. Schencki as in America, organisms in horse and man are identical. It is distinct from Zymonema farciminosum. 804 MEDICAL MYCOLOGY Mycelium 1.5-2;u in diameter, spores on a very short pedicel, often in groups of 6-10, spores ovoid to pyriform, sometimes nearly spherical in old cultures. Hypnospores formed in old cultures. On ordinary agar or glycerol agar, colonies star-shaped or irregularly dentate, circular, slightly elevated, whitish, shining, very adherent, penetrat- ing the medium 1-3 mm. On Sabouraud agar, colonies round, yellowish white, elevated, adherent, surface folded. On potato, colonies granular, verrucose, grayish white, dry, not abundant. With glycerol added, the growth is much more luxuriant. On coagulated ox serum, development slow, colonies small and with black points. In peptone meat broth or, better, in that to which glucose or glycerol has been added, colonies white, fioccose, attached to sides and bottom of the tube, liquid remaining clear even on shaking. In glucose or glycerol broth also, floating colonies as small islands. Milk coagulated in 2 weeks. Gelatin liquefied slowly after several weeks. Sporotrichum Carougeaui Langeron apud Brumpt, Precis Parasitol. ed 2, 1913 (fide Langeron Bull. Soc. Path. Exot. 15: 453-459, 1922, footnote) ; Fon- toynont & Carougeau, Bull. Soc. Path. Exot. 15 : 444-453, 1922. Rhinocladium Carougeaui Neveu-Lemaire, Precis Parasitol. Hum. 85, 1921. Found in cutaneous ulcerous lesions in the hands of a six-year-old child, also suffering from tuberculosis. Medication with KI helped, but did not cure at once. Ten years later the child was well. Fontoynont & Carougeau give ease history in detail. Mycelium septate, rampant, 2.5-5/j, in diameter (mean 3/i,), more volumi- nous than in Rhinocladium. Cells 8-10/^ long; conidia ellipsoid, 2 x 4;u,, becom- ing spherical, 4-4.5 and even 5/a in diameter when detached. Conidia attached by definite sterigmata, arising variously all along the hyphae. Occasionally yeast forms produced both in culture and in lesions. Best growth at 19°-22° C. Organism easily isolated on sugar media. On agar, colony creamy, white. On glycerol agar or maltose or glucose agars, groAvth is equally favorable, colony elevated, wrinkled, vermiculate, adherent, without the sharp peak in center characteristic of S. Beurnianni. Folds vary from 0.5 to 4 mm. thick. Color silvery white on glycerol or glucose agar, yellowish on maltose. On manioc with glycerol, growth less abundant than on agar, forming a white, thin, farin- aceous colonj^ without folds, becoming slightly tomentose in 2 weeks. On potato glycerol, colony 1-2 mm. thick, white or slightly grayish, surface irregular with coremia, and white flakes floating in the glj^cerol solution at the bottom of the tube. On carrot glycerol or turnip glycerol, colony similar but more humid, creamy, growth slow. On banana, colony folded, gray on reddish back- ground, spreading very little, formed of small, rounded, confluent colonies, slightly brown. On gelatin stab, surface colony folded, white, with inverted pine tree below; liquefaction on tenth day. On green bamboo bark, growth good, farinaceous, white. Liquid media (glycerol or sugars), show good growth in flakes, which settle to bottom but are easily suspended on shaking. Little or no development in broth. SPOROTRICHEAE 805 Sporotrichum Jeanselmei Bninipt & Laiigeron, Bull. Mem. Soc. Med. Hop. Paris III 29: 792-796, 1910; also note in III 29: 824, 1910. Bull. Soc. FrauQ. Derm. SyphiligT. 21 : 190-192. 1910. (First case, Jeanselme & Chevallier, Bull. Mem. Soc. Med. Hop. Paris III, 29: 784-792, 1910.) Rhinodadium Jeanselmei Grandinetti, Contr. Estudo Esporotricose Sao Paulo 53, 1934. Found in widespread lesions, closely resembling syphilitic gummata, on skin, bones, etc. Medication with large doses of KI promptly cleared up the lesions. Probably the case of Bedell (1914), from the eye, should be referred here. Cultures of this organism show all stages between blastospores fonned directly on conidia and well-developed mycelium. Occasionally, one spore has four spores attached by sterigmata or by a short germ tube, bearing the tuft of spores on its end, as in smuts. Mycelium, as usual in Sporotrichum, com- posed of elongate tine filaments, abundantly branched, tangled, septate; spores either in small tufts or isolated along the filaments. Tufts either terminal or at the ends of short, lateral branches. Each spore, borne on a short sterigma, brownish and spherical when detached. Tufts usually only 4-5-spored. Hyphae 1.5-2/x in diameter, spores 2.5-3.5/x in diameter, sterigmata 1-1.5 x 0.5-0.85/a. Sporotrichnm homhycimon Corda differs in having filaments thicker, spores hyaline, attached singly to branches 15-26/^ long and of nearly the same diameter as the principal filament. ^S^, Beurmanni has mycelium more slender and more abundant, 1.5-1.7/x in diameter. Short forms exceptional. Tufts of spores larger, containing 12-15 spores. Lateral spores closer together, ellipsoid, 3.5 X 2.6/x, or pyriform, 4 x 2.4a'- On most media, colonies compact, resistant, adherent. Organism grows on usual media. Growth best at 35°-37° C. On Sabouraud maltose agar, growth tomentose. On glucose peptone agar (first isolation), growth in 8 days, center of colony smooth, then a ring of striate radiating crests and out- side a "crown of rays." Colonies cream color to old ivorJ^ On potato with- out glycerol, white velvety colonies on fourth to eighth day. Thirty-four days later, colonies size of lentils, flat, furrowed, with radiating folds and little sharp peaks. On potato glucose in incubator, growth visible on sixth day, cream white, blackening on fourteenth day. On thirty-fourth day, surface cov- ered, coal black, in hollows, slightly ashy, powdery on ridges. On carrot, white colonies, becoming black with white border. Growth much slower than on potato. On gelatin, smooth pearly little points on the surface, radiating and filamentous on walls of the tube after 3-4 weeks. Center of colony a conic umbo ; periphery, a white zone with fine striations at bottom of a deep umbilical depression. In broth, a fragile pellicle composed of small white nodules surrounded by rays, mediiun remains clear. Gelatin not liquefied. Sporotrichum Schencki ]Matruchot, C. R. Acad. Sei. 150: 544, 1910. Sporotrichum sp. E. F. Smith in Schenck, Johns Hopkins Hosp. Bull. 9: 286-290; 1 pi, 1898. 806 MEDICAL MYCOLOGY Sporothrix SchencH Hektoen & Perkins, Jour. Exp. Med. 5: 77-89, Pis. 2, 3, 1900. Bhinocladnim Schenki Verdun & Mandoul, Precis Parasitol. 713, 714, 1924. Rhinocladium Schencki Grandinetti, Contr. Estudo Esporotricose Sao Paulo 18, 1934. Bhinotrichum Schenkii Ota, Jap. Jour. Derm. Urol. 28: [4], 1928. Hyphae about 2/x in diameter, irregularly staining with Gram '^s stain. My- celium formed of parallel strands of curved liyphae, few or no lateral branches. Spores rare along the hyphae, usually terminating them, 3-5ju, in diameter, ovoid or apiculate (Fig. 122), staining with Gram stain. Optimum temperature 30° -38° C, growth slow at room temperature. Colo- nies show straight furrows, not cerebriform. Lactose fermented, not sucrose. Culture remains colorless indefinitely, with a brownish pigment developing in the aerial portions. Spores hyaline, aerial fructification normal. In deeper and moister portions of the cultures, spores sprout below into short, irregular chains, 5-8 spores long. — Matruchot. The following data come from the original description. Colony on agar white, wrinkled, becoming brownish or even dark brown and velvety. Growth on potato yellow, becoming brown, medium darkening. Tufts of mycelium ap- pear in broth, also islets on the surface. Litmus milk is unchanged, with little growth occurring. No fermentation, no liquefaction of serum. Gelatin slowly liquefied. Chlamydospores appear in media poor in nutrients, but are not abundant, as in some strains of Sporotrichum Beurmanni. Davis (1914) believes that the chlamydospores do not afford a sufficient criterion for the separation of these two species. Var. Beurmanni (Matruchot & Ramond) Dodge, n. comb. Sporotrichum Beurmanni, Matruchot & Ramond, C. R. Soc, Biol. 57: 379, 1905. (First case Beurmann & Ramond, Ann. Derm. Syphiligr. IV, 4: 678, 1903.) Bhinocladium Beurmanni Vuillemin, in litt. apud Beurmann & Gougerot. Arch, de Parasitol. 15: 98, 1911. Sporotrichopsis Beurmanni Gueguen, apud Beurmann & Gougerot, Arch, de Parasitol. 15: 103, 104, 1911. Bhinotrichum Beurmanni Ota, Jap. Jour. Derm. Urol. 28: [4], 1928. Isolated from the pus in small tumors in the subcutaneous tissues. Tu- mors possess definite walls, become about the size of a peach stone, are at first painless, later becoming painful on pressure, finally filled with odorless, granular pus. Mycelium recumbent, 2/x in diameter, colorless, much branched, tangled. Fructifications are composed of large, cylindric masses, lOjji in diameter, some- times elongated. Spores terminal on long hyphae or on branches, solitary. SPOROTRICHEAE 807 variable in number, irregularly borne, pyriform, rarely on a sterigma, 1-2/x long by 0.5/x, in diameter. Spores become ovoid and brown, 3-5 x 2-4/a. Opti- mum temperature 22°-30° C, growth slow at 37° C. Colonies on Blanchetiere & Gougerot medium (carbohydrate 30 gm., pep- tone 10 gm., water 1,000 c.c.) at 22°-25° C, are confluent, white at first, rapidly Fig. 122. — Sporotrichum Schenki var. Beurmanni. 1, 2, germinating spores ; S, l), 15, SO, oldla on various media ; i, intercalary chlamydospore ; 5, yeastlike cells ; 6, young filament ; 7, terminal chlamydospore ; 8-lS, 16, IS, 2Z-Zi, conidia showing: variation in arrangement on hyphae in various media ; n, 19-21, mycelium with chlamydospores on various media. (After Moore 1934.) 808 MEDICAL MYCOLOGY and completely becoming dark brown when the fructifications form, cerebri- form, convoluted. Same on carrot or potato. Acid formation with glycerol, glucose, galactose, sucrose(?), maltose, inulin, starch (slight), but not with mannite, dulcite, dextrin, lactose. When CaCOg is present acetic acid is also formed. This variety is doubtfully distinct from the species, as has frequently been pointed out. Var. Greconis Dodge, n. var. See Mackinnon, Arch. Soc. Biol. Montevideo, Suppl. Fasc. 5: 1320-1327, 1931. Agrees with above, but does not acidify sucrose. Var. Fioccoi, Dodge, n. var. Sporotrichum epigacnim Aschieri, Atti 1st. Bot. R. Univ. Pavia IV 1: 199- 222, 8 figs., 2 graphs, 1930. Not Brunaud, Nouv. Fragm. Mycol. II, 11, Bor- deaux, 1888; Saccardo, Syll. Fung. 10: 534, 1892. Sporotrichosis ease of Fiocco. Conidiophores 2.5-4/a in diameter, varying in length with the media, septate, straight or variously contorted, with short lateral branches; spores sessile, 2.5-3/x in diameter, rounded or slightly elongate ; chlamydospores in- tercalary or terminal, spherical, 7-10/x in diameter, or elongate, 7 x 10/a. Conidia solitary or sympodially in small groups on short branches or in whorls, some- times suggesting conditions seen in Beauveria. Growth curves are given for 5° intervals between 5° and 30°, showing an optimum at 20°. Colonies on Pollacci agar whitish, floccose, smooth becoming rugose, ca- nary yellow in spots. Aschieri reports development of chestnut colonies, moist, surface verrucose covered with white conidial hyphae ; sometimes a sector, completely covered with conidial hyphae and surrounded by a yellow ring. In Saboviraud agar, colony smaller, verrucose, covered with a white layer of mycelium with a yellow margin 3 mm. broad. On potato, colony whitish, covered by a white cottony layer. On carrot, colony similar, but with a slightly yellowish color. On liquid media, a ring developed with a gelatinous mass in the bottom of the tube. Var. Oouncilmani (Wolbach) Dodge, n. comb. Sporotrichum Councilmani Wolbach, Jour. Med. Res. 36: 327-355, 4 pis., 1917. Ehinotrichum Councilma7ii Ota, Jap. Jour. Derm. Urol. 28: [4], 1928. Rhinocladium Councilmani Grandinetti, Contr. Estudo Esporotricose Sao Paulo 53, 1934. Isolated from a lesion in the knee, which followed a wound from a nail in an ash barrel, Boston. Pain developed in one week. Case unimproved after 2 months, with a variety of treatments. Knee joint fixed. In about 5 months, pain disappeared, but joint still completely fixed. Pathogenic to rabbits and guinea pigs, giving the typical lesion. SPOROTRICHEAE 809 Hyphae 1-1.25/1, in diameter, branching, irregular, the young sporiferous branch often somewhat swollen in the proximal region, septate, with the septa occurring at intervals of from 10-22/*. Spores hyaline, 1-celled, pyriform, 7-6 X 2.5-3. 5/t, solitary or usually in clusters, formed by successive apical proliferation at the tips of aerial sporophores, rarely sessile on the walls of the vegetative hyphae, the point of attachment being marked by a slight denticulation of the fertile branch. Pleomorphic mycelium is characterized by a free aerial growth of hyphae, abundant spore formation, large size of spores, absence of lateral spore clusters, and occurrence of septate branching fila- ments in lesions. Doubtful Position Sporotrichum congolensis (Baerts) Dodge, n. comb. Discomyces congolensis Baerts, Bull. Med. Katanga. 2: 67-75, 1925. Actinomyces congolensis Brumpt, Precis Parasitol. ed. 4, 1206, 1927. Lesions originally described by van Nitsen, Ann. Soc. Beige Med. Trop. 1 : 1920. Found in subcutaneous nodules of pea size, later growing to pigeon egg size, becoming adherent to the skin and softening. On puncture, these yield a clear, viscid liquid which soon becomes purulent, yielding mycelium. These ruptured nodules may either heal spontaneously or be secondarily infected. Some ulcers, with irregular borders, gradually extend, involving muscles, joints, rarely also bones. The pus shows small yellowish or grayish gelatinous grains, 1-2 mm. in diameter, some even being as large as 3-5 mm. Medication with arsenicals, salvarsan, and KI ineffective. In the grains, hyphae cylindric, long aseptate, glassy, hyaline, varying from 0.33-5/x in diameter, most being l-3/i, dichotomous, straight, curved or spiral, with lateral, rarely terminal, pyriform spores, 8-9 x 2-4/a. Organism not cultivated. This is evidently not an Actinomyces, probably an Indiella or Sporotrichum, but is difficult to place in the absence of figures or more adequate description. Rhinocladium guayaquilense Valenzuela. Brumpt, Precis Parasitol. ed. 4, 1329, 1927. Sporotriclium indicum Castellani & Chalmers, Man. Trop. Med. 622, 1910. Rhinocladium indicum Vuillemin apud Verdun & Mandoul, Precis Para- sitol. 115, 1924. Sporotrichum Beurmanni var. indicum Beurmann & Gougerot, Les Sporo- trichoses 143, 1912. Imperfectly described and cultures lost. Sporotrichum Lesnei (Vuillemin) Castellani & Chalmers, Man. Trop. Med. ed. 3, 1121, 1919. Rhinocladium Lesnei Vuillemin, Bull. Soc. Sci. Nancy 11: 139-144 (10-15), PI. 1, 1910. 810 MEDICAL MYCOLOGY Originally described as producing mycetoma pedis in Madagascar. Fon- toynont & Carougeau, m litt, published by Beurmann & Gougerot, Arch, de Parasitol. 15: 13, 1911, state that it was an impurity found in a secondary infection of a lesion of a different nature. Hyphae branched, septate, l.Sfx. in diameter, scarcely fuliginous, discrete or fasciculate, and then ascending or erect. Conidia oblong or ovoid, short pedicellate, fuliginous, 4-7 x 3-3.5/*, solitary on hyi^hae, occasionally or fre- quently with denticulate apex, cylindric or rarely nodose. The fasciculate hyphae simulate Graphium (Fig. 123). Sporotrichum lipsiense Benedek, Derm. Woch. 83: 1770-1777, 1926. Isolated from various superficial epidermal lesions accompanied by pru- ritus and intertrigo. Fig. 12Z.— Sporotrichum Lesnei Vuillemin. (After Vuillemin 1910.) Mycelium septate, 2-4/x, mostly 3/a, in diameter, cells 8-12/1, long, occasion- ally 20-28/x. No differentiated conidiophores, spores lateral or terminal ovoid and almost pyriform, becoming spherical after abjunction, sessile or with a short slender sterigma, 3-4/a in diameter. Chlamydospores 4 x 5-8 x 12/a. Sprouting forms 6 x 6-5 x Sfi, hyaline. Cultural optimum temperature 18°- 20°, growth ceasing at 25° C. On glucose agar, glycerol, potato or glycerol carrot, growth gray white, dull, moist, flat, becoming rough from tufts of hyphae and finally white velvety. Ferments glycerol and mannitol, not dul- citol, glucose, galactose, fructose, sucrose, maltose, lactose, dextrin, starch, or inulin. Sporotrichum bronchiale Montague, Plant. Cell. Nouv. 92 [cited by Sac- cardo, Syll. Fung. 4: 100, 1886]. Isolated from bronchomycosis, case of Gubler. SPOROTRICHEAE 811 Hyphae white, branched, repent or erect ; septate, cells 5-7/a in diameter ; conidia hyaline, spherical, 5/* in diameter. I have been unable to locate the original place of publication, and suspect that Saccardo may have been in error, as it is not in Montague's Cent. Plant. Cellulaires Nouv. No. 92, Ann. Sci. Nat. Bot. II, 20: 370-372, 1844. Montague did not recognize the species when he compiled his Sylloge Gen. Spec. PI. Cryptog. 1856, containing all the species he had previously described in sys- tematic order. Castellani & Chalmers do not quote an authority for their statements, and Jacobson copies Castellani & Chalmers. Rhinocladium parvidum Redaelli, 1923 ; Grandinetti, Contr. Estudo Espo- rotricose Sao Paulo, 43, 1934. Sporotrichum parvulum Brunaud. Isolated from lungs in cases clinically resembling tuberculosis. BIBLIOGRAPHY Achard, Ch. & Louis Ramond. 1909. Sporotricho-tuberculose, Bull. Mem. Soc. Med. Hop. Paris III, 27: 738-743. Adamson, H. G. 1908. Sporotrichosis, a resume of the literature relating to sporotrichal infections of the skin, Brit. Jour. Derm. Syphilis 20: 296-303. — . 1911. A case of sporotrichosis, Brit. Jour. Derm. Syphilis 23: 239-246, Pis. 1, 2, Figs. 1, 2. — . 1913. A case of sporotrichosis of the disseminated, ulcerating, gumma type, in which there occurred acute synovitis, Brit. Jour. Derm,. Syphilis 25: 33-37, 4 figs. — . 1913. A case of sporotrichosis simulating blastomycosis, Brit. Jmbr. Derm. Syphilis 25: 60-64, 2 figs. — . 1922. Sporotrichosis (with cultures) [described by T. Joekes], Proc. E. Soc. Med. Derm. Sect. 15: 18, 19. Agostini, Angela. 1931. On Blastomycoides lanuginosns Castellani, Jour. Trop. Med. Hyg. 34: 287-288, 2 figs. — . 1932. Sul Blastomycoides lanuginosus Castellani, Atti. 1st. Bot. E. Univ. Pavia TV, 3: 65-68, 2 figs. Aguiar Pupo, Joao de. 1920. Sporotrichoses no Brasil, Annaes Paulistas Med. Cirurg. 11: 200-207. Aleixo, A. 1921. E sporotrichose produzida por um esporotricho em forma de levedo, Brasil Med. 35: 2:283. Arndt, G. 1909. Vorlaufige Mitteilung iiber einen Fall von Sporotrichose der Haut, Berlin. Klin. Woch. 46: 1966-1968. — . 1910. Beitrag zur Kenntnis der Sporotrichose der Haut, mit besonderer Beriicksichtigung der Lymphangitis sporotrichotica. Experimentelle Sporotrichose, Derm. Zeitschr. 17: 24-53, 171-207, Pis. 1, 2. Aroeira Neves. 1929. Contribui^ao ao estudo da esporotrichose familiar, Brasil Med. 43: 92-94. Aschieri, Eugenia. 1929. Un Sporotrichum nuovo parassita dell'uomo, Atti 1st. Bot. B. Univ. Pavia IV, 1: 199-222, 8 figs. Auche, B. & A. LeDantec. 1894. fitude d'une nouvelle mucedinee pyogene parasite de I'homme (variete Botrytis), Arch. Med. Exp. I, 6: 853-861, 4 figs. Ballagi, Stepan. 1932. Mykologische Beschreibung der Acremoniosis, Arch. Derm. Syphilis 166: 405-407, 2 figs. Basile, Giovanni. 1915. Sporotricosi sperimentale con particolare riguardo alia oto-rino- laringologia, Ann. di Clin. Med. 6: 264-294, Pis. 1-S. 812 MEDICAL MYCOLOGY Battaglia, Mario. 3912. Contribute alio studio delle sporotricosi, Ann. Med. Nav. Colon. 18: 2:1-11. Bedell, Arthur J. 1914. A case of chronic sporotrichosis of the eye, Ann. Ophthamol. 23: 605-617, 5 figs. Bellucci, Luigi. 1925. Di alcuni importanti reperti micotici in oto-rino-laringologia (Acremonium tonsillare — Ozena nasale — Otite media da Sterigmatocystis ochracea), Atti B. Accad. Fisiocrit. Siena IX, 17: 17-46, 16 figs, [same title, Arch. Ital. Otol. 37: 1-21, S pis., 1926]. Benedek, Tibor. 1926. Beitrag zur Kenntnis der rein epidermal Sporotrichosen. Dermo- epidermitis sporotrichotica verursacht durch eine neue Art von pathogenen Sporo- trichum: Sporotrichum lipsiense nov. spec, unter besonderer Beriicksichtigung der Serumdiagnose und der experiment sUen Sporotrichose, Derm. Woch. 83: 1695-1703; 1733-1738; 1770-1777; 1803-1807; 1824-1827. Benham, Ehoda W. & Beatrice Keston. 1932. Sporotrichosis; its transmission to plants and animals, Jmir. Infect. Dis. 50: 437-458, 15 figs. Berti, Giuseppe. 1922. Kicerche sperimentali sull'azione tossica del dello Sporotrichum Beurmanni, Eiv. Biol. 4: 44-50. Beurmann, Lucien de. 1912. On sporotrichosis [translated by R. W. MacKenna], Brit. Med. Jour. 2: 289-296. — . 1912. Une forme uouvelle de sporotricliose. Elephantiasis sporotrichosique gommeux, ou pied de Madura sporotrichosique, BiiU. Mem. Soc. Med. Hop. Paris III, 34: 621- 628, 1 fig. — . 1914. Un cas de sporotrichose osseuse chronique a forme rhumatoide avec gomme mam- maire spontanement ouverte . . . Bull. Mem: Soc. Med. Hop. Paris III, 38: 379-384. Beurmann, Lucien de, Brodier and Gastou. 1907. Sporotrichose gommeuse disseminee avee lesions laryngees, Bidl. Mem. Soc. Med. Hop. Paris III, 24: 1060-1069. Beurmann, Lucien de & H. Gougerot. 1906. Les sporotrichoses hypodermiques, Ann. Derm. SypUUgr. IV, 7: 837-864; 914-922; 993-1006. — . 1907. Etiologie et pathogenic de la sporotrichose, Tribune Med. 40: 693-696. — . 1907. Sixieme cas de sporotrichose sous-cutanee et cutanee. Bull. Mem,. Soc. Med. Hop. Paris III, 24: 309-313. — . 1907. Les sporotrichoses sous-eutanees, Bidl. Mem. Soc. Med. Hop. Paris III, 24: 302- 305. — . 1907. Sporotrichoses tuberculoides, Ann. Derm. Syphiligr. TV, 8: 497-544; 603-635; 655- 679. — . 1907. Associations morbides dans les sporotrichoses. Onzieme observation de sporo- trichose; syphilis, tuberculose et sporotrichose. Bull. Mem. Soc. Med. Hop. Paris III, 24: 591-596. 12« observation. Ibid. 24: 596-610. — . 1908. Sporotrichoses americaines. Diffusion du Sporotrichum Beurmanni, Bull. Mem. Soc. Med. Hop. Paris III, 25: 733-741. — . 1909. Comparaison des sporotrichoses et des infections cocciennes. Sporotrichose aigues et subaigues disseminees. Sporotrichomes a evolution phlegmasique, Ann. Derm. Syphiligr. IV, 10: 81-98. — . 1910. Comparaison du Sporotrichum Jeanselmei et des Sporotrichum voisins, Bull. Mem. Soc. Med. Hop. Paris HI, 30: 818-823. — . 1910. Importance pratique du diagnostique de la sporotrichose et des autres mycoses facilite et difficulte de ce diagnostic, Bev. Med. Hyg. Trop. 7: 185-197. — . 1910. Sporotrichoses nord-americains. Bull. Mem. Soc. Med. Hop. Paris III, 30: 798-818. — . 1911. Etat actuel de la question des Sporotrichoses — ^Les progres accomplis — Les discus- sions botaniques, interet pratique, pronostique, therapeutique et eeonomique. Interet doctrinal des sporotrichoses, Arch. Derm. Syphilis 110: 25-74. — . 1911. Les SporotricJuim pathogenes. Classification botanique, Arch, de Parasitol. 15: 5-109, Pis. 1-5. SPOKOTRR'HEAE 813 * — . 1912. L' etat actuel do la question des luycoses, Biol. Med. 10. — . 1912. Les Sporotrichoses. Faris, YeVix Alcan. S25 :pp., 8 pis., 181 figs. Beunnann, Liicien de, H. Gougerot & Vaucher. 1907. Notes sur les sporotrichoses generalisees experinientales . . . B^dl. Mem. Soc. Med. Hop. Pans III, 24: 1000-1008. — . 1907. Note sur 1 'histologie des f ollicules sporotrichosiques experimentaux, Bull. Mem. Soc. Med. Hop. Paris III, 24: 1009-1013. — . 1908. Epi to his Aspergilhis roseus. Hence, eitlier A. conspersum or A. microsperum must be chosen as the type. Since the former was evidently the best known, it should be taken as the type. Bonorden, Saccardo and Lindau have followed this concept. Hyphae repent; conidiophores erect, unbranched, septate or not; conidia lateral, sessile, unicellular, or hyaline. A single pathogenic species has been referred here. It has been so little studied morphologically that its position here is uncertain. Acladium Castellanii Pinoy apud Castellani, Brit. Med. Jour. 2; 486, 1 pL, 1916. Pseudomicrosporon Castellanii Craik, Jour. Trop. Med. Hyg. 26: 184, 185, 1923. MISCELLANEOUS FUNGI IMPERFECTI 837 Found in Ceylon, Malay States, and Macedonia in small lesions usually diagnosed as syphilitic. Ulcers sharply defined, roundish or oval, with red granulating fundus. The purulent secretion dries up in thick yellow crusts covering the ulcers. Gummatous nodules and furuncular lesions may also be observed. Little or no pain, pruritus usually absent. Wassermann negative. Mercury and arsenic compounds have no effect. Lesions promptly cured by medication with KI (20 gm. t.i.d.). Hyphae 2/x in diameter with pseudoconidia of varying shape — cylindric, pyriform, or spherical — -attenuate at point of insertion. Pseudoconidia, 4 x 2/A. Occasionally chlamydospores in short chains, spherical, often terminal, 8-lOyu in diameter (Fig. 131). Organism grows on Sabouraud agar, glucose agar, carrot or potato. Colo- nies on carrot or potato whitish, covered with spiculated formations consisting of straight parallel filaments. On glucose agar, after 4-8 days, colonies small, amber yellow, becoming hemispheric and coalescing into a knotty mass. If colonies do not fuse, they increase and show radiating furrows. Colonies dimorphic, smooth (mostly below 20° C. in malt agar). Oidia and chlamydospores present; rough, with coremia bearing hyaline, pyriform, sessile conidia, 2 x S/j. — Craik (1923). MONOSPORIUM Monosporium Bonorden, Handb. allg. Mykol. 95, 1851. The type species is Monosporium agaricinum Bonorden. Bonorden described his genus very briefly as possessing branched hyphae bearing round or ovoid spores on the tips of the branches, the branches not regu- larly divided. He treats 15 species and mentions 2 others. M. corticola, M. agaricinum, M. spinosum (Fig. 132), M. memhranaceum, M. decumhens, M. viri- descens, M. reflexum, and M. acuminatum are described as new, most of the others evidently known to the author only from figures. The figures of M. acuminatum, M. spinosum, M. agaricinum, and M. memhranaceum resemble each other much more closely than they do the other members of Bonorden 's genus. Within this group, the choice of type species is arbitrary. Hyphae repent, septate, branched; conidiophores erect; septate or not, more or less dendroid, branched, branching often dichotomous ; branches usu- ally tapering to a point, bearing a single hyaline, smooth, unicellular, thin- walled, ovoid conidium. In the pathogenic species, the conidiophores are somewhat decumbent and are said to have been transferred by Saccardo to a new genus, Scedo- sporium, but I have been unable to find where this name was validly published. Saccardo did not recognize this name in later volumes of his Sylloge Fungorum. All of the pathogenic species so far reported have produced black grained mycetoma pedis. 838 MEDICAL MYCOLOGY Monosporium apiospermum Saccardo, Ann. Myc. 9: 254, 1911; Kadaeli, Giorn. Ital. Mai. Yen. Pelle 52: 109-116, 1 pL, 1911. Scedosporium apiospermum Saccardo, 1914. Aleurisma apiospermum Maire apud Montpellier & Gouillon, BuU. Soc. Path. Exot. 14: 285-290, 1921. Indiella americana Delamare & Gatti, C. R. Acad. Sci. April, 1929 ; fide Pena, Rev. Med. Cirurg-. Brasil 38: 142-147, 1 fig., 1930, who studied their culture. Isolated from cutaneous granuloma of the human foot, North America, Brazil, and Europe. Fig. 132. — Monosporium spinosuyn Bonorden. (After Saccardo.) Hyphae white, then slightly fuscous, cottony ; conidiophores not erect ; vaguely and sparingly branched, sparingly septate, 2.5-3yu,, branches ascending, slightly attenuate, terminated by a single conidium each ; conidia unicellular, pyriform, oblong or obovoid, truncate at base, 11 x 5.6-5.7/t, rarely almost spherical, guttulate, smooth, hyaline at first becoming light dirty rose yellow. Selerotia abundant in tissues of host and on certain culture media (Pig. 133). Colonies on Sabouraud agar after 5 days, of lentil size, raised, covered with a white velvet, surrounded on the eighth day by a circular furrow, center finally pale chamois in color with remainder a brown yellow. On potato growth same as above, but more luxuriant, substrate blackening on fifth day. On carrot the same, carrot blackening in 10 days. Growth on com, barley, or oats white, cottony, becoming mouse gray and arachnoid. Similar growth on bread crumbs, but deeper part greenish. Same, but less luxuriant, on MISCELLANEOUS FUNGI IMPERFECTI 839 beans or onions. In hay infusion (20%), slightly acid, hemispheric growth at bottom of tube on fifth day. Same in Sabouraud maltose broth, peptone glu- cose broth, potato decoction, or carrot decoction. Interesting cases have been reported by Montpellier (1921, 1924), Lin- hares (1917), Magalhaes (1919), Fonseca & Area Leao (1927), and Gay & I Fig. 133. — Monosporium apiospermum Saccaxdo. 1, 2, pyriform conidia ; S, i, crescent or fusiform conidia; 5-12, chlamydospores in various stages of development. (After Fonseca & ArSa Leao 1927.) Bigelow (1930). Unpublished observations of R. F. Smart in my laboratory indicate suggestions of copulation and sexuality but, unfortunately, Smart's studies were interrupted before completion. Sclerotia were very abundant in corn meal agar. 840 MEDICAL MYCOLOGY Var. Peperei Sartory, Champ. Paras. Homme Aijim. 681, 682, 1922. Monosporium sclerotiale Pepere, A. Soc. fr i Cultori di Sci. Med. e Natural! i Cagliari 18 juin 1914; Sperimentale 68: 543, 1914. Scedosporium sclerotiale Brumpt, Precis Parasitol. 1114, 1922. Isolated from a black grain mycetoma, grains 1-2 mm. in diameter. Path- ogenic to laboratory animals. Hyphae slightly brownish, rarely septate, 3-4/a, some 5-5. 5/x, branched, sometimes united into strands. Conidiophores slender and short, erect or de- cumbent, bearing a single terminal conidium, 10-14 x 4-7/1,, sometimes [young- er?], spherical or ovoid and smaller, 5-7/a, point of insertion yellowish; conidia with oil globules and opaque brown granules. Very rarely in old cultures, 2-3 spores appear at the end of a single conidiophore. Sclerotia common in old cultures. Growth under anaerobic conditions very slow. In very old drying out cultures, anaerobic cultures and colonies in collodion sacs in the peritoneum or under the skin of rabbit or guinea pig, sclerotioid growths are produced. Hyphal cells swollen somewhat irregular, contorted, having a pseudo- parenchyma without and more normal hyphae within; conidiophores abnor- mal, spores nearly round. Also coremia present at the periphery of the colony. On simple agar, colonies develop sloAvly, transparent, then white, slightly elevated, confluent. On Sabouraud agar, growth good on either glucose or maltose, colony round, center hemispheric, translucent or glassy, finally cov- ered with hyphae. Central elevation becomes irregular, mammillate, sur- rounded by a ridge, giving a somewhat crateriform appearance and ridge surrounded by a furrow, not blackening. On potato or potato glycerol, small white colonies with rapid groAvth, hemispheric with long cottony hyphae, in 6-7 days surface covered with a gray or gray brown coating with tendency to greenish or black. Old colonies completely convoluted, losing their cottony appearance. On beets, colonies white, much the appearance of potato. On banana, growth slow, colonies umbilicate, remaining white, although in old cultures brown guttation may appear. On gelatin, development not very good at 20° C. ; liquefaction slow and scarce. On serum or serum glycerol, colonies oval, central portion mammillate, hemispheric, translucent, glassy. In broth, with glycerol or glucose, colonies translucent, yellowish white, opaque, adherent, confluent, folded, then brownish; filaments from the margin climb the walls of the tube. On hay infusion + agar 2%, development rapid, colo- nies round, convex, with rapid formation of hyphae. On hay infusion, colonies spherical, mucilaginous, center dark, finally producing a white pulverulent pellicle which easily breaks up and falls to the bottom. On potato decoction, about the same. In beef broth, development slow, small colonies yellowish, tending to brown, surface granular, finely mammillate, adherent to the tube, with aerial colonies convex, white, or grayish white. In old cultures (30-35 days), colonies form a long, blackish cylinder, finally reaching the bottom of the tube, broth clear. Presence of sugars does not modify development. On peptone water, development slow, small white colonies adherent to the walls and finally making a thin white pellicle. MISCELLANEOUS FUNGI IMPERFECTI 841 Monosporimn Magalhaesi (Froes) Dodge, n. comb. Scedosporiuni Magalhaesi Froes, Do Mycetoma Pedis no Brazil 49, 1930 [name only] . Scedosporiuni sp. P. S. Magalhaes, Hyphomycetoma Nova Mycose. Rio de Janeiro, 1919. Monosporium tulanense (Castellani) Agostini, Jour. Trop. Med. Hyg. 35: 266-269, 5' Jigs., 1932. Blast omycoides iulanensis Castellani, Amer. Med. 32: 292, 1928; Amer. Jour. Trop. Med. 8: 386, 387, 1928. Aleurisma tulanense Ota & Kawatsure, Arch. Derm. Syphilis 169: 156-160, 1933. Isolated by Castellani from "blastomycosis" in Louisiana, organism studied by Agostini (1932). Pathogenicity unknown. Hyphae 1-2/x in diameter, branching, often in fascicles ; conidial mycelium 4-5/t in diameter ; conidia pyrif orm, 5-7 x 3-5//, ; arthrospores present ; chlamy- dospores 8-15ja, thick-Avalled, often with fat globules [mistaken by Castellani for ascospores] ; found both on media and in host tissues. On Pollacci agar, colonies white, adherent, slightly fluffy. No growth on blood agar. In liquid media, colonies coalescing into a mucilaginous mass. Milk not coagulated, gelatin and serum not liquefied at first but later some liquefaction takes place. No fermentation of any carbohydrates. var. mannitolfermentans Castellani, Jour. Trop. Med. Hyg. 36: 306, 1933. Differs from the species producing acidity in mannitol but not in other carbohydrates. Monosporium venezuelense Castellani, Jour. Trop. Med. Hyg. 36: 307, 1933. Isolated from a case of blastomycosis in Venezuela and from one in Cen- tral America. Conidia ovoid, 7 x ^AAfx ; sterigmata up to 10.5/i. Colonies becoming dark brown or even black on glucose agar. Gelatin rapidly liquefied. VERTICILLIEAE Mycelium hyaline or light colored, conidiophores differentiated, branched, conidia borne in verticils. Key to Genera Conidiophores sterile on ends of main brandies, conidia borne singly on short flask-shaped side branches. Pachyhasium. Conidiophores not sterile on ends of main branches. Conidia and branches not surrounded bj- a gel, borne singly. Conidia spherical, ellipsoid, ovoid, clavate, neither cylindric nor elongate. Conidia single on tips of branches, soon falling, not clavate. Verticillium. Conidia clavate, single, terminal, fertile branches ending in two clavate cells, which are perpendicular to each other. Ferticilliopsis. Conidia ?)4 in a group at tips of branches. Cladoiotryum. 842 MEDICAL MYCOLOGY Conidia cylindric or fusiform, elongate. Conidia acrogenous, single. Acrocylindriwrn. Conidia several on the tips of the branches. Tips of branches swollen. Calcarisporium. Tips of branches uncinate. TJncigera. Tips of branches neither swollen nor uncinate, sterigmata in a single row on one side. Coemansia. Conidia and branches more or less surrounded by a gel. Conidiophores several times verticillately branched. Acrostalagmus. Conidiophores with an unbranched main axis on which the small side branches are borne. Branches perpendicular, like sterigmata, simple with a small head of conidia. Harziella. Branches short, ovoid, conidia single. Gloiosphaera. Conidia borne in chains (sec also Penieillimv, etc.) Spicaria. Fig. 134. — Verticillium agaricinum (Link) Corda. (After Harz. ) No clear-cut pathogens are yet known from this group. Verticillium is the only common contaminant (Fig. 134). The position of Spicaria is not clear. Some have considered Scopulario'psis a synonym, in which case it should be transferred to the Aspergillaceae. Some species were originally described as Penicillium. Many of the species have not been adequately figured, so that one cannot be sure that the spore chain is formed in the same way as that of the Aspergillaceae, but it is very likely that such is the case. For the present, we have retained it as a separate genus and are treating Spicaria rubra from corneal lesions at this point. MISCELLANEOUS FUNGI IMPERFECTI 843 Spicaria rubra (Baquis) Dodge, n. comb. Verticillium ruhrum Baquis, Ann. di Ottabnol. 34: 945, 946, 1905, Isolated from lesions in cornea, keratomycosis. Not pathogenic to experi- mental animals. Chains of ellipsoid spores hyaline, 5 x 15/* in diameter, arranged in ver- ticils. Colony whitish velvety, becoming rose (color of peach blossoms). HAPLOGRAPHIEAE Mycelinm well developed, conidiophores differentiated; richly branched; conidia dark colored, in chains on tips of branches. Key to Genera Conidiophores erect, septate, each bearing a chain of conidia at its end. Catenularia. Conidiophores branched at the tip, each branch bearing a chain of conidia. Eaplographiitm. Conidiophores bearing more or less branched chains of conidia. Conidiophores dendroid, branched; conidia spherical or ovoid. Hormodendron. Conidiophores ending in dichotomously branched conidial chains; conidia cylindric, Hormiactella. CATENULARIA Catenularia Grove in Saccardo, Syll. Fung. 4: 303, 1886. The type species is Psilonia atra Corda. Conidiophores erect, septate, each bearing a chain of conidia at the tip; conidia brown, unicellular. Catenularia fuli^nea Saito, Jour. Coll. Sci. Imp. Univ. Tokyo 18: 5: 51, PI. 2, Fig. 4, 1904. Reported pathogenic by Joseph, Derm. Woch. 87: 1396-1398, 1928. Iso- lated from excoriated plaques with crusts on hands of boy. Intracutaneous injection produced small abscesses in mouse, from which the organism was recovered. Lesions were experimentally reproduced on skin of man. Mycelium dirty greenish color, with pleurogenous spherical spores, 4/x, in diameter. Chlamydospores intercalary, up to 40/* in diameter, with spore chains of elongate spores in hanging drops. On maltose agar, gray-brown colonies, becoming cacao brown. This species is too poorly described to place definitely in this genus. One suspects that it might have been referred to Dematium with greater propriety. The author may have been misled by the usual figure of Catenularia atra, which shows free conidia in juxtaposition to the conidiophore, so that they might be mistaken for lateral conidia. 844 MEDICAL MYCOLOGY HAPLOGRAPHIUM Haplographiuni Berkeley & Broome, Ann. Mag. Nat. Hist. Ill, 3 : 360, 1859. The type species is Haplograpliium deUcatum Berkeley & Broome. Hypliae repent, often not seen ; conidiophores erect, not branched, septate, brown, branching at the tip, each branch bearing a chain of spores, spherical or ellipsoid, green, brown, or almost hyaline, unicellular. This seems to be a black analogue of Penicillium, but it has not been sufficiently investigated in the saprophytic species to know whether the termi- nal branches are metulae and phialides or not (Fig. 135). Apparently these are in the pathogenic species. Haplographium DeBella-Marengoi Pollacci, Atti 1st. Bot. R. Univ. Pavia 18: 125, 126, PI. 30, 1921. [Case history, pathology, and cultural characters Fig-. 135. — Haplographium chlorocephalum (Fresenius) Grove. (After Saccardo.) given by deBella and Marengo, in Giorn. Ital. Mai. Ven. Pelle 63: 690-697, 1922.] Isolated from a gummatous cutaneous lesion on the jaw. Pathogenic to guinea pig and rabbit. Mycelium septate, abundantly branched, at first hyaline, then brown. Sterile hyphae repent, branched, hyaline, then brown, SAfi in diameter ; conidio- phores repent or erect, simple septate, black, 50-80/1. long, little or much branched above, phialides 10-12/x long, ending in chains. Gonidia spherical or ovoid, black, smooth, 4-5/* in diameter, ends pointed. Colony hemispheric, dirty white, then pea green with lanuginous center, then very dark grayish green. After 30 days, colony black, dry, rugose; reverse dirty white and then black. On glucose agar, colony black, zonate, orbiculate. MISCELLANEOUS FUNGI IMPEKFECTI 845 Var. equinum Pollacci, Riv. Biol. 10: 358-367, 1928; Bolognesi & Chiurco, Micosi Chirurgiche, 904, 1927. Isolated from cancro del feltone. Differs from the species in host, conidiophores longer, eonidia ovoid, 4.3 X 7.2/.. Pollacci thinks the species of Penicillium, imperfectly described by Rebaudi & Podesta (1922), belongs in Haplographium. It certainly is not a Penicillium. Var. pulmonale Redaelli, in Bolognesi, Atti 1st. Lombardo Sci. e Lett. 1925; Bolognesi & Chiurco, Micosi. Chinirg. 905, 906, 1927. Spores ellipsoid, slightly apiculate or rounded, sometimes with lateral de- formations; epispore thick, fuscous, 2.5-4.5 x 3.5-4/x. Differs from species with shorter branches bearing the chains and insertion of conidiophores, smaller ellipsoid eonidia. On carrot, colonies rounded, irregularly mammillate, elevated, dusty, velvety, grayish olive green, later confluent and darkening. HORMODENDRON Hormodendron Bonorden, Handb. allg. Mykol. 76, 1851. The type species was not designated. Since Bonorden mentions the habitat of Hormodendron olivaceum, it is probable that he had seen a specimen, while in the case of the others, he apparently knew them only from Corda'"s figures. Hence H. olivaceum may be taken as the type (Fig. 136). Hyphae repent, branched, septate. Conidiophores erect, septate, brown, branched. Conidial chains acrogenous on the branches; eonidia spherical or ovoid, olive green or brown, unicellular. In contrast to chains of spores in Penicillinni, cell division occurs simul- taneously throughout the greater portion of the branch ; the cells round up and produce spores. Several pathogenic species have been reported. They make a Avell-marked group, quite distinct from other pathogenic genera. Whether or not the reference to Hormodendron is correct is still uncertain. Hormodendron alg-eriensis ]\Iontpellier & Catanei, Ann. Derm. Syphiligr. VI, 8: 626-635, 1927. Found in lesions somewhat resembling those of sporotrichosis. On inocu- lation to rabbit, caused voluminous abscess. Sterile hyphae brown, 4/. in diameter, septate. Sporophores erect, bear- ing little chains of spores. Cell wall with little thickenings or tuberosities from which spores are borne. Spores ovoid to elongate, 5.5-11 x 3-4/a, with wall thickened at point of attachment. Colonies brown. Hormodendron madagascarensis (Verdun) Dodge, n. comb. Cladosporium sp. Gueg-uen, C. R. Acad. Sci. Paris 152: 412, 413, 1911. Cladosporimn madagascarensis Verdun, Precis Parasitol. 1912. Hormodendron sp. Langeron, Bull. Soc. Path. Exot. 15: 443, 1922. 846 MEDICAL MYCOLOGY Isolated from ulcers on the leg of a Malgache, twenty-eight years old, which developed after a bath in a brook. The primary lesion healed promptly but, after a forced march, the leg broke out into confluent nodosities followed by ulcerous tumors, from which oozed a seropurulent liquid with a fetid odor. The whole area between foot and thigh was invaded. Aseptic puncture of fresh tumors gave a blackish bloody liquid from which, after 10 days, the organism was regularly recovered. In pus and in the hypertrophied connec- tive tissue were numerous ovoid bodies, 3-4/t long, staining intense violet with Giemsa stain. Pathogenic to guinea pig and white mouse. Medication with KI ineffective. Mycelium chocolate brown, pulverulent, plane becoming cerebrif orm ; on carrot heaping up 1 cm. or more at point of inoculation. In hanging drop, the mycelium germinates oidium-like after 4 days, forming arbuscles of ovoid elements which decrease in size from base to summit of branches; later form- Fig. 136. — Hormodendron olivaceum Corda. (After Corda 1839.) ing elongated conidiophores. On slices of carrot, oidia are almost at once replaced by hyphae, cylindric, flexuous, branched, tending to become verticil- late, with the ultimate branches more or less breaking up to resemble oidia, reminding one of Cladosporium penicillioides. Oidia of first filaments germi- nating, 3-10 X 3-4/A. On solid media, cells of hyphae measure 2.5-3 x 15-25/*. Conidiiform cells of the long chains measure 3-4 x 2-4/i,. Hormodendron Fontojmonti Langeron apud Brumpt, Precis Parasitol. ed. 2, 1913; fide Langeron, Bull. Soc. Path. Exot. 15: 436-443, 1922; footnote Fontoynont & Carougeau, Bull. Soc. Path. Exot. 15: 424-435, 1922. Found in hodi potsy or parasitic achromia in Madagascar. Mycelium brown, septate, thick-walled, 2.5-7.8/* in diameter (mean 4/* on carrot). Sporophores well differentiated, dendroid. Chains of blastospores arise on tubercles or sides of last joint of conidiophore, the lower member of MISCELLANEOUS FUNGI IMPERFEUTI 847 chain elongates and gives rise to chains of spores, easily breaking up into single cells. Blastospores ovoid, more or less elongate, unicellular or septate with a single septum, thick-walled, thickened at disjunctors, giving a some- what eitriform appearance, 5 x 3-8 x 4fi (Fig. 137). Grows on usual media at room temperature, must be transferred in 15 days. Hormodendron Langeroni Fonseca, Area Leao & Nogueiro Penido, Sciencia Medica 5: 563-580, 2 pis., 1927. Found in ulcero-nodular mycosis. On the lower part of the internal side of the right leg, and oval ulceration, about 3 cm. long, with sharply defined Fig. 137. — Hormodendron Fontoynonti. (After Fraga 1930.) borders, in some places perpendicular to the skin surface, in other places the skin being more or less detached. The bottom of the ulcer is irregular, of dark red color, displaying a considerable purulent, malodorous secretion, the tissues under the ulcer edematous and infiltrated. Three small subdermic nodules noted, following a more or less straight line over the ulcer. These nodules are very painful. AVassermann positive. Clinically close to cutaneous lymphangitic type of sporotrichosis. Aerial mycelium of septate, undulating hyphae, 2-4/^ in diameter, some- times anastomosing, sometimes with crystal deposits; conidiophores usually more or less branched, the branches composed of cylindric or elongated ovoid 848 MEDICAL MYCOLOGY cells which are easily dissociated. The apical cell of the conidiophore, 7-12 X 3-4/A, bears a number of excrescences formed by a localized thickening of the wall. These excrescences bear cells in fascicles or whorls, which are clavate to more or less cylindric and produce chains of cells. The terminal cells of the chains, by repeated cell division, increase the length of the chains, the youngest spores at the ends. The typical spores are ovoid, 7-8 x 4-5/^ (2.5-14 X 2.5-6/a), or navicular, both poles being provided with disjunctors, or one may be rounded. Young colonies greenish, becoming dark green to black on most media. Growth smooth and moist, becoming cracked on drying out, sometimes covered with a thin layer of aerial hyphae. Colony form varies on different media, being subconic or crateriform on potato, smooth and confluent on carrot, often with radial furrows on agar. For further details, the original paper should be consulted. Hormodendron leproides (Leger & Nogue) Dodge, n. comb. Scopulariopsis leproides Leger & Nogue, Bull. Soc. Path. Exot. 15: 654- 661, 1922. Isolated from tAvo patients with mild lesions resembling leprosy, but with- out anesthesia. Mycelium never a coremium. Hyphae 2.5-4/x in diameter, regularly cylin- dric, old branches tangled. Septa prominent, 10-15/* apart, branching without regular alternation, often at a 25° angle. Conidia easily scattered on simple or divided sporophores, 20-60/a long. Basal conidium ovoid, retained for a rather long time by a very loose pedicel. These conidial mother cells give rise to basipetal chains of varying lengths. False branchings may be alter- nate or verticillate but never as large as the aspergillar head. Phialides not seen. Conidia rounded or citriform, each carrying as ornamentations two slight elevations on the longest axis where chains break apart, 4-6/a in di- ameter. Membrane thick, nucleus easily stained. Hormodendron rossicum Meriin, Arch. Derm. Syphilis 162: 300-310, 5 figs., 1930. Case by I. I. Chemiavski [Tschernjawski], Arch. Derm. Syphilis 157: 196-206, 1929. Found in dermatitis verrucosa, therapeusis effected by excision and medi- cation with KI. Antigens not specific. Pathogenic to rats, injections causing ulcers which show an infiltration of diffuse mononucleate cells and newly formed vessels. In the center, there is an increase of giant cells and poly- morphonuclears, with masses of spherical brown cells of the fungus which was reisolated from the experimental lesions. Mycelium 2.6yu in diameter, terminal cells (conidiophores) 2.6-3.4 x 2.6/1, spherical or ovoid. Conidia ovoid, outer conidia smaller. Conidia in rosette of 3 or 4 short chains of 2-6 members on ends of conidiophores which are only the free ends of hyphal branches ; pseudosclerotia formed. Original pus showed brown spherical cells ; does not reproduce by budding. MISCELLANEOUS FUNGI IMPERFECTI 849 Colonies on Sabouraud agar black, spherical, ovoid or irregular cells of deep greenish color, without mycelium. In subcultures, colonies are circular with raised center, black with a mouse gray velvet of young mycelium. Fasci- cles of hyphae appear. As culture becomes older, color becomes grayer, sur- face velvety and grooved. Growth good on bread or sugar agar, poor on rice, also on serum, where hyphae are not formed. Growth best at 37° C, very slow at room temperature. Growth good on the following medium: Peptone 2 gm., agar 2 gm., honey 8 gm., water 100 c.c. No gas on sugars. Slight acidity with sucrose, mannite, dulcite, galactose, maltose, glucose, or starch. Milk not coagulated, gelatin liquefied. Doubtful Species Oidium coeruleus cuticularis Greco, Argentina Med. 5 pis., 1909; Origine des Tumeurs 54-63, Figs. 11-14, 1916. Found on a young shepherd who was engaged in dipping sheep to cure them of the scab. Lesion began as red spots of desquamation, preceded by small vesicles arranged in the arc of a circle, gradually spreading to the fore- arms, pruriginous. Surface erythematous, red or somewhat lilac colored, slightly desquamatous, formed by the confluence of plaques still smaller, united more or less tangentially by their edges which are shown by the curved lines of small gray crusts, very adherent to a red surface, slightly redder than the central portions of the plaques covered with small furfuraceous grayish white scales. Cells spherical or ovoid, about 6/x in diameter, forming filaments 8-10 x 4/*, in 24 hours, which elongate to 120)U long by 1/a thick. Cells 5-6/^ long. Fila- ments variously branching, spores terminal on branches, with whole filament later breaking up into arthrospores. On Sabouraud glucose, growth is visible in 24 hours, colonies at 8 days 1 cm. in diameter, elevations 2-3 mm., color olive green, umbilicate with ele- vated margin, finally becoming dark green, almost black ; dry and separating from the medium with difficulty. Growth on Sabouraud maltose much the same as on Sabouraud glucose. Colonies on potato glycerol or plain potato abundantly cover the surface with a grayish, greenish olive color, powdered with black spores ; liquid filled Avith flocculent floating colonies, with grayish olive centers and white margins; liquid finally covered with dark olive green pel- licle, with a dirty gray powdery surface. Growth in broth or glycerol broth much the same as in the liquid of potato glycerol. This organism is problematical. The lesion suggests Epidermophyton while the description of colony, with black spores, suggests Hormodendron or some organism of that group. Blastomyces sp. Rudolf, Arch. Schifi:'s.-Tropenhyg. 18: 498, 1914. Found in the disease known as "Figueim" in Minas Geraes and Goyaz in Brazil (mossy foot?). Isolated from warts on back of foot causing a cauliflower papilloma. Pathogenic to monkeys and white rats. Colonies on Sabouraud agar dark brown to black, of appearance of mouse skin. 850 MEDICAL MYCOLOGY PERICONIEAE Mycelium well developed, conidiopliores differentiated, ending in a vesicle ; conidia black, not in chains. Key to the Periconieae Septa of conidiophore appearing as a black ring. Camptoum. Septa of conidiophore not as above. Conidia sessile on end of conidiophore, sterigma, if present, not highly developed. Conidia spherical to ovoid, tip of conidiophore more or less swollen. Periconia. Conidia elongate, tip of conidiophore not swollen. Conidiophore not branched. Gomphiriaria. Conidiophore branched. Synsporium. Conidia borne on highly developed sterigmata. Conidia not embedded in a gel. Stachybotrys. Conidia embedded in a gel. Gliobotrys. GOMPHINARIA GomphiiiKiria Preuss, Linnaea 24: 130, 1851. Acrotheca Auct. non Fuckel, Enum. Fung. Nassoviae 43, 1860. The type species is Gomphmaria amoena Preuss. Acrotheca was based on Acrotheca Gei Fuckel as a conidial stage of Depazea geicola. Hyphae repent, not very evident above the substrate ; conidiophore not branched, brown with simple, not swollen tip. Conidia fusiform to short cylindric, brown, several attached close together at the tip of the conidiophore, making a compact ball. Only a single parasitic species has been reported from this genus whose species are saprophytic and rather rare. Gomphinaria Pedrosoi (Biiimpt) Dodge, n. comb. Phialophora verrucosa A. Pedroso & J. M. Gomes, Bull. Soc. Med. Cir. Sao Paulo 3: 254, 1920; Gomes, Ibid. 3: 42, 43, 1 pi, 1920; Ann. Paulistas Med. Cir. 11: 53-61, Pis., 1-5, 1920; not Medlar, Mycologia 7: 200-203, 1915. Hormodendrum Pedrosoi Brumpt, Precis Parasitol. ed. 3, 1921. Acrotheca Pedrosoi Fonseca & Area Leao, C. R. Soc. Biol. 89: 762, 763, 1923. Trichosporium Pedrosoanum Ota, Jap. Jour. Derm. Urol 28: [4], 6, 1928. Found in nodular ulcers in a case suspected of leprosy at first. Warts covered by whitish crust which on removal leaves bleeding papillomatous surface. Mycelium black, septate, branched. Conidiophores slightly larger above, bearing a tuft of 3-15 ovoid or subfusiform spores more or less adhering in a gel (Fig. 138). I MISCELLANEOUS FUNGI IMPERFECT! 851 Colonies on Sabouraud agar black, smooth, penetrating the medium, craterifoi-m, later covered by a dark, ashy pubescence. On Loeffler's medium, growth slow, colonies flat, with irregular margins not penetrating the medium, no pubescence. On potato, colonies poor with long ashy tufts. Conidia pro- duction best on Dox & Czapek agar according to Fonseca. PIYALODIDYMEAE Conidia hyaline, 2-celled, elongate, ovoid, clavate, or pyriform. Only one species of Diplosporium has been reported to be pathogenic, but the following key is included since, very frequently, species of Trichothecium and Arfhrohotrys appear in cultures as contaminants. » Fig. 138. — Goniphinaria Pedrosoi. (After Langeron 1929.) Key to Genera Couidia single, not in chains. Both cells of the coniJium similar, siiiooth. Conidiophores rarely or not at all branched. Conidia typically pleurogenous Conidia arranged singly in a helix about the conidiophore. Haplariopsis. Conidia in wlioils (Fig. 139). Arthrobotrys. Conidia terminal. Conidia borne on sterigmata. Diplorhinotrichum. 852 MEDICAL MYCOLOGY I Fig. 139. — Arthrohotrys superha Corda. 1, conidium ; Z, swollen conidiophore ; 5, habit sketch. (After Corda 1839, 1840.) Fig. 140. — Trichothecium roseum Link. (After Matruchot. ) MISCELLANEOUS FUNGI IMPERFECTl 853 Conidia sessile. Conidia clavafe or pyrifoini, cells more or less equal. Conidiopliores not differentiated, conidia clavate, single. Didymopsis. Conidiopliores differentiated, conidia pyriform, either single or in small heads (Fig. 140). Trichothecmm. Conidia neither clavate nor pyriform, cells about equal. Diplosporium. Conidiophores branched in whorls as in Verticillium. Diplocladium. Conidial cells wholly dissimilar, at least the upper warted. Mycogone. Conidia in chains. Conidia formed as oidia on the conidiophore, which is small and irregularly branched. Hormiactis. Coiiidin foniH^l as cliains from the tips of conidiopliores, wliich branch in whorls. Didymocladium. DIPLOSPORIUM Diplosporium Link, in Linne Sp. PI. ed. 4 [Willdenow] 6: 1: 64, 1824; Bonorden, Handb. allg'. Mykol. 98, 1851. The type species is Diplosporium nigrescens Link. Mycelium of repent, septate, branched hyphae ; conidiophores erect, ir- regularly branched, bearing terminal, two-celled conidia. It is quite likely that considerable shifting of names will occur in this group unless action is taken at the next International Congress. Originally Link proposed this genus apparently in the sense of Cladotrichum of the cur- rent monographs, in which case the latter name should fall into synonymy with Diplosporium, and the species now placed in Diplosporium should be re- named, if it is decided to separate black-spored and hyaline-spored groups. Most modern monographers have apparently considered D. album Bonorden the type of Diplosporium. Diplosporium vaginae Nannizzi, Atti R. Accad. Fisiocrit. Siena IX, 17: 491-499, 1 Jig., 1926. Found in purulent vaginitis. Heads broadly effused, confluent, white, then pale ivory. Hyphae 3-6/x in diameter, ramulose, septate, conidiophores simple, subterete, 50-80 x 2.5-3)u,, apex a little head of conidia stuck together. Conidia oblong-elliptic, almost subfusif orm, 1-septate, constricted at septum, 15-20 x 5-6;u,, hyaline ; chlamydo- spores short, pedicellate, rarely sessile, 1-celled, adnate to a globulose or hemispheric vesicle, others bi- to pluricellular, wall thick, verruculose, 30-35/*, hyaline, sometimes in single chains (Fig. 141). Colony round, mammillate, lanuginous, white, covering surface of medium. PHAEOPHRAGMIAE Conidia ovoid, elongate, cylindric, fusiform, straight or curved, Avith two or more septa, dark colored ; occasionally single cells almost hyaline. 854 MEDICAL MYCOLOGY Key to Genera Conidia verticillate. Conidia acrogenous. Conidia subhyaline. Acrothecvum. Conidia black. Cacumisporium. Conidia acropleurogenous. Conidia curved, with the median cell longer and darker. Acrotheciella. Conidia straight, with the cells about equal. Spondylocladium. Conidia not verticillate. Conidiophores short and soft. Napicladium. Conidiophores long and quite rigid. Brachysporium. Fig. 141. — Diplosporium vaginae. (After Nannizzi 1926.) ACROTKECIUM Acrothecium Preuss, Linnaea 24: 111, 1851. f Acrothecium Corda, Icon. Fung. 2: 10, 1838 (as subgenus only). The type species is Acrothecium multisporum Preuss. Hyphae repent, not abundant. Conidiophores erect, unbranched, with a group of sterigmata at the tips, bearing a head of conidia. Conidia elongate, fusiform, dark colored or hyaline, with two or more septa. Acrotheciuin nigrum Ciferri, Ann. Parasitol. Hum. Comp. 7: 524-535, 1929. Isolated by Ochoterena from a case of black pinta in Mexico. i MISCELLANEOUS FUNGI IMPERFECTI 855 Mycelium of thick brown hyphae and hyaline slender hyphae, with inter- calary hypnospores, forming chains of vaiying lengths. Conidiophores simple or sparingly branched, erect, light brown, short, denticulate, with a black, small head, acrogenous or nearly so ; conidia rarely single, usually 2-20 in a head, 1-4-celled, usually 3-celled, pyriform, ellipsoid, ovoid, at first subclavate, subhyaline 1-celled, then septate, brown, with the middle cells darker and the end cells smaller and much lighter, base acute truncate, 16-30 x 6-ll)U,, mostly 22-26 X 8-9/x (Fig. 142). On Sabouraud agar, colonies velvety, gray to black, compact, pigment diffusing into the agar. Acrothecium obovatum Cooke & Ellis var. subcapitulatum Ashford & Ciferri, Mycologia 22: 180-185, 2 figs.. 1930. Saprophyte on human skin. Fig. 142. — Acrotliecium nigrum. 1, sprout mycelium ; Z, Conidiiferous mycelium ; 5, 5, abnormal conidia spores; i, normal conidia. (After Ciferri 1929.) SPONDYLOCLADIUM Spondylocladium Martins, Flor. Cryptog. Erlang. 355, 1817. The type species is Spondylocladnim fumosum Martius. Sterile hyphae repent, septate ; conidiophores erect, unbranched, stiff and dark colored ; conidia verticillate, several septate, brown. Spondylocladium atro-olivaceum Neves, Mem. Inst. Oswaldo Cruz 25: 323- 331. Pis. 85, 86, 1931. Isolated from small furunculoid abscesses in the middle portion of the naso-genal region, following an automobile accident. An intense inflamma- tory reaction and induration, suggesting sporotrichosis, Brazil. Conidiophores 34-55;u,, dark chestnut in color, septate, rigid, erect ; conidia clavate fusiform, ovoid or subovoid, either constricted or not in the middle, 856 MEDICAL MYCOLOGY ends acute or roimded, short pediceled or sessile, easily caducous, dark chest- nut at maturity, 7-10.5 x 2-5/^ ; sterile hyphae septate, aerial hyphae dendroid, creeping. Colony olivaceous, becoming very intense and finally black, filamentous at first, becoming glabrous in old cultures. Colonies on Sabouraud maltose have centers elevated, with radial furrows, some coremia in central portion, deep olive buff to dark olive buff, more or less zonate. On Sabouraud honey, center elevated, umbilieate, 4 radial furrows, from dark olive buff to dark olive, diffusing Saccardo's umber into the medium. On Sabouraud glucose, deep olive buff to dark olive, diffusing natal brown into the medium, reverse becoming olive brown. On potato glycerol, colonies mammillate, deep olive buff to dark olive buff, diffusing avellaneous or wood brown into medium. On carrot glycerol, growth similar to that on potato, citrine drab to deep olive, diffusing buffy brown into medium. On horse dung, small tufts, citrine drab. On rice grains, olive buff. PHAEODICTYEAE Mycelium dark colored, at least in age, conidiophores either simple my- celial branches or more highly specialized ; conidia more or less muriform, dark colored, quite variable in form. Key to Subtribes Conidiophores on differentiated hyplial branches. Conidiophores definitely differentiated. Conidia single on the tip of the conidiophore. Conidia in a head at the tip of the conidiophore. Conidia in chains or growing quite irregularly. Micronemeae. Macrosporieae. Dactylosporieae. AUernarieae. This group is largely saprophytic or parasitic on plants hence reports of human pathogens should be studied very critically before they are admitted. Considerable evidence has been accumulating that Alternaria (especially A. tenuis) is important in some cases of allergy (see Brown 1932). ALTERNARIA Alternaria Nees ab Esenbeck, System der Pilze 72, 1817. Sterile hyphae creeping, septate ; conidiophores single or in small bunches, septate not branched, short ; conidia inverted, clavate, usually with an elongate lighter tip, muriform and darker color below, mostly joined in long simple chains. The type species is Alternaria tenuis Nees ab Esenbeck. Alternaria tenuis Nees ab Esenbeck, System der Pilze 72, Fig. 68, 1817. Recently Borsook (1933) repeatedly isolated a fungus identified as Al- ternaria tenuis Nees (Cf. Elliott, John A., 1917. Taxonomic Characters of the Genera Alternaria and Macrosporium, Am. Jour. Bot. 4: 439-476, Pis. 19, 20). The lesions occurred following a splinter in the hand of a Roumanian Jewess MISCEIiLANEOUS FUNGI IMPERFECTI 857 in Canada. The lesions were dark macular, pustular and ulcerating, with scaly centers on forearm; they tended to heal spontaneously only to reappear over a period of 16 years. Macroconidia 4-8-celled with only transverse septa, or fusiform with both longitudinal and transverse septa, varying from "the size of a single leucocyte to that of 8-10." Optimum temperature 25° C, aerobic or aquatic aerobic. On Sabouraud agar, colony white, becoming brownish black in 3 weeks after conidia are produced. In subcultures conidia are produced much sooner, often in 5 days. On serum glucose agar, pellicle formed at surface. On beef broth a pellicle was produced. In the foregoing description there is little to prove identity with Alternaria tenuis Nees. The lesions suggest those produced by Hormodendron or Spondy- locladium. It is hoped that Borsook will follow his account by a more com- plete description of the organism and appropriate figures. STILBACEAE Vegetative hyphae in or upon the substrate, septate, branched, hyaline or dark colored. Fertile hyphae in parallel strands forming coremia which bear conidia at their tips. The ends of the coremia often form differentiated heads on which the conidia are borne ; in other forms the conidia are borne along the stalk as well. The stalk may be either simple or branched. The group is artificial, as coremia are formed by various groups of fungi, especially in old cultures and are often overlooked. In this case the organism would be placed among groups on the basis of its conidia without reference to coremia. In this work, organisms have been classified without reference to coremial production, but below a key to genera of Stilbaceae has been provided, together with indications of the position of various organisms treated previous were they to be treated in this family. Stysanus, bearing a brown coremium, often appears in cultures as a contaminant. Key to Genera Conidia, coremia, and hyphae hyaline or bright colored. HYALOSTILBOIDEAE. Conidia unicellular, hyaline, or bright colored. Conidia in chains. Coremimn. Conidia not in chains. Conidia bacilliforni. Clavularia. Heads not differentiated, conidia borne over whole surface. Isaria. Heads differentiated partially, conidia only on the top. Cilicio'podium. Heads differentiated. Each stalk bearing a single terminal head. Conidiophores simple. Stilbclla. Conidiophores branched. Dendrostilbella, 858 MEDICAL MYCOLOGY Stalks with lateral as well as terminal heads. Conidia in heads as in Cephalosporium. Tilachlidium. Conidial heads resembling Aspergillus. Gibellula. Conidia 2-celled, elongate, hyaline. Bidymostilbe. Conidia several-celled, elongate, hyaline. Conidia in chains. Symphyosira. Conidia not in chains. Conidia ovoid. Arthrosparmm. Conidia curved. Atractium. Coremia and hyphae dark colored, conidia hyaline or dark colored. PHAEOSTILBOIDEAE. Conidia unicellular, hyaline or dark colored. Conidia not in chains. Coremia composed of both hyaline and dark hyphae Ceratocladium. Coremial hyphae uniform. Heads solid, formed of conidiophores. Heads without spines. Sporocy'be. Heads with spines. Saccardaea. Heads formed by spreading of stalk hyphae and spores. Conidia ovoid or ellipsoid. Graphium. Conidia elongate, curved. Harpographium. Conidia in chains. Coremia with more or less definite heads or discs. Conidia spherical. Coremia with heads. Briosia. Coremia with discs. Heydenia. Conidia elongate. Antromycopsis. Coremia spreading brush or broom-shaped. Coremia fleshy. Stemmaria. Coremia not fleshy. Conidia ovoid, stalk of equal thickness throughout its length. Stysanus. Coremia fusiform, stalk thickened below. Graphiothecium. Conidia two-celled, greenish or brownish. Antroinyces. Conidia several-celled, hyaline or brownish. Coremia of agglutinated, thick-walled hyphae, dark colored. Arthrdbotrywm,. Coremia of loosely tangled hyaline or brownish hyphae. Isariopsis. Dendrostilbella Boydii Shear apud Boyd & Crutchfield, Amer. Jour. Trop. Med. 1: 258-268, 1921; Mycologia 14: 242, 1922. Coremial stage of Allescheria Boydii Shear (see p. 652). Tilachlidium Bogolepoffi Vuillemin, Bull. Soe. Myc. France 28: 113-120, PI. 6, 1912. Coremial stage of C ephalosporiiim Bogolepoffi (Vuillemin) Dodge (see p. B30). TUBERCULARIACEAE In this family are assembled forms which regularly produce conidiophores and conidia on compact cushions, known as sporodochia. The group is some- what artificial, and conidial forms vary greatly. The group contains a large MISCELLANEOUS FUNGI IMPEEFECTI 859 number of plant pathogens and saprophytes. It is doubtful whether any im- portant animal pathogens will be found here. In the four species so far reported, three are so briefly described that it is far from certain whether they belong- in this group rather than in Blast otrichum or Septocylindrium. Fusarium Moronei Curzi may have been a saprophyte accidentally present, but it has been shown to be pathogenic. This seems a true Fusarium, of which there are several hundred species described as plant pathogens. Fusarium Moronei Curzi, Atti 1st. Bot. K. Univ. Pa via IV, 1: 95-105, 2 figs., 1930; Atti 11. Accad. Naz. Lincei. Rend. CI. Sci. Fis. Mat. Nat. 11: 506- 508, 1930. From skin lesions of a dog. Mucor racemosus isolated from the same lesion. Lesions begin as subepidermal vesicles which become infiltrated. Pathogenic to white rats. Conidia falcate, end cells very attenuate ; apical cell often extended into a flagellum, the basal cell ending m a conspicuous pedicel ; typically 5- septate, 34-60 x 3.7-4.5/a, not less than 3-septate nor more than 7-septate. Sporodochia commonly tuberculiform, 0.5-2.5 mm. in diameter, pale yellow or even pseudopionnotes on the bare surface of the substrate ; or on fertile aerial hyphae either sparse or crowded. Intercalary chlamydospores always present in the mycelium, smooth or variously rugose, solitary or more often in chains or in various formless masses. Sclerotia rare, at first creamy white, then fer- ruginous, spherical, 0.5-2 mm. in diameter, composed of plectenchyma. Aerial mycelium abundant, at first white or rose color, then ochraceous, at length ferruginous from the numerous chlamydospores. No microconidia seen. Fusarium vinosum Greco, Origine des Tumeurs . . . 662-670, PI. 17, 1916. Isolated from lesions on nose, small nodular papules red yellowish lilac, with yellowish gray crusts, confluent. Rabbit died after inoculation, but author unable to isolate organism. Perhaps death caused by toxic products from disintegration of inoculum. Mycelium 2-4/x in diameter, branched, septate. Chlamydospores ovoid, up to 6/x long, or cylindric, 6 x lO/x. Conidia falciform, fusiform, 8-11 x 2-4/*, up to 7 cells. Colonies yellowish white, reverse rose, growth rapid, cottony, agar be- coming red. On Sabouraud agar, colonies radiate, red, slightly violaceous, growing to 2.5 cm. in 3 days, loose cottony, grayish white above, slightly yelloAvish at certain points. On carrot, the rose color is less pronounced and 6n potato the cottony growth is Avhiter. The organism grows, forming a thick, folded pellicle on glucose peptone broth, to which the juice of a potato has been added and filtered. This organism is too poorly described and figured to be identifiable as a Fusarium. The rather poor photomicrograph suggests Endomyces, Coccidioides or a similar organism, although no yeast cells were observed. Fusarium sp. Frei, Derm. Woch. 80 : 411-414, 1925. Patient 15-16 years old, contracted gonorrhea, which he treated at home with injections of cow's milk. Later he noticed turbid urine, 4 months before 860 MEDICAL MYCOLOGY acute gonorrheal urethritis, epididymitis, and prostatitis developed. Gon- ococcus disappeared in 14 days under the above-mentioned treatment. Ex- amination showed small white star-shaped colonies of mycelium in posterior urethra. The fungus is evidently a saprophyte as secondary invader. Therapy : AgNOg, 1-2%, applied directly to colonies with urethroscope. Spores cylindric to sickle-shaped, with or without septa, usually in groups of 2-8. Chlamydospores also present. Colonies dry, sunk in substratum, separable with difficulty, center um- bonate with radiating furrows, dull yellow to red brown, color diffusing into medium. At 37° C, little aerial mycelium; at room temperature, much aerial mycelium, which is white on gelatin. Growth good on broth, malt extract, milk, litmus milk, or urine, of star-shaped flocculi which both float and sink to form sediment. Gelatin not liquefied. Glucose and lactose not fermented. Milk peptonized. Fusariopsis Derrieni Parreiras Horta, Brasil Med. 33: 395, 1919 (nom. nudum) ; Tribuna Med. Rio de Janeiro 25: 233-256, 1919. Isolated from a case clinically diagnosed as diphtheria in Carrieu's clinic in Montpellier. False membranes and white points on uvula and left tonsil. Author unable to cultivate. Conidia fusiform, unicellular or 1-septate. Mycelium present. Fusariopsis was separated from Fusarinni solely on basis of human pathogenicity. Unknown or of Doubtful Position The following species are either too poorly described to place definitely or are known to me only by mention in secondary sources. Sarcinomyces Inkin'Oho, mentioned by Ota, Jap. Jour. Derm. Urol. 28: [4], 1928. Found on scrotum, in Formosa. Adenomyces Oruzi Dias 1917. Montoyella Bodini Castellani 1907; Castellani & Chalmers, Man. Trop. Med. ed. 2, 799, 1913. Colonies whitish or greenish. Reported by Castellani & Chalmers, 1913, from red pinta [p. 1514]. Muoor mycetomi Gelonesi, Ann. di Med. Nav. Colon. 33: 283-308, 8 figs., 1927. Isolated from a black grain mycetoma in Somaliland. When liquid from lesions was planted, tubes mostly remained sterile. Very rapid growth on banana, with "physiologic" serum at bottom of tube. Colonies ochre yellow, smooth, humid, adherent to the substrate, confluent, forming milky white, humid, dense pellicle. After two months, a white, cottony growth, 2-4 mm. above, hyphae erect, terminated by a black round little grain. The moist, mucous, waxy layer is composed of hyphae, 4-5/1 in diameter, densely interwoven, possessing granular protoplasm, regularly and frequently septate with branching both dichotomous and at various angles MISCELLANEOUS FUNGI IMPERFECTI 861 including right angle, with tormina I hypnospores. In the cottony layer hyphae 8-lOfji, hyaline, seplale, branches, sporangiophores \()-12/x, granular, rarely septate or lateral branching. Sporangium 40-5()/x long, spores spherical or ovoid, variable in size. Sporangium with black pigment granules. Columella evanescent ! N.B. In none of the other media is the first type of colony produced. BIBLIOGRAPHY Agostini, Angela. 1930. Glenosporella dermatitidis n. sp. causa di dermatomicosi umana, Atti 1st. Bot. E. Univ. Pavia IV, 2: 93-101. — . 1931. On Blastomycoides lanuginosus Castellani, Jour. Trop. Med. Hyg. 34: 287, 288, £ figs. — . 1931. Ricerclie biologielie sail' Aerostalagmus cinnabarinus Corda, Atti 1st. Bot. B. Univ. Pavi Bailey K. Ashford. 1930. A new Porto Eican species of Acremoniella, Mycologta 22: 62-68, 2 figs. Costantin & Eolland. 3889. Blastomyces, genre nouveau, Bull. Soc. Myc. France 4: 153-156, PI. 23. Craik, E. 1923. Acladium Castellanii Pinoy, Jour. Trop. Med. Eyg. 26: 184-186. Curzi, Mario. 1930. Intorno alia posizione sistematica di un Fusarium isolato dalla pelle del cane, Atti 1st. Bot. E. Univ. Paria TV, 1: 95-105, 2 figs. — . 1930. Prime osservazioni su la mutazione di un ifomicetc, Atti K. Accad. Naz. Lincei, Bend. CI. Sci. Fis. Mat. Nat. 11: 506-508. — . 1930. Su la mutazione di un ifomicete (Fusarium Moronei), Atti Secondo Congr. Naz. di Microbiol. Soc. Internas. di Microhiol. Sez. Ital. 8: 49-52. Englehardt, Willy. 1924. Uber zwei Falle von Hauterkrankungen, hervorgerufen durch ein sporotrichonartiges Monosporium, Derm. Woch. 79: 1405-1414, 11 figs. Fonseca filho, Olympio Oliveiro Eibeiro da & A. E. de Area Leao. 1923. Sur la systematique des champignons produisaut des chromoblastomycoses, C. li. Soc. Biol. 89: 762, 763. — . 1927. Sobre o Scedosporium apiospermum cogumelo productor de mycetomas na Italia e no Brasil, Bol. Inst. Brasileiro Sci. [Eio de Janeiro] 3: 24, 25. — . 1927. Sur un cas d'acladiose a Acladium Castellanii observe nu Bresil, C. B. Soc. Biol. 97: 1361, 1362. * — . 1930. Chromoblastomycose, Bol. Acad. Nac. Med. Bio de Janeiro 101: 18-40; *Medica- menta 9: 28, * — . 1930. Las cromoblastomycoses, Bev. Med. Cirurg. Brasil 38. Bexmion Soc. Argentina Pat. Begional del Norte 5: 328. Fonseca filho, Olympio Oliveiro Eibeiro da, A. E. de Area Leao &i J. C. Nogueira-Penido. 1927. Mycose de typo ulcero-nodular, semelhando a esporotrichose e produizida per uma especie de cogumelo do genero Hormodendrum, Sciencia Med. 5: 563-573, 2 pis. MISCELLANEOUS FUNGI IMPERFECTI 863 *Fonseca filho, Olympio Oliveiro Eibeiro tla & Eduardo RabcUo. 1927. Uma mycosc rara: Acladiose pelo Acladium Castollanii, A. Pathologia Geral. Fontoynont, Maurice & Humbert Boucher. 1923. Contribution a I'etude des mycoses de Madagascar, Ann. Derm. Syphiliqr. VI, 4: 209-23.'5; 318-344, 10 figs. Fontoynont, Maurice & J. Carougeau. 1922. Ktude sur le Hodi-potsy, dermatomycose mal- gache, Bull. Soc. Path. Exot. 15: 424-436. Fraga, Arminio. 1930. Sobre urn case de dermatite ulcero-nodular causada pelo Hormoden- drum Langeroni, Hev. Med. Cir. Brasil 38: 321-329, 1 pi. [English translation 330- 336]. Frei, W. 1925. Urethritis posterior chronica mycotica, Derm. Woch. 80: 411-414. Fresenius, G. 1850-1863. Beitrage zur Mykologie 1: 1-38, Pis. 1-4, 1850; 2: 39-80, Pis. 5-9, 1852. Gammel, John A. & Alan E. Moritz. 1933. Experimental monosporosis, Arch. Derm. Syphilol. 27: 100-106. Gay, Douglas M. & James B. Bigelovv. 1930. Madura foot due to Monosporium apiospermum, Amer. Jour. Path. 6: 325-335, Pi. "72. *Gomes, Jose Maria. 1920. Um novo caso de dermatite verrucosa, Bull. Soc. Med. Cir. Sao Paulo 3: 42, 43. Gougerot, H., Burnier &. J. Duche [histologie par Olga Eliascheff]. 1933. Mycose vegetante et ulcereuse due au (.'eplialosporium griseum. Bull. Soc. Frang. Derm. Syphiligr. 40: 417, 418. Gueguen, Fernand. 1911. Mycose cladosporienne de I'homme, C. E. Acad. Sci. 152: 412, 413. Hartmann, E. 1926. tJber ein gemeinsames Vorkommen von Cephalosporium und Tricho- phyton gypseum. Derm. Woch. 82: 565-569. Hopkins, J. G., Rhoda. W. Benham & Beatrice M. Kesten. 1930. Asthma due to a fungus — Alternaria, Jour. Am,. Med. Ass. 94: 6-10, 1 fig. Horta, P. Parreiras. See Parreiras Horta, P. Joseph, A. 1928. tJber Befunde von Pilzen der Gattung Dematiaceae bei oberflachlichen Hautaffectionen, Derm.. Woch. 87: 1396-1398. Kawatsure, Shuji. 1933. Tierexperimentelle Untersuchungen iiber die Erreger von sogenannten amerikanischen Blastomykosen: Scopulariopsis americana, Aleurisma tulanense and Coccidioides immitis, Arch. Derm. Syphilis 169: 173-199, 11 figs. Landrieu, Marcel. 1912. Les mycoses oeulaires. These de Paris, 136 pp. Lane, C. G. 1915. A cutaneous disease caused by a new fungus. Phialophora verrucosa, Jour. Cutan. Dis. 33: 840-846, S pis. Lang, Franz Josef. 1921. Durch Leptothrixinfektion bedingte Leberinfarcierung in einem Falle von callosem Ulcus duodeni, Arch. Path. Anat. Physiol. [Virchow] 234: 367- 377. Langeron, Maurice. 1913. Hormodendron Fontoynonti n. sp. et achromie parasitaire mal- gache, "Arch. Parasitol. 16:" [cited by Brumpt Precis de Parasitologic ed. 2, 1913, but never published due to war, therefore should be cited Langeron apud Brumpt. See Langeron, 1922, footnote.] — . 1922. Hormodendron Fontoynonti Langeron 1913, agent de 1 'achromie parasitaire mal- gache, Bull. Soc. Path. Exot. 15: 436-443. Leao, A. E. Area. See Area Leao, A. E. Leinati, Fausto. 1928. Micosi rare in animali. Osservazioni cliniche esperimentali, Atti R. Accad. Fisiocrit. Siena X, 3: 83-92, 3 pis. — . 1928. Sull'azione patogena di una nuova specie di Fusarium (F. Moronei), Biv. Biol. 10: 141-154, 10 figs. Maciel, Jesuino. 1930. ContribuiQao a historia das chromoblastomycoses brasileiras, ^er. Med. Cir. Brasil 38: 3-8, I pi. Maffei, L. 1929. Nuova specie di Cephalosporium causa di una cheratomicosi dell'uomo, Atti 1st. Bot. R. Univ. Pavia IV, 1: 183-198, 9 figs. 864 MEDICAL MYCOLOGY — . 1931. Micosi causata da una varieta di Halobyssua moniliformis Zukal, Atti 1st. Bot. K Univ. Pavia TV, 3: 10-28, 10 figs. Magalhaes, Octavio de. 1916. Aleurophora benigna n. g. n. sp., Brasil-Med. 30: 369, 370. Mantarro, Giovanni. 1931. Epidermomicosi acromizzanti da Hemispora stellata, Giorn. Ital. Derm. Sifilol. 72: 131-143. Margarinos Torres, G. 1927. Histologie pathologique de I'acladiose, C. B. Soc. Biol. 97: 1362-1364. Matruchot, Louis. 1892. Eecherches sur le developpement de quelques mucedines, These de Paris, 111 pp., 8 pis. Matruchot, Louis & Eamond. 1905. Un type nouveau de champignon pathogene chez I'homme, C. B. Soc. Biol. 57: 379, 380. Medlar, E. M. 1915. A new fungus, Phialophora verrucosa pathogenic for man, Mycologia 7: 200-203, Fig. 1. — . 1915. A cutaneous infection caused by a new fungus, Phialophora verrucosa with a study of the fungus. Jour. Med. Bes. 32: 507-521, Pis. 29-33. Mendelson, Ralph W. 1920. A case of Castellani's Acladiosis, Brit. Med. Jour. 2: 664, 1 fig. [Eeprinted in Jour. Trop. Med. Eyg. 24: 86, 1921.] Meriin, J. A. 1929. Gribok vydelennii iz sluchaia chromoblastomykoza, Mater. Mykol. Fytopat. 8: 90-100, 10 figs. — . 1930. Zur Mykologie der Chromoblastomykose (Der Erreger des europaischen Falles der Erkrankung), Arch. Derm. Syphilis 162: 300-310, 5 figs. — . 1932. Weitere Beobachtungen iiber den Erreger der europaischen Chromoblastomycosis, Arch. Derm. Syphilis 166: 722-729, 6 figs. Miller, Hiram E. & Howard Morrow. 1932. Cephalosporiosis, an unusual mycotic infection, Arch. Derm. Syphilol. 25: 294-303, 10 figs. Montpellier, J. 1922. Note biologique au sujet des mycetomes. Essais de vaccinotherapie, Bull. Soc. Path. Exot. 15: 7, 8. — . 1924. Deuxieme cas algerien de mycetome du pied ("type Pied de Madura") d clericus Dodge, 343, 345 Conori Dodge, 343, 344 Gotta (Neveu-Lemaire) Froilano de Mello, 346 guttulatu.cor Mich., 101-103, 108, 110 Aspergillus Scop., 113 hi'fi.dus Pres., 118 Gornealis V. Cav. & Sacc, 114 corymiifer Cohn, 112, 114 typica Lichtheimi Lucet & Cost., 112 criistaceus L., 641 dimorphosporus Lendn., *18 hiemalis Wehmer, *106 javanious Wehmer, 98 Mucedo L., *99, 107, 108 mycetomi Gelonesi, 860 niger Ciaglinski & Hewelke, 116 Prainii Chodat & Nechich, *20 pusillus Lindt, 111 racemosus Fres., *18, *20, *106, 110, 118, 859 racemosus Jakowski, 111 ramosus Lindt, 111, 113 Begnieri Lucet & Cost., 114 rMsopodiformis Cohn, 117 scarlatinosus Hallier, 110, 111 septatus Bezold, 117 sphaerosporus Hagen, 98 Truchisi Lucet & Cost., 114 Mucoraeeae classification, 109 key to pathogenic genera, 110 morphology, 98-109 Mucoreae, 100, 110 Mucosa, buccal, 176 Mucus, 341 Muguet, 204, 275 Mule, lymphatics, 169 skin, 169, 540 Muscles, 754 Mycelium, 17 of Trichophytoneac, *44!J Mycelohlastanon Ota, 202 albicans Ota, 274 halcardctim Ota, 249 bethaliense Ota, 231 brasiliense Ota, 157 hronchiale Ota, 254 candidum Ota, 270 CopellU Ota, 252 cutaneum Ota, 287, 288 var. Takahashi, 278 deoolorans Ota, 253 entericitm Ota, 264 enterocola Ota, 245 faecdle Ota, 254 Favrei Ota, 286 Mycelohlastanon — Cont 'd gifuense Taniguchi, 288 Gruetsii Ota, 289 Harteri Ota, 178 insolitum Ota, 258 Kartulis Ota, 260 KrU'Sei Ota, 231 linguae-pilosae Ota, 260 londineiise Ota, 252 macedonense Ota, 259 nnetalondinense Ota, 256 Nabarroi Ota, 253 mtidum Ota, 261 niveum Ota, 256 parabalcanicum, Ota, 250 parachalmersi Ota, 267 paraTcnosei Ota, 261 paratropicale Ota, 264 Pinoyi Ota, 285 pseudoalh leans Ota, 229 pseudotropicalis Ota, 259 psilose Ota, 279 pulmonare Ota, 298 roseu,in Ota, 293 rotvmdatum Ota, 220 tokioense Fujii, 235 tropicale Ota, 258 tumefaciens album Ota, 232 zeylanicum Ota, 241 Mycetoma, 355, 622, 630, 637, 652, 675, 683, 684, 687, 711, 716, 721, 726, 742, 745, 747, 750, 764, 793, 795, 840, 860 black grain, 680 pedis, 738, 739, 754, 756, 761, 810 Mycobacteriu/tn Lehm. & Neum., 269, 694, 705, 761 tuberculosis (Koch) Lehm. & Neum., 157, 254 Mycocandida Lang. & Tal., 191, 194, 207, 208, 293, 294 key to species, 293 inexpeotata Tal. & Mack., 296 mortifera (Red.) Lang. «fc Tal., 295 onychophila (Poll. & Nanu.) Lang. & Tal., 266, 293, 294 parapsilosis (Ashf.) Dodge, 293, 294 rosea (Zenoni) Dodge, 293 Shutetskyi Ota, 293 Mycoderma Pers., 178, 206, 207, 223, 463 key to species, 223, 224 nomenclature, 197, 198 asteroides Brumpt, 218 Bogolepoffi Jannin, 830 brasiliense Neveu-Lemaire, 157 brocianum Pinoy, 209 Caoi Jannin, 224, 225 cerevisiae Desmazieres, 327 concentricum Vuill., 490 cutaneum Brumpt, 211 dermatitidis Brumpt, 168 Dombrayi Vuill., 370 Gilchristi Jannin, 168 Griewanlci Neveu-Lemaire, 214 histosporocellularis Neveu-Lemaire, 179 immite Verdun & Mandoul, 149 infestans Fonseca & Area Leao, 209 I INDEX 887 Mycoderma — Cont 'd Issavi Mattl., 224, 22(5 Kieta Mattl., 224, 22G Erusci .Stovall & Bubolz, 231 lactis Desm., 287 matalensc Brumpt, 296 mesentericum Pers., 197, 223 mnltifermentans (Cast.) Dodge, 228 Muyaga Mattl., 224, 225 7hobile Sart., 223, 227 Nyaiisi Mattl., 223, 224 paranigosum (Cast. & Douglas) Dodgo, 219, 224, 225 pseudoalbicans (Neveu-Lemaire) Dodge, 229 piiilmoneum Vuill., 217 Babesalama Font. & Boucher, 23fi Boquettei Vuill., 492 rotundatum Brumjit, 220 Salcuranei Jannin, 679 suhtile (Blanch.) Verdun, 228 sulfureum (Beauverie & Lresieur) Dodge, 223, 227 vvni Grawitz, 274, 275 virulens (Norris) Dodge, 223, 224 sp. Bennett, 555 sp. Griewanki & Laveau, 214 sp. Sart. & Bailly, 795 Mycodermc LeDantec, 237 Mycogone Link, 853 Mycosis, generalized, 177, 792 Mycosphaerella Fragarme (Tul.) Kiel)., *123 Mycotorula Lang. & Tal., 272, *273 Mycotorula Will, 135, 136, 194, 207 nomenclature, 201 albicans Laiig. & Tal., 178, 274 albicans Tal. & Mack., 239 Candida Tal. & Mack., 242 craterica Will, 201 fainata Harrison, 218 interdigitalis Bed., 278 vvuris Cif. & Bed., 208 onycJiophila Ciarrocchi, 294 pulmonalis Cif. & Red., 217 radioplicata Will, 201 vesica Harrison, 236 Mycotornloides Lang. & Tal., 194, 207, 208, *290 key to species, 290, 291 nomenclature, 203 Aldoi (Pereira Filho) Lang. & Tal., 291, 292 argentina (Vivoli, Avellaneda & Bardessi) Dodge, 290, 291 triadis Lang. & Tal., 203, 290, 292 unguis (Weill & Gaudin) Lang. & Tal., 290, 291 Myxomycetes, 24 Myxotrichum Kunze, 429, 451 Myzeloblastanon (See Myceloblastanon) N Nadsonia Syd., *127, 105, 307 fulvescens (Nads. & Konokotina) Syd., *303 Bichteri Kostka, 303 Nails, 234, 239, 248, 266, 267, 291, 294, 296, 315, 440, 479, 500, 523, 524, 549, 622, 625, 627, 637, 638, 645, 647, 649, 673, 82B, 830 Naming of organisms, 75-96 Napicladviim Thiim., 854 Naso-genal region, 855 Nasopharynx, 151 Neck, 210, 278, 311, 341, 344, 486, 488, 493, 498, 646, 795, 801 Neisseria gonorrhoeae (Neisser) Trev., 337 Nematospora Peglion, *127, 130, 136 Coryli Peglion, *131 Gossypii Ashby & Nowell, *130 Nagpuri Dastur, 131 Neogeotrichum Magalhaes, 156, 157 pulmonenm Magalhaes, 157 Neomicrosporuin Sab. (subgenus), 447, 461, 537, 539, 545 N eotrichophyton Cast. & Chalm., 462 fiavum Cast. & Chalm., 529 plicatile Cast. & Chalm., 530 Nitrocellulose method for embedding fungi, 71, 72 Nitrogen requirements of fungi, 33, 34 Nocardia Trev., 704, 706 Actinomyces Trev., 730 africana Pijper & Pullinger, 754 alha Froilano de Mello & St. Antonio Fernandes, 725 albida Chalm. & Christ., 746 appendicis Chalm. & Christ., 733 asteroides Blanchard, 750 aurea Cast. & Chalm., 732 bicolor Froilano de Mello & St. Antonio Fernandes, 749 Berestneff, Chalm. & Christ., 754 bovis Blanchard, 730 brasiliensis Cast. & Chalm., 749 Bruni Chalm. & Christ., 752 buccalis Cast. & Chalm., 755 Candida Cast. & Chalm., 725 carnea Cast. & Chalm., 741 Carongeatd Cast. & Chalm., 711 Chalmersi Froilano de Mello & St. Antonio Fernandes, 734 Christophersoni Froilano de Mello & St. Antonio Fernandes, 723 Gonvoluta Chalm. & Christ., 745 cruoris Macfie & Ingram, 753 cylindracea Korte, 716 Dassonvillei (Liegard & Landrieu) Brumpt, 727 decu^sata Cast. & Chalm., 760 enteritidis Cast. & Chalm., 762 equi Chalm. & Christ., 758 farcinica Trev., 746 Foersteri Trev., 713 Foulertoni Chalm. & Ciirist., 760 fusca Cast. & Chalm., 757 Garteni Goug., 726 gedanensis Chalm. & Christ., 733 Genesii Froes, 761 goensis Froilano de Mello & St. Antonio Fernandes, 723 Gruberi Blanchard, 763 INDEX Nocardia — Cont 'd Ghwgueni Ota, 722 Eofmanni Chalm. & Christ., 7-12 hominis Cast. & Chalm., 760 indica Chalm. & Christ., 739 indicc Kanthack, 721 Israeli Cast. & Chalm., 714 Jollyi Vuill., 742 KrainsMi Chalm. & Christ., 718 Krausei Chalm. & Christ., 755 Lanfranchii Froilano de Mello & Pais, 731 Lasserrei Verdun, 763 Leishmani Chalm. & Christ., 740 lingualis Cast. & Chalm., 722 liquefaciens Cast. & Chalm., 731 londinensis Chalm. & Christ., 760 lutea Christ. & Archibald, 763 luteola Cast. & Chalm., 732 macrodipodidarum Fox, 747 Madurae Blanchard, 739 mexicana Ota, 721 minutissima Verdun, 741 mordore Chalm. & Christ., 729 Muris Froilano de Mello, 759 Nicollei Delanoe, 726 odorifera Cast. & Chalm., 763 panginensis Froilano de Mello & St. An- tonio Fernandes, 718 Pelletieri Pinoy, 764 Pijperi Cast. & Chalm., 756 Finoyi Froilano de Mello & St. Antonio Fernandes, 723 plii/ricolor Froilano de Mello & St. Antonio Fernandes, 763 pluricolor Temi, 763 Ponceti Verdun, 754 pretoriana Pijper & Pullinger, 747 pseudotuberculosis Froilano de Mello, 764 pulmonalis Cast. & Chalm., 758 putridogenes Froilano de Mello & Pais, 761 Bivierei Verdun, 721 Eogersi Froilano de Mello, 764 rosea Chalm. & Christ., 725 Bosenbachi Goug., 762 rubea Chalm. & Christ., 765 rubra Chalm. & Christ., 718 Sanfelicei Red., 751 Silberschmidti Froilano de Mello & St. An- tonio Fernandes, 711 somaliensis Chalm. & Christ., 765 tenms Cast., 715 Thibiergei Cast. & Chalm., 715 Thiryei Froilano de Mello & Pais, 729 transvalensis Pijper & Pullinger, 756 valvulae Froilano de Mello & Pais, 757 sp. Lasserre, 763 sp. Pijper, 756 Nodular organs, *450 Nodules, 233 intramuscular, 715 subcutaneous, 715 Nomenclature, botanical, 75-96 Norris' medium, 54 Nose, 151, 266, 341, 536, 801, 859 Nutrition of fungi, 32-34 O Octomyces Froilano de Mello & Gonzaga Fernandes, 163, 165, 180 Betteoicourti Froilano de Mello & Gonzaga Fernandes, 180 Etiennei (Potron) Dodge, 180 Oedocephalum Preuss, 824 Oidial formation, *20 Oidiodendron Robak, 282 Oidiomyces unguium Frei, 287 Oidiihm Link, 272, 274 nomenclature, 196 albicans Robin, 272, 274 var. Galli-Valerio, 259 var. Hasogawa, 282 aster oides Cast, & Chalm., 218 cocddioides Ophiils, 149 coeruleus cutic'ularis Greco, 849 cutaneu,7n Beurm., Goug. & Vaucher, 211 dermatitidis Ricketts, 168 epitheliomae Greco, 536 fmfur Zopf, 358 lactis Fres., 703 matalense Cast., 295 porriginis Mont., 558 protosooides Ophiils, 149 pulmoneum Bennett, 217 pulmoneumi Magalhaes, 156, 157 Quinclceanum Zopf, 556 roseum [non liquefaciens] Zenoni, 293 rotundatum Cast. & Chalm., 220 ScJioenleini Lebert, 558 Schoenleinii Zopf, 558 subtile Kotliar, 358 subtile- cv,tis Babes, 228 tonsurans Zopf, 531 tropicale Cast., 258 Vanderburgii Kohlbrugge, 244 22 Cao, 225 sp. Coley & Tracy, 299 sp. Favera, 214 sp. Miiller & Retzius, 558 sp. Sakurane, 679 sp. Spiethoff, 270 Stamm I Gentzsch, 230 Ointment, Swartz', 472 Whitfield's, 472 Oleina Tiegh., 165, 179, 180 lateralis Tiegh., *165, 179 nodosa Tiegh., 163, *165, 179 Onychogryposis, 291 Onychomycosis, 137, 143, 234, 248, 266, 267, 523, 524, 622, 625, 825, 830 (see also Nails) Onygena equina [Willd.] Pers., 653 Onygenaceae, 425, 653 Oogamy, 22 Oospora Sauvageau & Radais, 706 alba Sart., 744 alba I Sart., 761 alba var. toxique Sart., 719 algirus Sart., 682 aruierobies Sart., 717 asteroides Sauvageau & Radais, 750 Audouini var. equinum Bodin, 728 aiirea Sart., 732 bahiensis Sart., 740 INDEX 889 Oospora — Cont 'd boms Savaugeau & Eadais, T.'iO bra^iliensis Sart., 749 bronchialis Sart., 757 buccalis Roger, Bory & Sart., 755 cameli Mason, 7G2 oanis Sart., 748 caprae Sart., 748 carnca Sart., 741 catarrhalis Sart. & Bailly, 733 catemita Scliaede, 238 convoluta Sart., 745 consentrica Hanawa & Nagai, 490 cruoris Sart., 753 ouniculi Froilano de Mello & St. Antonio Fernandes, 712 cylindracea Sart., 716 d'Agatae Sacc, G49 decussatus Sart., 760 Dori Sart., 762 Doriae Sauvageau & Radais, 725 erysipeloidis Neum. & Lehm., 762 farcinica Sauvageau & Radais, 746 forme de Microsporum Bodin, 728 Forsteri Sauvageau & Radais, 714 fragilis Schaede, 238 ftisca Sart., 757 Garteni Sart., 726 gypsoides Sart., 724 Hoffmanni Sauvageau & Radais, 742 indica Kanthack, 739 Jollyi Sart., 742 lactis (Fres.) Sacc, 221 Lasserrei Sart., 763 ling%iulis Gueguen, 722 Uquefaciens Sart., 731 luteola Sart., 732 Madurae Lehm. & Neuni., 740 micetomae Sart., 738 minutissima Ridet, 741 mordore Sart., 729 tnusculorium sitis Lehm. & Neum., 711 odorifera Sart., 763 Pelletieri Thiroux & Pelletier, 764 Perieri Matr. & Antoine, 213 Ponceti Sart., 754 porriginis Sacc, 558 var. ceratophagus Sacc, 558 pvlmonalis Roger, Sart. & Bory, 758 var. acidoresistant Sart., 756 var. chromogene Sart., 752 pulmonea Sacc, 217 pntridogenes Sart., 761 Bivierei Sart., 721 rubra Wilbert, 765 Sommeri Sart., 739 Spitzi Sart., 752 spumalis Sart., 751 Thibiergei Sart., 715 Tozeuri Nicolle & Pinoy, 681 var. Brault & Masselot, 682 valvulas destr^ievs bovis Sart., 757 sp. Bachmann, 221 sp. Griitz, 802 sp. Sart., 712, 715, 719, 722, 730, 735, 756. 758, 765, 766, 795 Oospora Wallroth, 198 Organic compounds, use in sterilization, 46 Osteitis, purulent, 343 Osteoperiosteitis, 183 Otitis, catarrhal, 182 media, 351 suppurating, 766 {see also Ear) Otomyces pwptireus Wreden, 636 Otomycosis {see Ear) Ovary, 313 Ovularia SaCc, 835 Oxalis tuberosa Molin., 341 Oxygen requirements of fungi, 38, 39 Pachybasimn Sacc, 841 Paedlomyces Bainier, 642, 646 type of phialides, *614 Burci (Poll.) Thom, 642 Varioti Bainier, 642, *643 Palate, soft, 341 Palmar surfaces, 205, 480, 484, 486, 674, 788 Panki, 736 Papules, 210 Paracoccidioides Almeida, 145-147, 152, 153, 155 brasiliensis (Splendore) Almeida, 156, 179 Paraffin method for embedding fungi, 71 Paramycetoma, 711 Parasaccharomyces Beurm. & Goug., 207, 265, 266, 342 key to species, 265 nomenclature, 200 albicans Froilano de Mello & Gonzaga Fernandes, 274 Ashfordi Anderson, 279 Colardi Dodge, 265, 267 crateriformis Dodge, 265, 270 Rarteri Froilano de Mello, 178 intestinalis (Mattl.) Dodge, 265, 268 irritans (Mattl.) Dodge, 265, 268, 629 oosporoides (Zach) Dodge, 265, 266 var. GuggenJieimi Dodge, 267 paraclmlmersi (Cast. & Clialm.) Dodge, 265, 267 Sairibergeri Beurm. & Goug., 200, 265 subtilis Froilano de Mello & Gonzaga Fer- nandes, 228 Talicei Dodge, 265, 269 zeylanicus Froilano de Mello & Gonzaga Fernandes, 241 G Nye, 223 Parasitella simplex Bainier, *106 Parasitic achromia, 846 Parendomyces Queyrat & Laroche, 206, 208, 234, 238 key to species, 239 nomenclature, 200 alhus Queyrat & Laroche, 200, 238-240 asteroides Rischin, 210 Balzeri Goug. & Burnier, 212 BlancMrdi (Cast.) Dodge, 239, 243 Brocquii Beintema, 210 butantanensis (Gomes) Dodge, 239, 244 communis (Cast.) Dodge, 239, 245 enterocola (Macfie) Dodge, 239, 245 farciminosus Froilano de Mello et al., 169 Hessleris (Retger) Dodge, 239, 242 890 INDEX Pare7idomyces — Cont 'd inexorabilis (Mazza & Palamedi) Dodge, 239, 242 Krausi (Ota) Dodge, 239, 244 inacroglossiae (Cast.) Dodge, 239, 241 minor (Poll. & Nann.) Dodge, 244 periunguealis (Niiio) Dodge, 239 Perryi (Cast.) Dodge, 239, 243 pulmonalis Plaut aj)ud Mautner, 298 rugosus Ota, 219 Tokishigei Froilano de Mello & Gonzaga Fernandes, 169 vaginalis (Mazza & Los Rios) Dodge, 239, 243 Vanderhurgii (Kohlbrugge) Dodge, 239, 244 Vuillemini (Froilano de Mello & Gonzaga Fernandes) Dodge, 245 seylanicus (Cast.) Dodge, 239, 241 seylanoides (Cast, Douglas & Thompson) Dodge, 239, 240 sp. Forgues, 263 sp. Launois & Pinard, 299 Paronychia, 294, 830 Partlienogamy, 22 Parthenogenesis, 23 Pectinate organs, *450 Pedicellate chlamydospores in Trichoi^hy- toneae, *452 Pedogamy, 23 Pelvis, 149 Penicillimn Lint, 639, 642, 644, 826, 842 Bertai Tal. & Mack., 639, *640 Brefeldianmn Shear, *616 brevicaule Sacc, 643, 645 var. hominis Brumpt & Lang., 649 Burci Poll., 642, 798 Camemherti Thorn, conidial stage, *615 caseicolum Bainier, conidial stage, *615 crustacewrn [L.] Fr., 641 epigaemn Motta, 651 expansum Link, 639 Giordanoi Vuill., 641 glauawm Giordano, 641 glaucum Link, 641 miniimim, Siebenniann, 641 Montoyai Cast. & Chalm., 642 mycetogemtrm Mantelli & Negri, 640 mycetomagenum Mantelli & Negri, 639, 640 pictor Neveu-Lemaire, 642 prv/riosum Salisbury, 642 quadrifiduTti Salisbury, 642 spiculisponovi Lehm., 642 Wortmanni Klocker, *618 Penicilliumuhnlich Bezold, 641 Penicillus, biverticillate, 614 definition, 614 monoverticillate, 614 Penis, 151, 183, 245, 370, 437, 678 Pennsylvania medium, 52 Perianal folds, 278 Pericarditis, 713 Periconia (Tode) Bon., 850 Periconieae, 667, 850 key to genera, 850 Pericystis Betts, 145 alvei Betts, 145 Perionychomycosis, 239 Perithecium, 125 Peritoneum, 320 Peritonitis, 254, 750 Perleche, 176 Pferdehefe II Kraus, 283 Phaeodictyeae, 856 key to subtribes, 856 Phaeodidymeae, 668 Phaeophragmiae, 853 key to genera, 854 Phaeoscopulariopsis Ota, 651 Bestae (Poll.) Ota, 651 Paisi (Poll.) Ota, 651 Phaeostilboideae, 858 Pharyngitis, 232, 346 Pharjragolaryngitis, 344 Pharynx, 146, 212, 228, 250, 255, 263, 267, 341, 344 Phata, 736 Phialides, definition of, 613 types of, *614 Phialophora Thaxter, 833 verrucosa P'edroso & Gomes, 850 verucosa Thaxter, *833 Phialophoreae, 667, 833 Phototropism, 40 Phragmospore, 453 Phthisis, 143, 225 (see also Tuberculosis) Phycomyces Kunze, 101, 108 nitens [Agardh] Kunze, *100, 107, 108 Phycomycetes, 24, 25 Phymatotrichiim Bon., 835 Physiology of fungi, 31-40 Physospora Fr., 835 Pichia Hansen, *127, 136 Pichiaceae, 126, 127, 136 Piedraia Fonseca & Area Leao, 128, 130, 131 colombiana Dodge, 133 Eortai (Brumpt) Fonseca & Area Leao, 128, *129, 132 Sarmentoi Pereira f., 132 surinamensis Dodge, 133 venezuelensis Brumpt & Lang., 128, 133 Pilaira anomala (Cesati) Schroeter, 99 Pillars of fauces, 146, 263, 281, 341, 344 Piloholus Tode, 100, 130 crystallinus (Wigger) Tode, 101 microsporws (Klein) Bref., 101 oedipiis Mont., 101 roridus (Bolton) Pers. ex Fr., 99 Pinoyella Cast. & Chalm., 457, 476 simii Cast. & Chalm., 476, 477 Pinta, 642, 676 black, 855 red, 298 Piptooephalis Bary, 101, 104, 107 Freseniana Bary, 98 Pirella circinans Bainier, *99 Pityriasis, 541 alba, 369, 370 circinata, 361, 760 flava, 361, 369 marginata, 760 nigra, 678 INDEX 891 Pityriasis — Cont 'd rosea, 438, 515 sicca, 360 simplex capitis, 361 steatoides, 361 versicolor, 358, 361 versicolor flava, 628 Pityrosporum Malassez, 359 Pityrospanun Sab., 358 Cantliei Cast. So Chalm., 360 Malassesi Sab., 359 (male Cast. & Chalm., 359 pachydermatis Weidm., 370 rhinoserosum Sab., 370 Plantar surfaces, 205, 438, 439, 479, 480, 483, 674, 736, 788 Plasmogamy, 23 Plectascales, 425, 666 Plectenchyma, 18 Pleomorphism in Trichopliytoneae, 456, 457 Pleura, 181 Pleuritis, 713 Pleurococcus Beiffeli Kiichenmeister, 134 Pleurocystis Fresenii Bon., 110 Pleuropneumonia, 263 Pollacci medium, 52, 53 Polyactis Link, 835 Polymorphism, 20, 121 Polyp, 151 Pons, 167 Populus L., 160 Porrigo lupinosa, 440 Posadasia Canton, 147 capsulata Moore, 154 esferiformis Canton, 148, 149 pyriformis Moore, 155 Potassium iodide as therapeutic agent, 204 Potato decoction, Talice's, 186 Priestley 's method of staining hair and scrap- ings, 69 Proactinomyces Jensen, 705 Prosenchyma, 19 Prostate, 335, 765 Proteomyces Moses & Vianna, 206, 208, 273, 536 key to species, 208 nomenclature, 200 astermdes (Rischin) Dodge, 208, 210, *211 Balzeri (Goug. & Bumier) Dodge, 208, 211, 212, *213 hrocianum (Pinoy) Dodge, 208, 209 Brocquii (Beintema) Dodge, 208, 210 cornealis (Nannizzi) Dodge, 208, 215 cutaneus (Beurm., Goug. & Vaucher) Dodge, 211, *212 Faverae Dodge, 208, 214 goetuiis Dodge, 215 Grieivan'ki (Neveu-Lemaire) Dodge, 208, 214 infestans Moses & Vianna, 200, 208, *209 muris (Cif. & Red.) Dodge, 208 Perieri (Mat. & Antoine) Dodge, 208, 213 Protococcus ne})ulosu!i Kiitz., 327 Protomyces Unger, 101 inundatus Dangeard, 158 Protomyces — Cont 'd macrosporiis Unger, 157, *158, 159 pachydermus Thiimen, 157 Protomycetaceae, 127, 157 Pruritus, 115, 233, 235, 266, 286, 287, 339, 438, 490, 503, 506, 507, 542, 551, 789, 795, 810 ani, 497, 498 Psendococcidioides Fonseca, 147 Mazzai Fonseca, *148, 149 Pseudodysidrosis, 479, 480 Psexidomicrosporon Castellanii Craik, 836 Pseudomixis, 23 Pseudomonilia Geiger, 207, 208, 246, 295 key to species, 295 nomenclature, 200 albomargiTMta Geiger, 295 alessandrina (Panayotatou) Dodge, 295, 297 inexpectata (Mazza, Nino & Egiies) Dodge, 295, 296 matalensis (Cast.) Dodge, 295 Pseudomycodei-ma Will, 206, 207, 235, 236 key to species, 236 nomenclature, 201 fecale Dodge, 236, 237 matalense Cif., 296 Mazsae Dodge, 236, 237 Eabesalama (Font. & Boucher) Dodge, 236 Tanakae Dodge, 236, 237 vesica (Harrison) Dodge, 236 vini Will 201, 235 Pseudoparenchyma, 19 Pseudopityriasis capitis, 518 Pseudosaccharomyces Laer, 325, 341, 342, 347 Pseudosaccharomyces Bussei Bamberger, 200, 265, 266 Pseudotuberculosis (see Tuberculosis) Psilonia atra Corda, 843 Psoriasis, 115, 244, 340, 489 Pubis, 431, 437 Pubocrural region, 486 Pulliilarin Berkhout, 672-674 Jeanselmei (Lang.) Dodge, 675 pullulans (Bary) Berkhout, 674 Pus, 254, 730, 733 staining of, 70 Pustules, 210, 233, 266, 351, 486, 505, 518, 735 Pycnia, 19 Pycnospores, 19 Pyemia, 749 Pyrexia, 153, 222, 225, 231, 257, 285, 350, 651 Pyronema confluens (Pers.) Tul., *122, *123 Q Qaercus glanduUfera Blume, 288 R Rabbits, muzzle, 712 Radium, influence on fungi, 40 Rales, 356 Bamulaspora Lindr., 835 892 INDEX Eat, 751 blastomycosis, 209 bronchopneumonia, 759 generalized nodular lesions, 733 skin, 553 Rat-bite fever, 759 Recovery of organisms, 64, 65 Bedaellia Cif., 207, 289 nomenclature, 203 elegans Cif., 203, *289 Red flap, 437 Reproductive structures, 19 Respiratory tract, 204, 229, 245, 268, 269. 347 Rheumatism, 713 Rhinoceros, skin, 370 Ehinocladium asteroides Grandinetti, 802 Beurmanni Vuill., 806 var. asteroides Vuill., 802 Carougeaui Neveu-Lemaire, 804 coprogenum Sacc. & Marchal, 800 Dori Neveu-Lemaire, 762 Fonsecai Grandinetti, 801 Gougeroti Verdun, 678 guayaquilense Valenzuela, 809 indicwn Vuill., 809 Jeanselmei Grandinetti, 805 Lesnei Vuill., 809 parvulum Red., 811 SchencM Grandinetti, 806 Schenki Verdun & Mandoul, 806 Bhinosporidium Mincliin & ramtham, 147, 151 Bhinosporidiurn Kinealyi Mincliin & Fam- tham, 151 Seeheri (Wernicke) Seeber, 151, *152 Bhinotrichum Corda, 786 asteroides Verdun, 802 Be%(,rmanni Ota, 806 SchencTcii Ota, 806 Rhizoids, 18 Bhizomucor Lucet & Cost., 115 parasiticus Lucet & Cost., 115 septatus (Bezold) Lucet & Cost., 117 Bhisopus Ehrenberg, 98, 101, 110, 111, 115 key to pathogenic species, 115 arrhisus Fischer, *18 CohnU Berl. & Toni, 117 eqivirms Cost. & Lucet, 115, 117 var. annamensis Bernard, 116 niger (Ciaglinski & Hewelke) Bartli., *99, 116 nigricans Ehrenberg, 98, *106, 107, 115 Orysae Went & Gaerl., 98 parasiticus (Lucet & Cost.) Lendner, *99, 116, 118 reflexus Bainier, *99 rhizopodiformis (Cohn), Zopf, 115, 117 Bhodomyces erihbescens Apel, 272 Kochii Wettstein, 272 Bhodotorula hronchialis Lodder, 349 glutinis Harrison, 351 mucilaginosa Harrison, 353 race ruhrorugosa Lodder, 353 var. pararosea Lodder, 353 var. plicata Lodder, 353 var. sanguinea Lodder, 217 Bhodotorula — Cont 'd pallida Lodder, 209 rubra Lodder, 352 Sannei Lodder, 352 Rhopalomyces Corda, 824 Ringworm, 435, 497, 506 Rooster, 553 S SahouroAvdia Lodge (as subgenus), 445, 524, 525, 527, 528 Sabouraudites Ota & Lang., 463, 537 Sabouraudites ( Aleuramma) depauperatum Ota So Lang., 551 iris Ota & Lang., 548 lacticolor Ota & Lang., 501 persicolor Ota & Lang., 486 Viannai Ota & Lang., 431 Sahouraihdites (Aleurocloster) asteroides Ota •Sa Lang., 504 var. chibaensis Ogata, 506 Audoidni Ota & Lang., 549 caninus Ota & Lang., 509 farinulentus Ota & Lang., 502 felineus Lang. & Miloch., 542 felineus Ota & Lang., 497 flavescens Ota & Lang., 543 gallinae Ota & Lang., 555 granulosus Ota & Lang., 496 griseus Ota & Lang., 499 gypseus Ota & Lang., 430, 553 interdigitalis Ota & Lang., 481 plurizoniformis Brumpt, 484 pubescens Ota & Lang., 544 radiolatiis Ota & Lang., 505 radioplicatus Brumpt, 498 ruber Ota & Lang., 487 var. blanche Hashimoto, Irizawa & Ota, 500 scorteus Brumpt, 503 tardus Ota & Lang., 550 umbonatus Ota & Lang., 549 Urenae Ochoterena, 517 villosus Ota & Lang., 541 violaceus Ota & Lang., 554 xanthodes Ota & Lang., 544 Sabouraudites (Closteramma) equinus Ota & Lang., 540 fulvus Ota & Lang., 541 lanosus Ota & Lang., 542 Quincheanus Ota & Lang., 556 Sabouraud's conservation agar, 51, 52 glucose agar, 186 test agar, 52 Saccardaea Cavara, 858 Saccharomyces Meyen, *127, 307, 317 key to species, 317, 318 albicans Reess, 274, 275 am/inae Vuill., 317-*319 anmilatus Negroni, 304, 318, 322, *323 anomalus Hansen, 142 apiculatus Reess, 341 Balzeri Cast. & Chalm., 212 Beauveriei Froilano de Mello, 142 Blanchardi Guiart, 317, 320 Breweri Neveu-Lemaire, 338 Cantliei Cast., 360 INDEX 893 Saccharomyces — Cont 'd capillitti Oud. & Pekelharing, 359 cerevisiae Meyen, 205, 3U0, *302, 317 Chcvalieri Guill., 304 ellipsoideus Hansen, 303, 304 equi Marcone, 169 Etiennei Potron, 180 farciminosus Vuill., 169 Ferrani Froilano de Mello, 317, 319 Fresenii Schroet., 345, 351 fflutinis Carbone, 354 glutinis Colin, 351 granulatits Vuill. & Legrain, 317, *318 granulom