=) .. mas so oe 8 - = =e oe ‘ . a t : “s’ ta th ” J APN hi he be ' ye eT ea lds ot ad ‘ Ye 4 eu 2a) Oy tebe® O84 1 Vs ‘ 7 ea aie’ o4t 4 Q ghey ‘ p Agri) ist ag, etsy Bah DS ae it MF oe tet tt oe | Me " Wane “a . ' ~t e% AA, ated wad et Vs “9 mer | rat: t Mi > ed ey toh a PAT nr Trae OT et ’ 4 A i P ve i HAMA SENN TE i i! A, » J ; a af ‘ vey A cab Sais res . ” _" : ae ae BAY, a Maley yeh nek ited aaa bs uh 0 Praha \ % , ae + Pp «J HOA }y vied e et +3 * ay WF ft tated ribet hak iat att! wo t i aks Aha ye \*, ip ahh 7 . ape aen Tea 1) ae? aangh het fers \ ‘e , hi et “eh aoa, SinteG te ME DCS Me Lee TA AG 1: Ghee i? Ctl ane ey hail ane R Pale ft aThT an PO 14 | J PS ar ; Sy a AAP tg Aten HI % Pr ard at fin, h h e's 4 we atid thet ’ P| city he} nei uw ft} all } ” a Vit teey 2G ty te Yay f f ; i, aS q i) ‘ i _ 4 } + q 44 v u er Ce oe +9 a ve | Cals i das se‘ & °. . aT he ea auth } fet 1, ft PY, hi fei! ve ’ iv ted is a ala “4 yw rr ae 4 “ P ee gin! oat Phy i, 4 ad . f yd he te Sie Fare CPt Get 248 rae < wos oe tig ; : } i wen a bet ral a ry f vate a) Pt @ gia gel at i eon ties aw ites Bee, ‘ of Ay ‘ pasgee 4 ; 7. Sreaae% es * ° ; “ . ie EA ag ect Eee a eee sate ihre wee yt iB rate ae sit or art ’ he ened ~ 1S ee t. wBy art Ae a ytnet § dupes tee fe ¢ rm hy! fy. re Pe a’ Aut ~s . Pee 5 Bie) la Te Sed pagina fe down +t ast hékent ant oat a a ay moots We sh "res avd gers : tS Vee YAO Wed) Ae en he sh vw pd , pis ” ‘“ ' oie. ae aie: ’ wot My a) b> ty 4 m4 * ety mii’. ts Me. 4 ve pape y « { <.. | i § LARATIONS e -THE INTER- BAL COMMISSI N. ON ICAL (NOMENCLATURE VOLOME Klee * 1S et planes C54)... COLE. La Shapes aeeinticd by Order of de Inter 1, . 40% eanic kd PN wreetchetare ii; | iene me the Infernetional Udrmamiesian on 7 wre Dy the Inesroatanal Trustnt as Pubic: 4h ocen"s Clabe’ London, 3. W 7 . = Hy Se LT Le _ . = 4 A — ET > [y vale var © ‘ea OPINIONS AND DECLARATIONS RENDERED BY THE INTER- NATIONAL COMMISSION ON ZOOLOGICAL NOMENCLATURE VOLUME 7 Edited by FRANCIS HEMMING, C.M.G., C.B.E. Secretary to the International Commission on Zoological Nomenclature LONDON : Printed by Order of the International Trust for Zoological Nomenclature and Sold on behalf of the International Commission on Zoological Nomenclature by the International Trust at its Publications Office 41, Queen’s Gate, London, S.W.7 1954 (All rights reserved) »-\. INTERNATIONAL COMMISSION ON “Rol . ZOOLOGICAL NOMENCLATURE COMPOSITION AT THE TIME OF THE ADOPTION OF THE * OPINION ” PUBLISHED IN THE PRESENT VOLUME A. The Officers of the Commission President: Dr. Karl Jordan, Ph.D., F.R.S. (British Museum (Natural History), Zoological Museum, Tring, Herts, England). Vice-President : Dr. James L. Peters (Museum of Comparative Zoology, Cambridge, Massachusetts, U.S.A.). Secretary : Mr. Francis Hemming, C.M.G., C.B.E. (London, England). B. The Members of the Commission Class 1949 Senor Dr. Angel CABRERA (La Plata, Argentina). Mr. Francis HEMMING (London, England) (Secretary to the Commission) Dr. Karl JORDAN (British Museum (Natural History), Zoological Museum, Tring, Herts, England) (President of the Commission). Dr. Th. MorTENSEN (Universitetets Zoologiske Museum, Copenhagen, Denmark). Dr. Joseph PEARSON (Tasmanian Museum, Hobart, Tasmania, Australia). Class 1952 Senhor Dr. Afranio do AMARAL (S. Paulo, Brazil). Professor J. Chester BRADLEY (Cornell University, Ithaca, N.Y., U.S.A.). Professor Lodovico di CAporIACco (University of Parma, Italy). Professor J. R. DyMoND (University of Toronto, Canada). Dr. James L. PETERS (Museum of Comparative Zoology, Cambridge, Massdchusetis, U.S.A.) (Vice-President of the Commission). Dr. a E. Vokes (United States Geological Survey, Washington, D.C., U.S.A.). Class 1955 Professor Dr. Hilbrand BoscHMA (Rijksmuseum van Natuurlijke Historie, Leiden, The Netherlands). Dr. William Thomas CALMAN (Tayport, Fife, Scotland). Professor Teiso EsAki (Kyushu University, Fukuoka, Japan). Professor Béla HANKO (University of Debrecen, Hungary). Dr. Tadeusz JACZEwsKI (Polish Zoological Museum, Warsaw, Poland). Dr. Norman R. STOLL (Rockefeller Institute for Medical Research, Princeton, New Jersey, U.S.A.). C. Alternate Members of the Commission at the Session held in Paris in 1948 Professor Enrique BELTRAN (Instituto Mexicano de Recursos Naturales Renovables A.C., Mexico City, Mexico). Dr. Edward HINDLE (Zoological Society of London, London, England). Dr. Arturo Ricardo JorGE (Museu Bocage, Lisbon, Portugal). Professor Harold Kirsy (University ol California, Berkeley, California, U.S.A.). Dr. Henning LEMcHE (Kgl. Veterinzr- og Landbohgjskole, Zoologisk Labora- torium, Copenhagen, Denmark). Professor Kamel MANsouR (King Fouad University, Cairo, Egypt). Professor Z. P. METCALF (North Carolina State College of Agriculture and Engineering, University of North Carolina, Raleigh, North Carolina, U.S.A.). Mr. N. D. RILey (British Museum (Natural History), London, England). Professor Ragnar SpArck (Universitetets Zoologiske Museum, Copenhagen, Denmark). Professor Victor van STRAELEN (Institut Royal des Sciences Naturelles de Belgique, Bruxelles, Belgium). cay L. Usincer (University of California, Berkeley, California, IV INTERNATIONAL COMMISSION ON ZOOLOGICAL NOMENCLATURE (continued) C. The Staff of the Secretariat of the Commission Honorary Secretary : Mr. Francis Hemming, C.M.G., C.B.E. Honorary Personal Assistant to the Secretary: Mrs. M. F. W. Hemming Honorary Archivist : Mr. Francis J. Griffin, A.L.A. Administrative Officer: Mrs. S. C. Watkins, M.A. Mrs. J. H. Newman Secretariat : Mrs. Prudence Goldman | Mrs. E. M. Lewis Indexer ; Miss Joan Kelley, B.Sc. Translator : Mrs. R. H. R. Hopkin INTERNATIONAL TRUST FOR ZOOLOGICAL NOMENCLATURE Chairman : The Right Hon. Walter Elliot, C.H., M.C., F.R.S., M.P. Managing Director and Secretary: Mr. Francis Hemming, C.M.G., C.B.E. Publications Officer : Mrs. C. Rosner ADDRESSES OF THE COMMISSION AND THE TRUST Secretariat of the Commission : 28 Park Village East, Regent’s Park, London, N.W.1. Offices of the Trust : 41 Queen’s Gate, London, S.W.7. FOREWORD Each of the volumes of the present work so far published has consisted of a group of consecutively published Opinions and Declarations. The present volume however, comprises only a single Opinion, namely Opinion 283, the Opinion embodying the decisions in regard to the names properly applicable to the human malaria parasites taken by the International Commission on Zoological Nomenclature at its Session held in Paris in 1948. This arrangement has been adopted for two reasons. First, the length of this Opinion with its enclosures make it more con- venient for library use that it should constitute a single unit rather than be bound up as a part of a larger volume. Second, the interest attaching to the present Opinion extends over a much wider field than does that of most other Opinions rendered by the International Commission. For this reason also, it has been judged that for library purposes the most convenient arrangement would be for it to be issued as a single volume, complete with its own index. FRANCIS HEMMING Secretary to the International Commission on Zoological Nomenclature 28 Park Village East, Regent’s Park, LONDON, N.W.1, England. 15th May, 1954. a 2 eS eRe ee ae a - — es — A i a , | ; pe ee | es liyid cole tie? 2 a anh : : : \ y" . AO iW) wy. i f Z et ae y wy ak ae) eh | zi a) 4 a j ee ee Jy y , : f | : . nl Delhi Shke chit: gee ; ol ha & ae Lan SA at BN) x eres: y eer aly | 1 Oe ; Lr teeg crt aa i Vil TABLE OF CONTENTS OPINION 283. VALIDATION, UNDER THE PLENARY POWERS, OF THE GENERIC AND SPECIFIC NAMES COMMONLY USED FOR THE MALIGNANT TERTIAN MALARIA PARASITE AND THE QUARTAN MALARIA PARASITE RESPECTIVELY Page The Statement of the Case Investigations undertaken during the period 1943-1944 by Mr. Francis Hemming, Secretary to the Inter- national Commission on Zoological Nomenclature 13 Application submitted in 1944 jointly by Dr. Curtis W. Sabrosky and Dr. Robert L. Usinger, with a Supplement thereto submitted in 1946 __... ne 15 The Subsequent History of the Case Registration of the Present Application ge ee 23 Issue in 1944 of an appeal to specialists for advice on problems involved in the present case a Lis 23 Comments received from specialists in response to the appeal issued in 1944: Comment by Colonel Paul F. Russell (Chief, Division of Parasitology, Army Medical School, Army Medical Center, Washington, D.C., U.S.A.) 26 Comment by Dr. David Weinman (Columbia University, College of Physicians and Surgeons, De Lamar Institute of Public Health, New York, N.Y., U.S.A.) a my oe ve 26 Completion by Mr. Francis Hemming in 1947 of the Survey of the History of the Nomenclature of the Human Malaria Parasites begun in 1943 and the submission of that Survey to the Commission _... 27 Vill Page Issue in 1947 of Public Notices regarding the Possible Use by the International Commission of its Plenary Powers in the present case ... ah e Ee 28 The Decision taken by the International Commission on Zoological Nomenclature at its Session held in Paris in1948__.... ey a ie ga ae ed 29 APPENDIX TO ‘‘ OPINION ” 283 The Human Malaria Parasites: A Proposal for the Stabilisation of Nomenclature on the Basis of Current Usage ‘ By FRANCIS HEMMING, C.M.G., C.B.E., Secretary to the International Commission on Zoological Nomenclature Introductory... vp ey! vg bh 4. i 57 The Specific Trivial Names bestowed (or alleged to have been bestowed) upon the Human Malaria Parasites in the Period 1881-1897 tf ae a =e a 71 The Names of the Nominal Genera established (or alleged to have been established) for the Human Malaria Para- sites in the Period 1881-1897... fe ¥y a ee Synonymy of the Specific Names bestowed (or alleged to have been bestowed) upon the Human Malaria Para- sites in the Period 1881-1897 and of the names of the Nominal Genera established (or alleged to have been established) for those Species in the same Period ... 185 Original references for the names given, or = to the Human Malaria Parasites ve Synonymy of the Specific Trivial Names bestowed (or alleged to have been bestowed) upon the Human Malaria Parasites Names bestowed upon, or used for, the Malignant Tertian Malaria Parasite Names bestowed upon, or used for, the Quartan Malaria Parasite .. Names bestowed upon, or used for, the Benign Tertian Malaria Parasite Names of the Nominal Genera established (or alleged to have been established) for the Human Malaria Parasites The Correct Names for the Human Malaria Parasites under a strict application of the Régles Comparison of the names applicable to the Human Malaria Parasites under a strict application of the Régles with the names currently applied to those species ... aes ffs ne am Proposal for the Stabilisation of the Names for the Human Malaria Parasites on the Basis of Current Usage The Specific Trivial Name for the Quartan Malaria Parasite $Y a gen we me The Specific Trivial Name for the causa Tertian Malaria Parasite The Specific Trivial Name for the eaeae Tertian Malaria Parasite Certain terms consisting of Latin adjectives cited in the genitive case published as technical designations, but not as names, for human malaria parasites IX Page 186 188 189 190 191 192 195 196 199 200 203 203 The generic name Oscillaria as used by Laveran in 1881 The generic name Plasmodium Marchiafava & Celli, 1885 The generic name Laverania Feletti & Grassi, 1889 ... Proposed clarification of Article 14 of the “‘ Régles ” on the question of the status of terms consisting of Latin adjectives cited in the genitive case when published as though constituting names for species or subspecies ... SUMMARY OF RECOMMENDATIONS Recommendations for the stabilisation of the names of the human malaria parasites on the basis of current usage Recommendations for the clarification of Article 14 of the Régles in relation to the status of terms consisting of Latin adjectives cited in the genitive case when pub- lished as though constituting names for species or subspecies APPENDICES Appendix 1 :—The discovery by Golgi of the Quartan Malaria Parasite. By Colonel Sir Rickard Christo- phers, C.L.E., O.B.E., F.R.S. ine ays Appendix 2 :—Reproduction in facsimile of the paper entitled Sui parasiti della malaria by Feletti (R.) & Grassi (B.) as published in the serial publication Riforma medica, Napoli, in its issue of 15th January, 1890 and of the supplement to the foregoing paper entitled Sui parasiti della malaria : aggiunta della nota preliminare published in the same serial publication in its issue of Ist March, 1890 Page 204 205 207 211 213 221 223 225 INDEX XI to the names and technical designations which have been given, or have been alleged to have been given, to the Human Malaria Parasites Cytamoeba Danilewsky, 1891 Cytosporon Danilewsky, 1891 Cytozoon Danilewsky, 1891 ... Ematozoo (correction of ematozoo) Antolisei & Angelini, 1890 Ux HR a a aay: i falciforme, Ematozoo (correction of ematozoo), Antolisei & Angelini, 1890 se ae a, falciforme, Haematozoon (correction of shaematozoon), Thayer & Hewetson, 1895 falciparum, Haematozoon, Welch, 1897 Haemamoeba Feletti & Grassi, Dec. 1889 ... Haemamoeba Labbé, 1894 Haematomonas Mitrophanow, 1883 Haematomonas Osler, 1886 Haematophyllum Metschnikoff, 1887 Haematozoon (correction of haematozoon) Thayer & Hewetson, 1895 Haemocytsporon Danilewsky, 1891 ... Haemosporidium Lewkowicz, 1897 .. hominis, Laverania, Danilewsky, 1891 immaculata, Haemamoeba, Grassi, 1891 irregularis, Plasmodium malariae, Kruse, 1892 irregularis, Haemamoeba febris, Sakharov, 1892 laverani, Haemamoeba, Labbé, 1894 Page 176 176 176 175 112 146 15] ge 178 166 166 © 169 179 176 18] 132 119 134 137 140 XII Laverania Feletti & Grassi, Dec. 1889 malariae, Haemamoeba, Feletti & Grassi, Dec. 1889 malariae, Laverania, Feletti & Grassi, Dec. 1889 ... malariae, Oscillaria, Laveran, 1881 malariae, Plasmodium, Marchiafava & Celli, 1885 Oscillaria, Schrank, 1832 Oscillaria Laveran, 1881 Plasmodium Marchiafava & Celli, 1885 praecox, Haemamoeba, Grassi & Feletti, May 1890 quartana, Haemamoeba laverani, var. Labbé, 1894 quartanae, Haemosporidium, Lewkowicz, 1897 quartanae, Plasmodium malariae, M. & C. Varieta A., Celli & Sanfelice, 1891 ... quartanae, Plasmodium malariae, Kruse, 1892 quotidianae, Plasmodium malariae M. & C., Varieta C., Celli & Sanfelice, 1891 sedecimanae, Haemosporidium, Lewkowicz, 1897 ... tertiana, Haemamoeba laverani var., Labbé, 1894 ... tertianae, Haemosporidium, Lewkowicz, 1897 tertianae, Plasmodium malariae M. & C., Varieta B., Celli & Sanfelice, 1891 tertianae, Plasmodium malariae, Kruse, 1892 tropica, Plasmodium, Koch, 1899 undecimanae, Haemosporidium, Lewkowicz, 1897 ... vigesimo-tertianae, Haemosporidium, Lewkowicz, 1897 vivax, Haemamoeba, Grassi & Feletti, May 1890 Page 173 95 105 ae 83 159 159 164 107 140 148 124 134. 124 148 140 148 124 134 152 148 148 106 XIil DATE OF PUBLICATION OF THE PRESENT VOLUME The whole of the present volume was published as a single unit on 28th December 1954. INSTRUCTIONS TO BINDERS The present volume should be bound up as follows :—T.P.— XII—[XVI], 1—225, coloured wrapper (cover) in which the volume was published. i : f o; eae | or + Ai ali illae ite ee ate ray 3 oe ‘idee | ee ay aa ‘ eh no “| oi Pe, vs is 7 er > en | 7 so” Lae a. +e an z aa Ot ete Shy PAY AMIS it ’ widow , ¢: fT by dn? fy -- : vat Oe =) 4 ~ a ey 7 i) i, | > ae ‘ME @ ; _ esis) ; - ; eo A rat SAA Se Tita ae A! oe: — -* a ee , hie 7 6 ; o or, oa ; ‘ re Fp, vue ea ni sh ae ie iy e . Ang | net: * & i ¢ i) aT i a, | | | bey r Edited by FRA NCIS HEMMING, C.M.G., C.B.E. | ‘ ‘Secretary to the Commission VOLUME 7. Tn FEB 15 1°58 - “ Piensa Parasite and the ie en siwikce Parasite Tespectively LONDON : _ Printed by Order of the International Trust for Zoological Nomenclature and r “Sok Id on ‘behalf of the International Commission on Zoological Tee omenclature by the International Trust at its Publications Office a «41, Queen’s Gate, London, 8.W.7 1954 Price Six Guineas twa? er m7. . ea? 5 mae 4 a « ‘ ~ Lae a z ad | Bair La - Aes < oy ‘Si = (All rights reserved) INTERNATIONAL COMMISSION ON ZOOLOGICAL NOMENCLATURE COMPOSITION AT THE TIME OF THE ADOPTION OF THE OPINIONS PUBLISHED IN THE PRESENT VOLUME A. The Officers of the Commission President: Dr. Karl Jordan, Ph.D., F.R.S. (British Museum (Natural History), Zoological Museum, Tring, Herts, England). Vice-President : Dr. James L. Peters (Museum of Comparative Zoology, Cambridge, Massachusetts, U.S.A.). Secretary : Mr. Francis Hemming, C.M.G., C.B.E. (London, England). B. The Members of the Commission . Class 1949 Senor Dr. Angel CABRERA (La Plata, Argentina). Mr. Francis HeEMMING (London, England) (Secretary to the Commission) Dr. Karl JorDAN (British Museum (Natural History), Zoological Museum, Tring, Herts, England) (President of the Commission). Dr. Th. MorTENSEN (Universitetets Zoologiske Museum, Copenhagen, Denmark). Dr. Joseph PEARSON (Tasmanian Museum, Hobart, Tasmania, Australia). Class 1952 Senhor Dr. Afranio do AMARAL (S. Paulo, Brazil). Professor J. Chester BRADLEY (Cornell University, Ithaca, N.Y., U.S.A.). Professor Lodovico di CAPoRIACcO (University of Parma, Italy). Professor J. R. DyMonpD (University of Toronto, Canada). Dr. James L. PETERs (Museum of Comparative Zoology, Cambridge, Massachusetts, U.S.A.) (Vice-President of the Commission). Dr. Gan) E. Voxes (United States Geological Survey, Washington, D.C., Class 1955 Professor Dr. Hilbrand BoscHmMa (Rijksmuseum van Natuurlijke Historie, Leiden, The Netherlands). Dr. William Thomas CaLMAN (Tayport, Fife, Scotland). Professor Teiso EsAkiI (Kyushu University, Fukuoka, Japan). Professor Béla HANKO (University of Debrecen, Hungary). Dr. Tadeusz JACZEWSKI (Polish Zoological Museum, Warsaw, Poland). Dr. Norman R. Stout (Rockefeller Institute for Medical Research, Princeton, New Jersey, U.S.A.). C. Alternate Members of the Commission at the Session held in Paris’ in 1948 Professor Ehrique BeELTRAN (Instituto Mexicano de Recursos Naturales Renovables A.C., Mexico City, Mexico). Dr. Edward HINDLE (Zoological Society of London, London, England). Dr. Arturo Ricardo JorGe (Museu Bocage, Lisbon, Portugal). Professor Harold Kirsy (University ol California, Berkeley, California, U.S.A.). Dr. Henning Lemcue (Kgl. Veteriner- og Landbohgjskole, Zoologisk Labora- torium, Copenhagen, Denmark). Professor Kamel MANsour (King Fouad University, Cairo, Egypt). Professor Z. P. MercatF (North Carolina State.College of Agriculture and Engineering, University of North Carolina, Raleigh, North Carolina, U.S.A.). - Mr. N. D. Rivey (British Museum (Natural History), London, England). nah si — SpARCK (Universitetets Zoologiske Museum, Copenhagen, enmarK). Professor Victor van STRAELEN (Institut Royal des Sciences Naturelles de Belgique, Bruxelles, Belgium). TOs hS L. Ustncer (University of California, Berkeley, California, OPINIONS AND DECLARATIONS RENDERED BY THE INTER- NATIONAL COMMISSION ON ZOOLOGICAL NOMENCLATURE Edited by FRANCIS HEMMING, C.M.G., C.B.E. Secretary to the Commission VOLUME 7. Parti. Pp. 1—226, 1 pl. and 5 facsimiles 4 OPINION 283 Validation, under the Plenary Powers, of the generic and specific names commonly used for the Malignant Tertian Malaria Parasite and’ the Quartan Malaria Parasite respectively Also published with this Part: Title Page, Foreword; Table of Contents. LONDON : Printed by Order of the International Trust for Zoological Nomenclature and Sold on behalf of the International Commission on Zoological Nomenclature by the International Trust at its Publications Office 41, Queen’s Gate, London, S.W.7 1954 Price Six Guineas (All rights reserved) Issued 28th December, 1954 INTERNATIONAL COMMISSION ON ZOOLOGICAL NOMENCLATURE COMPOSITION AT THE TIME OF THE ADOPTION OF THE PRESENT “ OPINION ” A. The Officers of the Commission President: Dr. Karl Jordan, Ph.D., F.R.S. (British Museum (Natural History), Zoological Museum, Tring, Herts, ’ England). Vice-President : Dr. James L. Peters (Museum of Comparative Zoology, Cambridge, Massachusetts, U.S.A.). Secretary : Mr. Francis Hemming, C.M.G., C.B.E. (London, England). B. The Members of the Commission Class 1949 Senor Dr. Angel CABRERA (La Plata, Argentina). Mr. Francis HEMMING (London, England) (Secretary to the Commission) Dr. Karl JorDAN (British Museum (Natural History), Zoological Museum, Tring, Herts, England) (President of the Commission). Dr. Th. MorTENSEN (Universitetets Zoologiske Museum, Copenhagen, Denmark). Dr. Joseph PEARSON (Tasmanian Museum, Hobart, Tasmania, Australia). Class 1952 Senhor Dr. Afranio do AMARAL (S. Paulo, Brazil). Professor J. Chester BRADLEY (Cornell University, Ithaca, N.Y., U.S.A.). Professor Lodovico di CApoRIACCO (University of Parma, Italy). Professor J. R. DyMonbD (University of Toronto, Canada). Dr. James L. Peters (Museum of Comparative Zoology, Cambridge, Massachusetts, U.S.A.) (Vice-President of the Commission). Dr. Pe E. VoKes (United States mene) Survey, Washington, D.C., U.S.A.) Class 1955 Professor Dr. Hilbrand BoscHMA (Rijksmuseum van Natuurlijke Historie, Leiden, The Netherlands). Dr. William Thomas CALMAN (Tayport, Fife, Scotland). Professor Teiso EsAki (Kyushu University, Fukuoka, Japan). Professor Béla HANKO (University of Debrecen, Hungary). Dr. Tadeusz JACZEwskI (Polish Zoological Museum, Warsaw, Poland). Dr. Norman R. SToii (Rockefeller Institute for Medical Research, Princeton, New Jersey, U.S.A.). C. Alternate Members of the Commission at the Session held in Paris in 1948 Professor Ehrique BELTRAN (Instituto Mexicano de Recursos Naturales Renovables A.C., Mexico City, Mexico). Dr. Edward HINDLE (Zoological Society of London, London, England). Dr. Arturo Ricardo JorGE (Museu Bocage, Lisbon, Portugal). Professor Harold Kirsy (University ol California, Berkeley, California, U.S.A.). Dr. Henning LemMcue (Kgl. Veteriner- og Landbohgjskole, Zoologisk Labora- torium, Copenhagen, Denmark). Professor Kamel MANsourR (King Fouad University, Cairo, Egypt). Professor Z. P. Metcatr (North Carolina State College of Agriculture and Engineering, University of North Carolina, Raleigh, North Carolina, U.S.A.). Mr. N. D. RiLey (British Museum (Natural History), London, England). Professor Ragnar Sparck (Universitetets Zoologiske Museum, Copenhagen, Denmark). Professor Victor van STRAELEN (Institut Royal des Sciences Naturelles de Belgique, Bruxelles, Belgium). eke oe L. UsINcer (University of California, Berkeley, California, | OPINION 283 VALIDATION, UNDER THE PLENARY POWERS, OF THE GENERIC AND SPECIFIC NAMES COMMONLY USED FOR THE MALIGNANT TERTIAN MALARIA PARASITE AND THE QUARTAN MALARIA PARASITE RESPECTIVELY RULING :—(1) The incorrect particulars relating to the generic names Plasmodium and Laverania and their type species given in Opinion 104 and, in consequence, in the Official List of Generic Names in Zoology are hereby cancelled. (2) The Plenary Powers are hereby used for the pur- poses and to the extent specified below:— (a) The under-mentioned specific names published for the Malignant Tertian Malaria Parasite are here- by suppressed for the purposes both of the Law of Priority and of the Law of Homonymy: (i) malariae Laveran, 1881, as published in the com- bination Oscillaria malariae; (1) malariae Mar- chiafava & Celli, 1885, as published in the. combination Plasmodium malariae; (iii) malariae Feletti & Grassi, 1889, as published in the com- bination Laverania malariae (in so far as this was published as a new name and does not represent a transfer of Oscillaria malariae Laveran, 1881, to the genus Haemamoeba); (b) The under-mentioned specific names published for the Malignant Tertian Malaria Parasite are hereby suppressed for the purposes of the Law of Priority but not for those of the Law of Homo- nymy: (1) falciforme Antolisei & Angelini, 1890, as published in the combination Ematozoo falci- forme; (ii) laverani Labbé, 1894, as published in JAW 2 51955 OPINIONS AND DECLARATIONS the combination Haemamoeba laverani; (iil) falciforme Thayer & Hewitson, 1895, as pub- lished in the combination Haematozoon falci- forme; (c) The indication, by monotypy of Plasmodium malariae Marchiafava & Celli, 1885 (the Malignant Tertian Malaria Parasite) as the type species (1) of the genus Plasmodium Marchiafava & Celli, 1885, and (ii) of the genus Haematophyllum Metschnikoff, 1887 (the name of which was published as a substitute name (nom. nov.) for the name Plas- modium Marchiafava & Celli, 1885, in the erroneous belief that that name was notavailable under the Rég/es) is hereby set aside, and, in place of the foregoing species, the species Haemamoeba malariae Feletti & Grassi, 1889 (the Quartan Malaria Parasite) is hereby designated as the type species both of Plasmodium Marchiafava & Celli, 1885, and of Haematophyllum Metschnikoff, 1887; (dq) The under-mentioned specific names are hereby validated as follows: (i) the name malariae Feletti & Grassi, 1889, as published in the com- bination Haemamoeba malariae, to be the name of the Quartan Malaria Parasite, notwithstanding the fact that, prior to the suppression, under the Plenary Powers, of the names specified in (a) above, that name was an invalid secondary homonym; (ii) falciparum Welch, 1897, as pub- lished in the combination Haematozoon falci- parum, to be the name of the Malignant Tertian Malaria Parasite; (ce) The indication, by monotypy, of Laverania malariae Feletti & Grassi, 1889, or, as the case may be, of Oscillaria malariae Laveran, 1881 (being names for the Malignant Tertian Malaria Parasite sup- pressed under the Plenary Powers, under (a) above) as the type species of the genus Laverania Feletti & Grassi, 1889, is hereby set aside, and, in OPINION 283 5 place of the foregoing species, the species Haema- tozoon falciparum Welch, 1897, is hereby desig- nated to be the type species of the genus Laverania Feletti & Grassi, 1889; (f) The generic name Laverania Feletti & Grassi, 1889 (type species, by designation under the Plenary Powers, under (e) above: Haematozoon falci- parum Welch, 1897, validated under the Plenary Powers, under (d) above, as the name of the Malignant Tertian Malaria Parasite) is hereby validated. (3) The under-mentioned generic names are hereby declared to be either invalid or not required for the reasons severally stated below against the names in question :— (a) Oscillaria Laveran, 1881 (in so far as Laveran pub- lished this as a new name and not as Oscillaria Schrank, 1823): invalid, because a junior homonym of Oscillaria Schrank, 1823, that name retaining, under Article 1, its status under the Law of Homonymy, notwithstanding the fact that the genus so named is currently treated as belonging to the Vegetable Kingdom; (b) Haematophyllum Metschnikoff, 1887: invalid be-- cause a junior objective synonym of Plasmodium Marchiafava & Celli, 1885 (these two nominal genera having the same nominal species as type species) ; (c) Haemamoeba Feletti & Grassi, 1889; invalid be- cause a junior objective synonym of the name Plasmodium Marchiafava & Celli, 1885 (con- sequent upon the designation under the Plenary Powers, under (2)(c) above, of Maemamoeba malariae Feletti & Grassi, 1889, the nominal species which is the type species, by monotypy, of Haemamoeba Feletti & Grassi, 1889, to be also the type species of Plasmodium Marchifava & Celli, 1885); 6 OPINIONS AND DECLARATIONS (d) Ematozoo Antolisei & Angelini, 1890: not required, because the type species (Ematozoo falciforme Antolisei & Angelini, 1890) of the genus so named is a junior subjective synonym of Haemato- zoon falciparum Welch, 1897, designated under the Plenary Powers, under (2)(/) above, to be the type species of the genus Laverania Feletti & Grassi, 1889; ‘ (e) Haematozoon Thayer & Hewetson, 1895: not re- quired because Haematozoon falciforme Thayer & Hewetson, 1895, the type species, by monotypy, of the genus so named, is a junior subjective synonym of Haematozoon falciparum Welch, 1897, designated under the Plenary Powers, under (2)(f) above, to be the type species of the genus Laverania Feletti & Grassi, 1889 ; (f) Haemosporidium Lewkowicz, 1897: invalid because a junior objective synonym of Plasmodium Mar- chiafava & Celli, 1885 (consequent upon the designation under the Plenary Powers, under (2)(c) above, of Haemamoeba malariae Feletti & Grassi, 1889, the type species of Haemosporidium Lewkowicz, 1897 (by selection under Article 30, Rule (g)) to be the type species also of Plas- modium Marchiafava & Celli, 1885); (4) The under-mentioned specific names, each of which was published as the name of a new avian parasite but in the description of each of which there appeared an incorrect statement that the parasite in question had been found in the blood of human malaria patients, are hereby declared not to be available as specific names for the Malignant Tertian Malaria Parasite (the parasite mis- identified with the avian parasite concerned by the authors in question), these specific names adhering, under the Rég/les, to the avian parasites from which the original OPINION 283 7 descriptions of these parasites were drawn up by their respective authors :— (a) praecox Grassi & Feletti, 1890, as published in the combination Haemamoeba praecox; ; (b) immaculata Grassi, 1891, as published in the com- bination Haemamoeba immaculata. (5) The under-mentioned terms consisting of Latin adjectives published in the genitive case in agreement not with the generic name (as required by Article 14(1)(a)) but with the specific name, either expressed or understood, are hereby declared to have been published not as sub- specific names of human malaria parasites, but as techni- cal designations for those species, and, in consequence, the Latin adjectives of which these terms are composed possess no status as subspecific names in zoological nomenclature :-— (a) guartanae Celli & Sanfelice, 1891, as published in connection with the specific name Plasmodium malariae; (b) tertianae Celli & Sanfelice, 1891, as published in connection with the name Plasmodium malariae; (c) quotidianae Celli & Sanfelice, 1891, as published in connection with the name Plasmodium malariae : (d) tertianae Kruse, 1892, as published in connection with the name Plasmodium malariae ; (e) guartanae Kruse, 1892, as published in connection with the name Plasmodium malariae; (f) irregularis Kruse, 1892, as published in connection with the name Plasmodium malariae; (g) tertianae Lewkowicz, 1897, as published in connec- tion with the generic name Haemosporidium; 8 OPINIONS AND DECLARATIONS (h) guartanae Lewkowicz, 1897, as published in connec- tion with the generic name Haemosporidium; (i) undecimanae Lewkowicz, 1897, as published in connection with the generic name Haemospori- dium; (j) sedecimanae Lewkowicz, 1897, as published in connection with the generic name Haemospori- dium; (k) vigesgimo-tertianae Lewkowicz, 1897, as published connection with the generic name Haemospori- dium. (6) It is hereby placed on record that there is no such generic name as Haematomonas Osler, 1886, Osler in the passage in question not having published a new generic name but having referred to the previously published name Haematomonas Mitrophanow, 1883. (7) It is hereby recalled that, under the Ruling given in Opinion 101, the under-mentioned generic names and the under-mentioned specific name published for human malaria parasites by Danilewsky in 1891 possess no status in zoological nomenclature :— (a) the under-mentioned generic names: (i) Cytamoeba Danilewsky, 1891; (ii) Cytosporon Danilewsky, 1891; (iti) Cytozoon Danilewsky, 1891; (iv) Haemocytosporon Danilewsky, 1891; (b) the specific name hominis Danilewsky, 1891, as published in the combination Laverania hominis. (8) It is hereby placed on record that the name quartana Labbé, 1894, as published as a subspecific name in the combination Haemamoeba laverani var. quartana, is an available name but is not required for the Quartan Malaria Parasite, it being a junior subjective synonym of the specific name malariae Feletti & Grassi, 1889, as published in the combination Haemamoeba malariae, validated under the Plenary Powers, under (2)(d)(i) above. OPINION 283 9 (9) It is hereby placed on record that the under- mentioned alleged specific names, not having been published, are cheironyms, and, in consequence, possess no status in zoological nomenclature: — (a) irregularis Sakharov, erroneously alleged to have been published in 1892 as a subspecific name in the combination Haemamoeba febris irregularis ; (b) tropica Koch, erroneously alleged to have been published in 1899 in the combination Plasmo- dium tropica. (10) The specific name vivax Grassi & Feletti, 1890, as published in the combination Haemamoeba vivax, is hereby declared to be the oldest available specific name for, and, therefore, the valid specific name of, the Benign Tertian Malaria Parasite. (11) It is hereby placed on record that the specific name ftertiana Labbé, 1894, as published as a subspecific name in the combination Haemamoeba laverani var. tertiana, is an available name, but is not required for the Benign Tertian Malaria Parasite, it being a junior sub- jective synonym of the specific name vivax Grassi & Feletti, 1890, as published in the combination Haema- moeba vivax. (12) The following particulars in regard to the generic names Plasmodium and Laverania, together with the Televant bibliographical references, are hereby inserted in the Official List of Generic Names in Zoology in place of the particulars deleted therefrom under (1) above :— (a) Plasmodium Marchiafava & Celli, 1885 (gender : neuter) (type species by designation under the Plenary Powers: Haemamoeba malariae Feletti & Grassi, 1889 (the Quartan Malaria Parasite)); 10 OPINIONS AND DECLARATIONS (b) Laverania Feletti & Grassi, 1889 (gender : feminine) (type species, by designation under the Plenary Powers : Haematozoon falciparum Welch, 1897 (the Malignant Tertian Malaria Parasite)) (generic name to be used by authors who consider the Malignant Tertian (or Aestivo-Autumnal) Malaria Parasite to be generically distinct from the Quartan Malaria Parasite). (13) The under-mentioned generic names or alleged generic names are hereby placed on the Official Index of Rejected and Invalid Generic Names in Zoology with the Name Numbers severally cited below :— (a) the under-mentioned generic names rejected under 3) above, to be entered as Name Nos. 88 to 91 : t Haemamoeba Feletti & Grassi, 1889; (ii) Haematophyllum Metschnikoff, 1887; (ii) Haemo- sporidium WLewkowicz, 1897; (iv) Oscillaria Laveran, 1881 (in so far as this was published as a new name and not as Oscillaria Schrank, 1823); (b) the reputed but non-existant generic name Haema- tomonas Osler, 1886, rejected under (6) above, to be entered as Name No. 92; (c) the under-mentioned generic names rejected under (7) above, to be entered as Names Nos. 93 to 96: (1) Cytamoeba Danilewsky, 1891; (ii) Cyto- sporon Danilewsky, 1891; (ai1) Cytozoon Dani- lewsky, 1891; (iv) Haemocytosporon Danilewsky, 1891. (14) The under-mentioned specific names are hereby placed on the Official List of Specific Names in Zoology ~ as Names Nos. 112 to 114: (i) falciparum Welch, 1897, as published in the combination Haematozoon falciparum (the Malignant Tertian Malaria Parasite); (ii) malariae Feletti & Grassi, 1889, as published in the combination Haemamoeba malariae (the Quartan Malaria Parasite); (iii) vivax Grassi & Feletti, 1890, as published in the combination Haemamoeba vivax (the Benign Tertian Malaria Parasite). | ' sl tt tie li ie as OPINION 283 11 (15) The under-mentioned specific names are hereby placed on the Official Index of Rejected and Invalid Specific Names in Zoology with the Name Numbers severally cited below :— (a) the under-mentioned names suppressed under the Plenary Powers, under (1)(a) above, for the pur- poses both of the Law of Priority and of the Law of Homonymy, to be entered as Name Nos. 45 to 47: (i) malariae Laveran, 1881, as published in the combination Oscillaria malariae; (i) malariae Marchiafava & Celli, 1885, as published in the combination Plasmodium malariae; (iii) malariae Feletti & Grassi, 1889, as published in the combination Laverania malariae ; (b) the under-mentioned names suppressed under the Plenary Powers, under (1)(b) above, for the pur- poses of the Law of Priority but not for those of the Law of Homonymy, to be entered as Name Nos. 48 to 50: (i) falciforme Antolisei & Angelini, 1890, as published in the combination Ematozoo falciforme; (41) falciforme Thayer & Hewetson, 1895, as published in the combination Haemato- zoon falciforme; (iii) laverani Labbé, 1894, as published in the combination Haemamoeba laverani; (c) the under-mentioned specific names published as_ the names of avian parasites, in so far as in the original description the names concerned were, as noted in (4) above, incorrectly applied also to the Malignant Tertian Malaria Parasite, the names concerned to be entered as Name Nos. 51 and 52: (4) immaculata Grassi, 1890, as published in the combination Haemamoeba im- maculata; (ii) praecox Grassi & Feletti, 1890, as published in the combination Haemamoeba praecox; (d) the under-mentioned terms, which have been alleged to have been published as names but which have been ruled in (5) above to have been published as OPINIONS AND DECLARATIONS technical designations, the entries now to be made to be given the Name Nos. 53 to 63 : (i) irregularis Kruse, 1892, as published in connection with the name Plasmodium malariae ; (ii) quartanae Celli & Sanfelice, 1891, as published in connection with the name Plasmodium malariae ; (iii) quart- anae Kruse, 1892, as published in connection with the name Plasmodium malariae ; (iv) quart- anae Lewkowicz, 1897, as published in connection with the generic name Haemosporidium; (v) quotidianae Celli & Sanfelice, 1891, as published inconnection with the name Plasmodium malairae; (vi) sedecimanae Lewkowicz, 1897, as published in connection with the generic name Haemospori- dium; (vii) tertianae Celli & Sanfelice, 1891, as published in connection with the name Plasmo- dium malariae; (viii) tertianae Kruse, 1892, as published in connection with the name Plasmodium malariae; (ix) tertianae Lewkowicz, 1897, as published in connection with the generic name Haemosporidium; (x) undecimanae Lew- kowicz, 1897, as published in connection with the generic name Haemosporidium; (xi) vigesi- motertianae Lewkowicz, 1897, as published in connection with the generic name Haemospori- dium ; (e) the name hominis Danilewsky, 1891, as published in the combination Laverania hominis, which, as noted in (7) above, is invalid under the Ruling given in Opinion 101, this name to be entered as Name No. 64; (f) the under-mentioned alleged names ruled in (9) above to be unpublished cheironyms, to be entered under the Name Nos. 65 and 66: (i) irregularis Sakharov, erroneously alleged to have been published in 1892 in the combination Haemamoeba febris irregularis ; (ii) tropica Koch, erroneously alleged to have been published in 1899 in the combination Plasmodium tropica. OPINION 283 13 I.—THE STATEMENT OF THE CASE Detection of errors in “ Opinion” 104: In January 1943, Mr. Francis Hemming, Secretary to the International Commission on Zoological Nomenclature, when checking certain miscellaneous documents which in 1939 had been transferred to his charge by Dr. C. W. Stiles on his election as Secretary to the Commission, found an offprint of a paper entitled ““ The Correct Name of the Malignant Tertian Malaria Parasite’? written jointly by Sir Rickard Christophers, C.I.E., F.R.S., Colonel I.M.S. (ret.) (Leverhulme Fellow under the Medical Research Council, London School of Hygiene and Tropical Medicine, London) and J. A. Sinton, M.D., D.Sc., Lieut-Colonel I.M.S. (ret.) (Manson Fellow, London School of Hygiene and Tropical Medicine), which had been pub- lished in December 1938, Brit. med. J. 1938 (vol. 2) 1130-1134. It became immediately apparent from this paper that the entries in regard to the generic names Laverania and Plasmodium made in the Official List of Generic Names in Zoology by the Ruling given in the Commission’s Opinion 104 (1928, Smithson. misc. Coll. 73 (No. 5):25-28) contained a number of serious errors which would need to be corrected by the International Commission. 2. Accordingly, in January 1944 Mr. Hemming entered into correspondence with Sir Rickard Christophers (at this time of the Department of Zoology, Cambridge University, Cambridge) with a view to elucidating certain questions as a preliminary to the preparation, for the consideration of the International Com- mission, of a paper drawing attention to the errors in Opinion 104 and submitting proposals for their rectification by the use by the Commission of its Plenary Powers for the purpose of stabilising the nomenclature of the human malaria parasites on the basis of the practice currently adopted by malariologists. This corres- pondence was extremely protracted, fresh nomenclatorial diffi- 14 OPINIONS AND DECLARATIONS culties constantly arising when known difficulties were overcome. A preliminary draft of the required application was completed by Mr. Hemming in the early part of 1944, but a number of important matters still remained to be settled. Mr. Hemming’s corres- pondence with Sir Rickard Christophers and Brigadier Sinton was still in progress when in August 1944 an application to the Commission on similar lines to that in preparation by Mr. Hemming was received from Dr. Curtis W. Sabrosky and Dr. Robert L. Usinger (then of the United States Public Health Service, Office of Malaria Control in War Areas, Atlanta, Georgia, U.S.A.). In correspondence between Mr. Hemming and Dr. Usinger, under cover of a letter from whom the foregoing joint application had been transmitted to the International Commission, it was agreed that the application submitted by the latter jointly with Dr. Sabrosky and the application upon the preparation of which Mr. Hemming had been engaged since the beginning of 1943 should be laid before the Commission as complementary applica- tions, each designed to secure the same general ends. It was not long after this that further nomenclatorial problems came to light, including, in particular, the discovery recorded by Sabrosky in 1946 that the material on which the nominal species Plasmodium malariae Marchiafava & Celli, 1885, was based was not homo- geneous and, although consisting primarily of examples of the Malignant Tertian Malaria Parasite, included also examples of the Benign Tertian Malaria Parasite. It was not until January 1948 that Mr. Hemming succeeded in obtaining photostat copies of the last of the papers which it was necessary to consult in order to make sure that it contained no further surprises. Throughout this long period, Mr. Hemming’s paper was under constant revision on questions of detail, and, when the last of the required photostats was received, it was not long before the final version was completed. By April 1948 the application in its final form was submitted to the International Commission. 3. Mr. Hemming’s application: Mr. Hemming’s application contains a detailed survey of the history of the names, generic and specific, given to the human malaria parasites from the time when OPINION 283 15 the first of these species was named by Laveran in 1881 up to the year 1897, in which the name Haematozoon falciparum was published by Welch as a name for the Malignant Tertian Malaria Parasite. At the conclusion of his application, Mr. Hemming submitted a series of proposals designed, not only to stabilise existing nomenclatorial practice but in addition to sweep away, with the aid of the Commission, the large number of invalid names and reputed but non-existent names which in the past have contributed so largely to the confusion in which the names for the human malaria parasites have been engulfed. Inevitably therefore Mr. Hemming’s paper is both complicated and lengthy. In these circumstances, it has been decided that the most con- venient course will be, while incorporating Mr. Hemming’s paper in the official records of the present case, to print it in a separate Appendix rather than actually to include it in the body of the present Opinion. Mr. Hemming’s proposals which are sum- marised in Part 6 of his paper will be found on pages 213 to 222 of the present Opinion. 4. Joint Application submitted by Dr. Sabrosky and Dr. Usinger: As explained in paragraph 2 above, Dr. Robert L. Usinger on 7th July 1944 addressed a letter (which was not however received until 22nd August) to the International Commission covering an application for the use, by the Commission, of its Plenary Powers for the purpose of stabilising the nomenclature of the human malaria parasites on the basis of the existing practice of malario- logists. In this paper, Drs. Sabrosky and Usinger accepted the conclusion reached by Christophers and Sinton that, contrary to the view held till then, it was the Malignant Tertian Malaria Parasite and not the Quartan Malaria Parasite which had been described by Marchiafava & Celli when in 1885 they had estab- lished the nominal genus Plasmodium; but they followed previous authors in assuming that Marchiafava & Celli had not cited any nominal species at the time when they established the foregoing genus, and in consequence of this, they treated the nominal species Oscillaria malariae Laveran, 1881, as the type species of 16 OPINIONS AND DECLARATIONS this genus. It transpired in subsequent correspondence that the applicants were not then aware of the rule laid down by the International Commission that all applications submitted to it should, as a preliminary, be published in the Bulletin of Zoological Nomenclature and had already submitted their paper for publica- tion in the serial publication Science, in which it was published within about a week of its receipt in the Offices of the Commission (Sabrosky & Usinger, Ist September 1944, Science 100 (No. 2502): 190-192). The problem dealt with in the foregoing application was further examined by Dr. Sabrosky in 1946 when he published a paper (Sabrosky, 25th October 1946, Science 104 (No. 2704): 401-402), in which he stated that an examination of Marchiafava & Celli’s paper of 1885 had satisfied him that, contrary all previously published statements, the foregoing authors when establishing the nominal genus Plasmodium, had established also a new nominal species to which they had given the new name Plasmodium malariae. In the same paper, Dr. Sabrosky reported that the detailed descriptions, the case histories and the figures on Plate VI published by Marchiafava & Celli, when establishing the © foregoing nominal species, had been critically analysed by Dr. Martin Young (National Institute of Health, Bethesda, Maryland) who had come to the conclusion that, although, as previously stated by Christophers & Sinton (1938), the material on which Marchiafava & Celli had based their nominal genus Plasmodium consisted predominantly of examples of the Malignant Tertian Malaria Parasite, it included also—a point that had not been noted by Christophers & Sinton—some examples of the Benign Tertian Malaria Parasite. Since this later paper by Dr. Sabrosky rendered out of date in certain particulars the application which he and Dr. Usinger had submitted to the Commission in 1944, Mr. Hemming, as Secretary to the Commission, wrote to the applicants, asking how they wished this matter to be dealt with when the case came to be laid before the Commission. As the result of this exchange of correspondence it was agreed (letters dated 2lst and 26th October 1947 from Dr. Sabrosky and Dr. Usinger respectively) that the original application should be laid before the Commission in the form in which it was submitted OPINION 283 17 in July 1944, notwithstanding the modifications rendered neces- sary by the evidence adduced in Dr. Sabrosky’s paper of 1946, but that in addition Dr. Sabrosky’s (1946) paper should be submitted to the Commission simultaneously with the original application, to which it would thus form a supplement. The two documents are as follows:— (1) Joint Application by Dr. Curtis W. Sabrosky and Dr. Robert L. Usinger submitted under cover of a letter from Dr. Usinger dated 7th July 1944 NOMENCLATURE OF THE HUMAN MALARIA PARASITES By CURTIS W. SABROSKY and ROBERT L. USINGER (United States Public Health Service, Office of Malaria Control in War Areas, Atlanta, Georgia, U.S.A.) The names for the malarial parasites of man have long been the subject of discussion, much of which has hopelessly intermixed zoo- logical and nomenclatorial considerations. Stiles (1928, Opinion 101, p. 13) has aptly remarked that “‘ the nomenclature of the parasites of malaria in man and birds represents one of the most confusing chapters in the entire history of zoological nomenclature.” The problem has been very ably and painstakingly reviewed by Christophers and Sinton.* We have carefully studied their paper, we concur in their conclusions, and we agree that strict adherence to the International Rules of Zoological Nomenclature would result in great confusion. The nomenclature adopted in Opinion 104 is clearly the best solution to the problem and should be maintained. However, we feel it necessary to point out that Opinion 104 did not provide an official answer, notwithstanding the fact that zoologists have for years regarded it as the final decision in the matter. NOMENCLATURE OF THE HUMAN MALARIA PARASITES UNDER A STRICT INTERPRETATION OF THE RULES Oscillaria Laveran, 1881 (type, O. malariae Laveran, 1881, the parasite of malignant tertian malaria; by monotypy). Malario- logists are now agreed that Laveran had the sexual forms of this species, only, in his first paper cited above. * R. Christophers & J. A. Sinton, 1938, British Medical Journal, 1938 (vol. 2): 1130—1134. 18 OPINIONS AND DECLARATIONS = Plasmodium Marchiafava & Celli, 1885 (type, P. malariae Laveran, 1881, parasite of malgnant tertian malaria; by monotypy). — Haemamoeba Feletti & Grassi, 1890 (type, H. malariae Feletti & Grassi, 1890, parasite of tertian and quartan fevers in man; by monotypy). The name has been shown to include two species, and therefore is not really monotypic. Grassi & Feletti later in 1890 restricted malariae Feletti & Grassi to the parasite of quartan fever and proposed H. vivax for the tertian-producing form. Result: the type of Haemamoeba is H. malariae Feletti & Grassi, as restricted by Grassi & Feletti to quartan fever. In the genus Plasmodium it is a secondary homonym of malariae Laveran, 1881, and the next oldest available name, quartanae Celli & Sanfelice, 1891, is valid. —Laverania Feletti & Grassi, 1889 (cf. footnote, Christophers & Sinton, 1938, p. 1133) (type, Oscillaria malariae Laveran; by monotypy). = Haematozoon Welch, 1897 (type, Haematozoon falciparum Welch 1897; by monotypy). Conclusions, under a strict interpretation of the Rules: (1) If zoologists agree on one genus for the malarial parasites, its name, as shown above, would be Oscillaria Laveran, because this name is not invalidated by previous use in botany (Code, Article 1) or by its unsuitability (Article 32). It will be noted that all five generic names were monotypic as originally proposed and that three—Oscillaria, Plasmodium, Laverania—have identical type species and would be isogenotypic synonyms whatever classification is used. (2) If zoologists decide that two genera are required, the name Oscillaria would be strictly correct for the parasite of malignant tertian fever, and Haemamoeba for the other two. (3) The names of the classical species of malaria under a single genus would be as follows :— Oscillaria vivax (Grassi & Feletti, 1890)—Tertian malaria. Oscillaria quartanae (Celli & Sanfelice, 1891)—Quartan malaria (to replace malariae Feletti & Grassi, 1890 nec Laveran 1881). Oscillaria malariae Laveran, 1881—Malignant tertian malaria. OPINION 283 19 It is generally agreed that such conclusions would result in great confusion in medical and zoological literature. The generic name _ Plasmodium and the specific names malariae (quartan)} and falciparum (malignant tertian) have long been accepted by malariologists through- out the world. The usage was supported by Opinion 104 in which Laverania and Plasmodium, together with 55 other generic names, were placed on the Official List of Generic Names with “ malariae (as restricted to quartan fever) ’’ designated as the type of Plasmodium and falciparum Welch (1897) designated as type of Laverania. The matter would thus appear to have been settled but for the following significant statement by Stiles (1928) in the presentation of the case for Opinion 104: *“The Secretary has personally checked these names and believes that they are all nomenclatorially available and valid, and that, there- fore, they can be adopted in harmony with the Rules instead of as nomina conservanda.” Contrary to this usage in Opinion 104, it is clear that malariae as used in the combination Laverania malariae by Grassi & Feletti was not a homonym but was the original malariae of Laveran, that falci- parum therefore was an unnecessary substitution, that falciparum after all was not the next oldest available name, that ‘‘ malariae (as restricted to quartan fever) ’’ could not be the valid type of Plasmodium Marchiafava & Celli, 1885, since the Plasmodium of that date, was based on malariae Laveran, the parasite of malignant tertian fever, and that Oscillaria Laveran, 1881, antedates both of the above generic names. To have arrived at any of the conclusions stated in the Opinion would therefore have required a Suspension of the Rules. Inasmuch as the Rules were not suspended for any of the names approved in Opinion © 104, we submit that the names Laverania and Plasmodium hold a place on the Official List in direct contravention of the Rules, rather than being maintained and protected by them. In the present instance, we are faced with an Official List containing certain names which are not official in the sense that the action neces- sary to make them so was never taken. We can find no justification for believing that names placed on the Official List, merely in the absence of any expressed “‘ objection, question, or adverse comment ” (Opinion 75, p. 35) at the time, are thereby conserved to eternity and not subject to critical evaluation. Since the names Plasmodium malariae (quartan) and falciparum (malignant tertian) are generally accepted in zoological and medical literature and since it was apparently t According to Christophers & Sinton, 1938, the name malariae Laveran was first applied to quartan malaria erroneously by Liihe (1900). 20 OPINIONS AND DECLARATIONS the intention of the International Commission to fix these names, we respectfully request that the International Commission on Zoological Nomenclature legalize Opinion 104 as it applies to the malaria parasites by suspending the Rules and taking the following action :— 1. Suppress the generic name Oscillaria Laveran (1881) in favour of Plasmodium Marchiafava & Celli (1885). 2. Suppress the species name malariae Laveran (1881), and any other names for the parasite of malignant tertian malaria, in favour of falciparum Welch (1897). 3. Establish malariae Feletti & Grassi (1889, 1890) nec malariae Laveran (1881) as the valid name for the parasite of quartan malaria. 4. Designate as the type of Plasmodium Marchiafava & Celli (1885)— Haemamoeba malariae Feletti & Grassi (1889, 1890). 5. Designate as the type of Laverania Feletti & Grassi (1889)— Haematozoon falciparum Welch (1897). In summary, the actions recommended above would legalize existing practice as follows :— Plasmodium vivax (Grassi & Feletti, 1890) parasite of tertian malaria. Plasmodium malariae (Feletti & Grassi, 1889, 1890), parasite of quartan malaria. Plasmodium falciparum Welch (1897), parasite of malignant tertian malaria. (2) Supplement to Application submitted by Dr. Sabrosky and Dr. Usinger on 7th July 1944 CORRECTION ON THE NOMENCLATURE OF HUMAN PLASMODIUM By CURTIS W. SABROSKY (United States National Museum, Washington, D.C., U.S.A.) In the nomenclatorial and zoological confusion in the names for the human malaria parasites (Sabrosky & Usinger, Science, 1944, 100, 190-192; Beltran. Gaceta Med. Mexico, 1944, 74, 61-74), one further point has been discovered. OPINION 283 21 It has usually been considered that there were only two different proposals involving malariae as a new specific name—Oscillaria malariae Laveran, 1881, and Haemamoeba malariae Feletti & Grassi, 1890. Actually it now appears that there were three ! Careful reading of Marchiafava & Celli (Fortschr. Med., 1885, 3: 791-797) shows that they too were proposing what they regarded as a distinct new form, quite unlike any previously described: ‘‘ Aus dem Gesagten geht hervor, dass die beschriebenen K6rperchen nicht verwechselt werden diirfen mit irgend welchen zufalligen oder patho- logischen Dingen, die man bisher inden roten Blutscheiben bemerkt hat . . . so scheint es uns nicht fernliegend, sie als parasitare Organismen anzusprechen und ihnen den Namen Plasmodium malariae zu geben.” The name was italicized and unquestionably intended as a formal scientific name. This proposal of Plasmodium as a new generic name has been accepted, but the new specific name has apparently been quite -generally overlooked. _ Of the three identical specific names, it is now generally agreed that malariae Laveran applied only to the parasite of falciparum or malig- nant malaria. Malariae Feletti & Grassi, which originally included both quartan and tertian parasites, was later restricted by Grassi & Feletti (Arch. ital. biol., 1890, 13: 300) to the quartan parasite, at which time they named the tertian parasite vivax. The situation is not so clear, however, with reference to the malariae of Marchiafava & Celli, 1885. Their detailed descriptions, case histories, and figures in Plate VI have been critically analysed by Martin Young, of the National Institute of Health, as follows:— It seems to me definitely that most of the infections that they saw were falciparum. However, they seem occasionally to have run across a vivax infection. Some of the descriptions are definitely of vivax segmenters while others of the descriptions, especially where they mention the finding of crescents, are definitely falciparum. The generalized figures on the plate are difficult to identify, with some suggestive of vivax and falciparum. The confusion arises from the fact that some of the cases they were looking at were very severe infections of falciparum. In such cases, it is not uncommon for the developmental forms of falciparum to be found in the blood stream. Therefore, from pictures with so little detail, it is hard to tell whether the forms shown are young stages of vivax or older stages of falciparum. 22 OPINIONS AND DECLARATIONS Throughout the paper, Marchiafava & Celli referred frequently to such characteristics of falciparum or malignant malaria as the comatose fever, rapid onset of death, remarkable number of parasites (especially in the capillaries of the brain), and the presence of crescents. Besides this evidence, it may be noted that their cases originated during a very severe epidemic in Rome and the Pontine Marshes, where falciparum is the principle species of malaria. There seems to be little doubt, therefore, that Plasmodium malariae. Marchiafava & Celli was based mainly on the malignant tertian parasite (falciparum). The genign tertian parasite (vivax) was seen, but. there is no evidence of quartan. If malariae Marchiafava & Celli be considered to represent one species only, then the malignant tertian form would have to be con- sidered the genotype of Plasmodium, in which case Oscillaria Laveran, Plasmodium Marchiafava & Celli, and Laverania Feletti & Grassi are isogenotypic, all based on the same species (properly called malariae Laveran, under strict interpretation of the Rules of Nomenclature, but generally known as falciparum). On the other hand, if malariae Marchiafava & Celli be regarded as originally composed of two species, it would then appear that the type has never been restricted to a single entity because the species name has been so long overlooked. Even though the malignant tertian parasite is unquestionably the basis of Marchiafava & Celli’s descrip- tion, it appears that at this late date considerable confusion could be avoided by restricting the name malariae Marchiafava & Celli to the benign tertian form. If such action were taken, and considering that all the human malaria parasites are congeneric (as they are usually regarded), the name would then be a homonym, and the correct name would be the next valid and available name, hence vivax Grassi & Feletti, 1890 (=malariae Marchiafava & Celli, 1885, nec malariae Laveran, 1881). Thus (1) vivax would become the genotype of Plas- modium Marchiafava & Celli; (2) it would not be necessary to suspend the Rules of Nomenclature in order to designate a type for Plasmodium; and (3) the status of Laverania as a possible generic name for the malignant parasite (if segregated) would not be disturbed. In order not to complicate any other action by the International Commission on Zoological Nomenclature, formal designation of the above is withheld, and it is presented as a suggestion to be considered as a part of the whole problem. OPINION 283 23 II.—THE SUBSEQUENT HISTORY OF THE CASE 5. Registration of the present application: Immediately upon a _ decision being taken by the Secretary that it was necessary to sub- mit an application for the correction of the erroneous entries regarding the generic names Laverania and Plasmodium in the Official List of Generic Names in Zoology made under the directions given in Opinion 104, a Registered File was opened (on 17th January 1943) for the present case, the number so allotted being Z.N. (S) 143. 6. Investigations undertaken by Mr. Francis Hemming in the period 1943-44: Thoughout 1943 and 1944, the Secretary was in correspondence in regard to this case with Sir Rickard Christophers and Brigadier Sinton. Full particulars of this correspondence have been given by Mr. Hemming at the appro- priate points in his application. 7. Receipt in 1944 of the Joint Application prepared by Dr. Curtis W. Sabrosky and Dr. Robert L. Usinger: On 22nd August, 1944 the joint application, dated 7th July 1944, prepared by Dr. Curtis W. Sabrosky and Dr. Robert L. Usinger, reproduced in the first section of paragraph 4 of the present Opinion was received in the Offices of the Commission. 8. Arrangements made for the concurrent submission of the independent applications received: In a letter dated 6th September 1944, Dr. Robert L. Usinger wrote :— ... Our paper in Science, [i.e. the joint application by Dr. Sabrosky and Dr. Usinger reproduced in paragraph 4(1) of the present Opinion] will serve as a supporting appeal from an entirely independent source. I might add that this support is considerable, since our paper was approved by the leading malariologists of the Public Health Service, Office of Malaria Control in War Areas, by malariologists of the National Institute of Health, and by the Chairman of the Committee on Malaria Terminology of the National Malaria Society. 9. Issue in 1944 of an Appeal to Specialists for advice on the problems involved in the present case: In October 1944 it was decided to take further steps to bring this important case to the notice of interested workers, especially in Europe, where in the 24 OPINIONS AND DECLARATIONS war conditions then obtaining it was unlikely that such workers would have an opportunity of seeing the issue of Science containing the paper by Drs. Sabrosky and Usinger. Accordingly, the following note was prepared by the Secretary and despatched to the serial publications Nature and the British Medical Journal; a copy was also sent to Science. This note was published in the first and third of these periodicals in their respective issues of 2nd December 1944 (Nature 154:701; Science 100:404, 405); It appeared in the British Medical Journal in January 1945 (Brit. med. J. 1945 (4385) :85):— Generic and Specific Trivial Names of the Tertian and Quartan Malaria Parasites By FRANCIS HEMMING Secretary to the International Commission on Zoological Nomenclature The Official List of Generic Names in Zoology was established by the International Congress of Zoology in order to promote stability in zoological nomenclature by placing on record the correct names of the - principal genera in each of the Classes and Orders of the Animal Kingdom, together with their type species. Hitherto names have been placed on the Official List in Opinions rendered by the International Commission on Zoological Nomenclature, but in 1943, the Inter- national Commission decided that it was desirable to make the Official List more readily available and accordingly decided to publish it as soon as possible in convenient book form and with a full index. The preparation of the Official List for publication in this way, which was begun in 1943, involved the checking in detail of all the relevant bibliographical and other references to the generic names concerned and their type species. In the course of this work, errors were detected in a number of the Opinions containing decisions relating to the Official List. These errors are being brought at once to the attention of the International Commission with a view to their rendering an Opinion as soon as possible containing such rectifications as may be necessary. Among the errors detected were errors in the entries in Opinion 104 (published in 1928) relating to the author’s name and date of publica- tion of the generic name for the Malignant Tertian Malaria Parasite (Laverania Feletti & Grassi, 1889). Further, in the case both of this name and of that for the Quartan Parasite (Plasmodium Marchiafava & Celli, 1885) the type species was found to have been cited under a name which was not the correct name under the International Code of Zoological Nomenclature. OPINION 283 Zo The names of these parasites, as recorded in Opinion 104, are the names now universally employed for these species in the enormous medical and technical literature relating to malaria and it would clearly be as wrong as it would be impracticable to attempt to introduce changes in such names merely on grounds of zoological nomenclature. In the present case such changes would be particularly undesirable, since they would involve the transfer of the specific trivial name *“ malariae’ from the Quartan Parasite (on which it was bestowed by Grassi & Feletti, 1890) by which name this species is universally known to the Malignant Tertian Parasite on which in 1881 it had been independently bestowed by Laveran (and by which name this species is never called). Transfer of trivial names in this way cause great confusion and the only solution in such a case is for the International Commission to use its Plenary Powers to suspend the rules in order to validate the names currently in use. It was accordingly decided early in 1943 to invite the International Commission to deal with this question under its Plenary Powers and, for this purpose, a thorough investigation into the highly complicated literature of these names was made, with the assistance of Sir Rickard Christophers and Brigadier J. Sinton, whose paper “‘ The correct name of the Malignant Tertian Malaria Parasite,” published in 1938 (Brit. med. 6. 1938 (vol. 2) 1130-1134) must form the starting point of any work on this subject. In the course of this investigation, names previously overlooked were brought to light and other unsuspected nomenclatorial difficulties were disclosed. A paper setting out in detail the present position under the International Code and containing recommendations to the Commission for placing the whole matter on a satisfactory footing has been prepared and will appear in the next part of the Bulletin of Zoological Nomenclature’, the Official Organ of the International Commission on Zoological Nomenclature. Quite recently the Officer in Charge of Malaria Control in War ~ Areas, Atlanta, Georgia, U.S.A., communicated to the International Commission an application prepared by Drs. Curtis W. Sabrosky and Robert L. Usinger, U.S. Public Health Service, drawing attention to the errors in Opinion 104 and requesting the International Commission to use its Plenary Powers to suspend the rules for the purpose of validating existing nomenclatorial practice in regard to these parasites. This application has since been published in the issue of Science of the Ist September 1944. It is extremely gratifying to the Executive Committee of the International Committee to find that malariologists in the United States, working independently, have reached substantially identical conclusions in regard to this matter, since this should greatly facilitate the early adoption by the International Commission of an Opinion setting this matter at rest once and for all. * Owing to printing difficulties arising in the immediate post-war period the paper here Teferred to was not published in the Bulletin of Zoological Nomenclature. It is published for the first time in the Appendix to the present Opinion. 26 OPINIONS AND DECLARATIONS In order to secure the widest support for the action proposed to be taken, the Executive Committee, on behalf of the International Com- mission, invite expressions of opinion from specialists, concerned in any aspect of the malaria problem. Such communications, which should be addressed to the International Commission on Zoological Nomenclature at its Publications Office, at 41, Queen’s Gate, London, S.W.7, will at once be published in the Commission’s Official Organ, the Bulletin of Zoological Nomenclature, in order that the whole of the material relating to this case may be before the Commissioners when reaching their decision. 10. Comments elicited by the Public Notices issued in 1944: The publication of the foregoing note elicited letters of support for the action proposed from: (1) Colonel Paul F. Russell, Medical Corps, (Chief, Division of Parasitology, Army Medical School, Army Medical Center, Washington, D.C., U.S.A.); (2) Dr. David Weinman (Columbia University, College of Physicians and Surgeons, De Lamar Institute of Public Health, New York, N.Y., U.S.A.). These letters are reproduced in the immediately following para- graphs. 11. Comment received from Colonel Paul F. Russell: On 27th | January 1945, Colonel Paul F. Russell, Army Medical Corps (Chief, Division of Parasitology, Army Medical School, Army Medical Center, Washington. D.C., U.S.A.) wrote the following letter to the International Commission :— Reference note by Mr. Hemming in Nature of 2nd December 1944 just in hand. I thoughly agree that it would be not only impractical but illogical at this late date to make the changes in generic and in specific trivial names of the tertian and quartan malaria parasites, although such changes might be in strict accord with a narrow reading of the law. It would seem to be sound procedure for the International Commission to use its Plenary Powers to avoid widespread and persisting confusion. 12. Comment received from Dr. David Weinman: On 27th February 1945, Dr. David Weinman, M.D. (Columbia University, OPINION 283 OF College of Physicians and Surgeons, De Lamar Institute of Public Health, New York, N.Y., U.S.A.) addressed the following letter to the Secretary to the Commission:— Thank you for sending the reprint on ‘* Generic and Specific Trivial Names of the Tertian and Quartan Malaria Parasites.” I am in whole-hearted agreement with your position as stated in this article to wit: that transfer of the specific name malariae from the guartan to the malignant tertian parasite would be the cause of great confusion. This seems to be one of those cases foreseen in the section “Suspension of Rules in Certain Cases,’ where power to suspend the Rules is conferred upon the International Commission when “‘ the strict application of the Rég/es will clearly result in greater confusion than uniformity.” I wish you complete success in this endeavour. 13. Further investigations undertaken by Mr. Hemming: At the beginning of 1945 there still remained a number of difficult questions relating to the names published for—or alleged to have been given to—the human malaria parasites which required elucidation before a comprehensive presentation of the problems involved could be laid before the International Commission. The extreme rarity of some of the serial publications concerned made progress inevitably slow, and it was not until early in 1948 that a conclusion was reached in regard to the last of these problems. The following specialists assisted in the unravelling of the pro- blems involved: (1) Miss I. M. Bellis (The Wellcome Foundation, London); (2) Sir Rickard Christophers, C.I.E., F.R.S. (Depart- ment of Zoology, Cambridge University, Cambridge); (3) Dr. G. Robert Coatney (Assistant Chief, Subdivision on Malaria, Division of Tropical Diseases, National Institute of Health, U.S. Public Health Service, Bethesda, Maryland, U.S.A); (4) Dr. Curtis W. Sabrosky (United States National Museum, Division of Insects, Washington, D.C., U.S.A); (5) Professor Robert L. Usinger (University of California, Division of Entomology and Parasitology, Berkeley, California, U.S.A); Dr. C. M. Wenyon, C.M.G., C.B.E., F.R.S. (The Wellcome Foundation, London). In addition, 28 OPINIONS AND DECLARATIONS Dr. Robert Coatney and Dr. C. F. W. Muesebeck (United States Department of Agriculture, Bureau of Entomology and Plant Quarantine, Washington, U.S.A) were kind enough to furnish photostat copies of some of the rarest of the early papers in which — names had been given to genera or species of human malaria parasites. Dr. Coatney also was good enough to furnish a separate of an important paper not otherwise readily obtainable. To all these specialists grateful thanks are offered by the Com- mission for the assistance so rendered. 14. Receipt of Dr. Sabrosky’s Supplementary Statement: On 10th January 1947 the Commission received from Dr. Sabrosky a copy of the paper containing certain corrections of the account of the history of the generic name Plasmodium Marchiafava & Celli, 1885, given in the application regarding the names of the human malaria submitted by himself and Dr. Usinger in July 1944. As has already been explained, it was subsequently arranged, at the request of Dr. Sabrosky and Professor Usinger, that this paper should be treated as a supplement to their original applica- tion. It has accordingly been reproduced as such in paragraph - 4(2) of the present Opinion. 15. Issue of Public Notices: On Sth November 1947 Notice of the possible use by the International Commission on Zoological Nomenclature of its Plenary Powers in the present case was issued to the serial publications prescribed by the Ninth International Congress of Zoology, Monaco, 1913. The publication of the Notices elicited no objection to the action proposed. 16. Completion of examination of bibliographical problems: The completion early in 1948 of the consultations referred to in paragraph 13 made it possible for Mr. Hemming to complete the outstanding portions of his survey of the names given, or alleged to have been given, to the human malaria parasites which formed the basis of his application to the Commission. This document was completed on 23rd April 1948. Mr. Hemming’s paper is given in the Appendix attached to the present Opinion. OPINION 283 29 III.—THE DECISION OF THE INTERNATIONAL COMMISSION ON ZOOLOGICAL NOMENCLATURE 17. The two applications submitted in the present case were considered by the International Commission on Zoological Nomenclature at the Fourteenth Meeting of its Paris Session held at the Sorbonne in the Amphithéatre Louis-Liard on Monday, 26th July 1948 at 2030 hours. The following is an extract from the Official Record of the Proceedings of the International Commission, summarising the discussion which then took place (1950, Bull. zool. Nomencl., 4:594-612) :— On being invited by the Acting President to open the discussion on the proposal which, jointly with Dr. C. W. Sabrosky, he had submitted to the Commission on this subject, ALTERNATE COMMISSIONER ROBERT L. USINGER (U.S.A.) said that when he had taken part in preparing that application, he had examined the nomenclatorial problems involved and had satisfied himself that the action under the Plenary Powers there recom- mended was necessary, if the gravest confusion in malariological literature was to be avoided. The detailed considerations in- volved were, however, no longer fresh in his memory. He would therefore greatly prefer that this problem should be placed before the Commission by the Acting President, who, he knew, was thoroughly familiar with every aspect of this case. * * * * * [Here followed (1950, Bull. zool. Nomencl., 4:596-612) a summary of the statement made by Mr. Hemming summarising the case as pre- sented in the then unpublished application which he had submitted to the International Commission. For the text of Mr. Hemming’s application see the Appendix to the present Opinion]. * * * * * IN THE DISCUSSION which ensued, the view was expressed on all hands that it was the clear duty of the Commission to use its Plenary Powers in the manner proposed, in order to prevent the catastophic confusion not only in the systematic literature of 30 OPINIONS AND DECLARATIONS Protozoa, but also—and, in this case, this was of much greater — importance—in the vast medical and technical literature of © malariology. The Secretary to the Commission was congratu- lated upon the masterly fashion in which he had assembled the complex mass of data which it was necessary to consider in order to determine the correct position as it existed under the Régles. 18. The following is an extract from the Official Record of the Proceedings of the International Commission, giving the decision reached by it on this case at the foregoing meeting (Paris Session, 14th Meeting, Conclusion 59) (1950, Bull. zool. Nomencl., 4: 594-624) :— THE COMMISSION agreed :— (1) to cancel the incorrect particulars relating to the generi names Plasmodium and Laverania contained :— (a) in Opinion 104; (b) in consequence of (a) above, in the Official List of — Generic Names in Zoology; (2) to use its Plenary Powers :— (a) to suppress for all purposes the under-mentioned trivial names published for the Malignant Tertian Malaria Parasite :— malariae Laveran, 1881, as published in the - binominal combination Oscillaria malariae ; malariae Marchiafava & Celli, 1885, as pub-— lished in the binominal combination Plasmo- dium malariae ; malariae Feletti & Grassi, 1889, as published in the binominal combination Laverania malariae (in so far as this was published as a new name and not as the trivial name malariae Laveran, 1881); OPINION 283 3] (b) to suppress for all purposes other than Article 35 the under-mentioned trivial names published for the Malignant Tertian Malaria Parasite :— falciforme Antolisei & Angelini, 1890, as pub- lished in the binominal combination Ematozoo falciforme ; laverani Labbé, 1894, as published in the bi- nominal combination Haemamoeba laverani ; falciforme Thayer & Hewetson, 1895, as pub- lished in the binominal combination Haema- tozoon falciforme; (c) to set aside the indication, by monotypy, of Plasmo- dium malariae Marchiafava & Celli, 1885 (the Malignant Tertian Malaria Parasite) as the type species (i) of the genus Plasmodium Marchiafava & Celli, 1884, and (ii) of the genus Hamatophyllum? Metschnikoff, 1887 (the name of which was published as a substitute name (nom. nov.) for Plasmodium Marchiafava & Celli, 1885, in the erroneous belief that that name was not available under the Régles), and in place of the foregoing species to designate Haemamoeba malariae Feletti & Grassi, 1889 (the Quartan Malaria Parasite) to be the type species both of the genus Plasmodium Marchiafava & Celli, 1885, and of the genus Hdmatophyllum Metschnikoff, 1887; | (d) to validate the undermentioned trivial names :— malariae Feletti & Grassi, 1889, as published in the binominal combination Haemamoeba malariae, to be the name for the Quartan Malaria Parasite, notwithstanding the fact that, prior to the suppression under the Plenary Powers of the trivial names consist- ing of the word “ malariae,’’ specified in (a) above, that name had been an invalid secondary homonym; 2 See paragraph 24 of the present Opinion. Ww tN OPINIONS AND DECLARATIONS falciparum Welch, 1897, as published in the binominal combination Haematozoon falci- parum, to be the name for the Malignant Tertian Malaria Parasite: (e) to set aside the indication, bymonotypy, of Laverania malariae, Feletti & Grassi, 1889, or, as the case may be, Oscillaria malariae Laveran, 1881, being names for the Malignant Tertian Malaria Para- site suppressed under (a) above, as the type species of the genus Laverania Feletti & Grassi, 1889, and in the place of the species so named to designate Haematozoon falciparum Welch, 1897, to be the type species of that genus; (f) to validate the generic name Laverania Feletti & Grassi, 1889 (type species, by designation under the Plenary Powers, under (e) above: Haemato- zoon falciparum Welch, 1897, validated under the Plenary Powers, under (d) above, as the name for the Malignant Tertian Malaria Parasite); (3) to declare the under-mentioned generic names to be invalid or not required for the reasons severally stated below against the names in question:— Generic name Reason why generic name cited in Col (1) is invalid or not required (1) (2) Oscillaria Laveran, 1881 Invalid because a homonym of (in so far as Laveran Oscillaria Schrank, 1823, that published this as a new name retaining under Article 1 name and not as Oscill- its rights under Article 34, not- aria Schrank, 1823) withstanding the fact that the genus so named has been transferred to the Vegetable Kingdom; Hdmatophyllum?Metschni- koff, 1887 Haemamoeba_ Feletti Grassi, 1889 Ematozoo Antolisei & Angelini, 1890 OPINION 283 33 Invalid because an objective synonym of Plasmodium Mar- chiafava & Celli, 1885, the two nominal genera having the same nominal species as type species; & Invalid because the type species of this genus (Haemamoeba malariae Feletti & Grassi, 1889) has, under (2)(c) above, been designated under the Plenary Powers to be the type species of the genus having the older name Plasmodium Marchiafava & Celli, 1885; Not required because its type species (Ematozoo falciforme Antolisei & Angelini, 1890) isa subjective synonym of Haema- tozoon falciparum Welch, 1897, designated under the Plenary Powers, under (2) (f) above, to be the type species of the genus Laverania Feletti & Grassi, 1889; Haematozoon Thayer & Not required because its type Hewetson, 1895 species (Haematozoon falci- forme Thayer & Hewetson, 1895) is a subjective synonym of Haematozoon falciparum Welch, 1897, designated under the Plenary Powers, under (2)(f) above, to be the type species of the genus Laverania Feletti & Grassi, 1889; ® See paragraph 24 of the present Opinion. OPINIONS AND DECLARATIONS Haemosporidium Invalid because its type species, Lewkowicz, 1897 Haemamoeba malariae Feletti & Grassi, 1889 (by selection under Article 30, Rules (g) and Opinion 46) is the same nominal species as that which, under the Plenary Powers, has under (2)(c) above, been desig- nated as the type species of the genus having the older name Plasmodium Marchai- fava & Celli, 1885 ; (4) to declare that the under-mentioned trivial names, each of which was published as the name of a new avian parasite but in the description of each of which there appeared an incorrect statement that the parasite in question had been found in the blood of human malaria patients, were not available as trivial names for the Malignant Tertian Malaria Parasite (the parasite misidentified with the avian parasite concerned), these trivial names adhering under the Régles to the avian parasites, from which the original descriptions of these parasites were drawn up by their respective authors :— praecox Grassi & Feletti, 1890, as published in the binominal combination Haemamoeba praecox; immaculata Grassi, 1891, as published in the bino- minal combination Haemamoeba immaculata; (5) to declare that the under-mentioned terms consisting of Latin adjectives published in the genitive case, in agreement not with the generic name (as required by Article 14(1)(a)) but with the specific trivial name, either expressed or understood, were published not as subspecific trivial names for human malaria parasites, but as technical designations for those species and that OPINION 283 35 the Latin adjectives in question accordingly possess no status under the Rég/es as subspecific trivial names :— quartanae Celli & Sanfelice, 1891, as published in connection with the binominal combination Plas- modium malariae ; tertianae Celli & Sanfelice, 1891, as published in connection with the binominal combination Plas- modium malariae; quotidianae Celli & Sanfelice, 1891, as published in connection with the binominal combination Plasmodium malariae; tertianae Kruse, 1892, as published in connection with the binominal combination Plasmodium malariae; quartanae Kruse, 1892, as published in connection with the binominal combination Plasmodium malariae; irregularis Kruse, 1892, as published in connection with the binominal combination Plasmodium malariae; tertianae Lewkowicz, 1897, as published in connection with the generic name Haemosporidium; quartanae Lewkowicz, 1897, as published in connect- ion with the generic name Haemosporidium; undecimanae Lewkowicz, 1897, as published in con- nection with the generic name Haemosporidium; sedecimanae Lewkowicz, 1897, as published in con- nection with the generic name Haemosporidium; vigesimo-tertianae Lewkowicz, 1897, as published in connection with the generic name Haemosporidium; (6) to place on record :— (a) that there was no such generic name as Haemato- monas Osler, 1886, Osler in the passage in question not having published a new generic name but having referred to the previously published name Haematomonas Mitrophanow, 1883; 36 OPINIONS AND DECLARATIONS (b) that the undermentioned generic and trivial names published for human malaria parasites by — Danilewsky, 1891, possessed no status under the Régles, the paper in which they were published — having been declared by the Commission in ~ Opinion 101 to be unavailable for nomenclatorial purposes :— (1) the generic names :— Cytamoeba Danilewsky, 1891 Cytosporon Danilewsky, 1891 Cytozoon Danilewsky, 1891 Haemocytosporon Danilewsky, 1891 (ii) the trivial name hominis Danilewsky, 1891, as published in the binominal combination — Laverania hominis; (c) that the trivial name guartana Labbé, 1894, as pub- — lished as a subspecific trivial: name in the trino- minal combination Haemamoeba laverani var. — quartana, is not required for the Quartan Malaria Parasite, it being a junior subjective synonym of © the trivial name malariae Feletti & Grassi, 1889, “ as published in the binominal combination — Haemamoeba malariae, validated under the Plen- — ary Powers under (2)(d) above; (d) that the under-mentioned alleged trivial names, not having been published, were cheironyms and accordingly possessed no status under the Régles :— irregularis Sakharov, erroneously alleged to have been published in 1892 as a subspecific trivial name in the trinominal combination Haemamoeba febris irregularis ; tropica Koch, erroneously alleged to have been published in the binominal combination — Plasmodium tropica in 1899 ; OPINION 283 37 (7) to declare that the trivial name vivax Grassi & Feletti, 1890, as published in the binominal combination Haemamoeba vivax, is the oldest available trivial name for, and therefore the valid trivial name of, the Benign Tertian Malaria Parasite; (8) to. declare that the trivial name tertiana Labbé, 1894, as - published as a subspecific trivial name in the trinominal combination Haemamoeba laverani var. tertiana, is not required for the Benign Tertian Malaria Parasite, being a subjective synonym of the earlier published trivial name vivax Grassi & Feletti, 1890; (9) to substitute the following particulars in regard to the generic names Plasmodium and Laverania in the Official List of Generic Names in Zoology in place of the particulars deleted therefrom in accordance with (1) above :— Plasmodium Marchiafava & Celli, 1885 (type species by designation under the Plenary Powers: Haema- moeba malariae Feletti & Grassi, 1889) (the Quartan Malaria Parasite); Laverania Feletti & Grassi, 1889 (type species, by designation under the Plenary Powers: Haemato- zoon falciparum Welch, 1897) (the Malignant Tertian Malaria Parasite) (generic name to be used by authors who consider the Malignant Tertian (or Aestivo-Autumnal) Malaria Parasite to be generically distinct from the Quartan Malaria Parasite); (10) to place the undermentioned generic names and alleged generic names on the Official Index of Rejected and Invalid Generic Names in Zoology :— Cytamoeba Danilewsky, 1891 (a name _ possessing no status under the Régles, the Commission having ruled (Opinion 101) that the paper in which it was published is not available for nomenclatorial purposes) ; 38 OPINIONS AND DECLARATIONS Cytosporon Danilewsky, 1891 (a name possessing no status under the Régles, the Commission having ruled (Opinion 101) that the paper in which it was published is not available for nomenclatorial purposes); Cytozoon Danilewsky, 1891 (a name possessing no status under the Régles, the Commission having ruled (Opinion 101) that the paper in which it was published is not available for nomenclatorial purposes); Haemamoeba Feletti & Grassi, 1889 (invalid because a junior objective synonym of Plasmodium Mar- chiafava & Celli, 1885, as defined under the Plenary Powers in (2)(c) above, but available for the purposes of Article 34); Haematomonas Osler, 1886 (a cheironym based upon a misreading of a passage referring to the generic name Haematomonas Mitrophanow, 1883); Hamatophyllum* Metschnikoff, 1887 (invalid because of a junior objective synonym of Plasmodium Marchiafava & Celli, 1885, but available for the purposes of Article 34); Haemocytosporon Danilewsky, 1891 (a name posses- sing no status under the Régles, the Commission having ruled (Opinion 101) that the paper in which it was published is not available for nomenclatorial purposes) ; Haemosporidium Lewkowicz, 1897 (invalid because a junior objective synonym of Plasmodium Mar- chiafava & Celli, 1885, as defined under the Plenary Powers in (2)(c) above but available — for the purposes of Article 34); | Oscillaria Laveran, 1881 (in so far as published by — Laveran as a new name) (invalid as a junior — homonym of Oscillaria Schrank, 1823); | * See paragraph 24 of the present Opinion. OPINION 283 39 (11) to place the undermentioned trivial names on the Official List of Specific Trivial Names in Zoology :— falciparum Welch, 1897, as published in the binominal combination Haematozoon falciparum (the Malig- nant Tertian Malaria Parasite); malariae Feletti & Grassi, 1889, as published in the binominal combination Haemamoeba malariae (the Quartan Malaria Parasite); vivax Grassi & Feletti, 1890, as published in the binominal combination Haemamoeba vivax (the Benign Tertian Malaria Parasite) ; (12) to place the undermentioned trivial names and alleged trivial names on the Official Index of Rejected and Invalid Specific Trivial Names in Zoology:— falciforme Antolisei & Angelini, 1890, as published in the binominal combination Ematozoo falciforme (suppressed under the Plenary Powers, under (2)(b) above, for all purposes other than Article 35); falciforme Thayer & Hewetson, 1895, as published in the binominal combination Haematozoon falci- forme (suppressed under the Plenary Powers, under (2)(b) above, for all purposes other than Article 35); hominis Danilewsky, 1891, as published in the binominal combination Laverania hominis (a name possessing no status under the Régles, the Com- mission having ruled (Opinion 101) that the paper in which it was published is not available for nomenclatorial purposes) ; immaculata Grassi, 1891, as published in the bino- minal combination Haemamoeba immaculata (not applicable to the Malignant Tertian Malaria Para- site, but available for the avian parasite, on which the description by the original author was based); OPINIONS AND DECLARATIONS irregularis Kruse, 1892, as published in connection with the binominal combination Plasmodium malariae (published as a technical designation, not as a trivial name, and in consequence possessing no status under the Régles); irregularis Sakharov, erroneously alleged to have been published in 1892 as a subspecific trivial name in the trinominal combination Haemamoeba febris irregularis (a cheironym possessing no status under the Régles); laverani Labbé, 1894, as published in the binominal combination Haemamoeba laverani (suppressed under the Plenary Powers for all purposes other than Article 35, under (2)(b) above); malariae Feletti & Grassi, 1889, as published in the binominal combination Laverania malariae (in so far as this was a new name and not merely a use of the trivial name malariae Laveran, 1881) (sup- | pressed for all purposes under the Plenary Powers under (2)(a) above); malariae Laveran, 1881, as published in the binominal combination Oscillaria malariae (suppressed for all purposes under the Plenary Powers under (2)(a) above); malariae Marchiafava & Celli, 1885, as published in the binominal combination Plasmodium malariae (suppressed for all purposes under the Plenary © Powers under (2)(a) above); praecox Grassi & Feletti, 1890, as published in the binominal combination Haemamoeba praecox (not — applicable to the Malignant Tertian Malaria — Parasite, but available for the avian parasite, on — which the description by the original authors was — based); OPINION 283 41 quartanae Celli & Sanfelice, 1891, as published in connection with the binominal combination Plas- modium malariae (published as a technical designa- tion, not as a trivial name, and in consequence possessing no status under the Régles); quartanae Kruse, 1892, as published in connection with the binominal combination Plasmodium malariae (published as a technical designation, not as a trivial name, and in consequence possessing no status under the Régles); quartanae Lewkowicz, 1897, as published in con- nection with the generic name Haemosporidium (published as a technical designation, not as a trivial name, and in consequence possessing no status under the Régles); quotidianae Celli & Sanfelice, 1891, as published in connection with the binominal combination Plas- modium malariae (published as a technical designa- tion, not as a trivial name, and in consequence possessing no status under the Régles); sedecimanae Lewkowicz, 1897, as published in connection with the generic name Haemosporidium (published as a technical designation, not as a trivial name, and in consequence possessing no status under the Régles); tertianae Celli & Sanfelice, 1891, as published in connection with the binominal combination Plas- modium malariae (published as a technical designa- tion, not as a trivial name, and in consequence possessing no status under the Régles); tertianae Kruse, 1892, as published in connection with the binominal combination Plasmodium malariae (published as a technical designation, not as a trivial name, and in consequence possessing no status under the Rég/es); OPINIONS AND DECLARATIONS tertianae Lewkowicz, 1897, as published in connection with the generic name Haemosporidium (published as a technical designation, not as a trivial name, and in consequence possessing no status under the Régles); tropica Koch, erroneously alleged to have been published in 1899 as a specific trivial name in the binominal combination Plasmodium tropica (a cheironym possessing no status under the Rég/es); undecimanae Lewkowicz, 1897, as published in con- nection with the generic name Haemosporidium (published as a technical designation not as a trivial name, and in consequence possessing no status under the Rég/es); vigesimotertianae Lewkowicz, 1897, as published in connection with the generic name Haemosporidium (published as a technical designation, not as a trivial name, and in consequence possessing no ~ status under the Rég/es); (13) to place on record their grateful thanks to the proto- zoologists, bibliographers and other specialists who, by furnishing information and advice on systematic and bibliographical questions or by supplying photostat copies of rare papers needed in the course of the present investigation into the nomenclature of the human malaria parasites or otherwise, had contributed to the successful elucidation of the complex problems involved; (14) to congratulate Secretary Hemming on the masterly fashion in which he had marshalled the evidence in the light of which the decisions now taken had been reached; (15) to render an Opinion recording the decisions specified in (1) to (12) above. OPINION 283 43 19. Having taken the decision set out in Point (5) of the Con- clusion quoted from the Official Record in paragraph 18 above, that certain Latin adjectives cited in the genitive singular in gram- matical agreement not (as required by Article 14) with the generic name but with a specific name, either expressed or understood, were not acceptable as specific (or subspecific) names, the Com- mission at the same Meeting (Paris Session, 14th Meeting, Conclusion 60) decided to recommend the International Congress of Zoology to insert words in the Régles for the purpose of making it clear that the publication of a Latin adjective in this way did not constitute the valid publication of that adjective as a specific or subspecific name and therefore that the provisions previously agreed upon for the automatic correction of infringements of certain Articles of the Régles, including Article 14, did not apply in such a case. This recommendation was later approved both by the Section on Nomenclature (1950, Bull. zool. Nomencl., 5 : 121-122) and by the Congress in Plenary Session. The following is an extract from the Official Record of the Proceedings of the Commission setting out the foregoing recommendation (1950, Bull. zool. Nomencl., 4 : 624-625) :— THE COMMISSION agreed:— (1) to recommend that words should be inserted in the Régles to make it clear:— (a) that the provision relating to the automatic cor- rection of orthographical and other infringements of Articles 14-16, 18 and 20, which it had been agreed to recommend should be inserted in the Régles, did not apply to a case where a sub- specific or infra-subspecific form of a species, the trivial name of which consisted of a noun in the genitive case, was denoted by an adjective in grammatical agreement not with the generic name (as required by Article 14) but with the specific trivial name, either expressed or understood; 44 OPINIONS AND DECLARATIONS (b) that an adjective used in the manner indicated in (a) above for the purpose of distinguishing a sub- species or infra-subspecific form did not acquire thereby the status of a subspecific, or infra- subspecific, trivial name; (2) to report to the Section on Nomenclature the recom- mendation specified in (1) above and the conclusions reached in regard to the nomenclature of the human malaria parasites, as recorded in Conclusion 59 above®; (3) to invite the Acting President, in his capacity as Secretary to the Commission, forthwith to submit to the Section on Nomenclature the Report referred to in (2) above. 20. The original references for the names placed on Official Lists and Official Indexes by the Ruling given in the present Opinion are given in paragraph 250 of the-application submitted by Mr. Hemming reproduced in the Appendix to the present Opinion (see pp. 186—188). 21. The decision taken in the present case was reported to, and approved by, the Section on Nomenclature of the Thirteenth International Congress of Zoology, Paris, 1948, at its Sixth Meeting held on Monday, 26th July, 1948 (1950, Bull. zool. Nomencl., 5 : 121-122). 22. The Ruling given in the present Opinion was concurred in by the sixteen (16) Commissioners and Alternate Commissioners present at the Paris Session of the International Commission, namely :-— Beltran vice Cabrera; Boschma; Bradley; di Caporiacco; Hemming; Hindle vice Jordan; Jorge vice do Amaral; Kirby vice Stoll; Lemche vice Dymond; Mansour vice Hanko; Metcalf vice Peters; Riley vice Calman; Rode; Sparck vice Mortensen; van Straelen vice Richter; Usinger vice Vokes. > The Conclusion here referred to has been quoted in full in paragraph 18 of the present Opinion. OPINION 283 45 23. The Ruling given in the present Opinion was dissented from by no Commissioner or Alternate Commissioner present at the Paris Session. 24. When the present case was considered by the International Commission at its Paris (1948) Session, the treatment to be accorded to names based upon words containing letters bearing diacritic marks was regulated by Article 20 of the Régles which provided that in the formation of names consisting of words derived from languages using the Latin alphabet “the exact original spelling, including diacritic marks, is to be retained.” It was for this reason that in the application submitted in this case and in the Commission’s decision on it the German umlaut was retained over the first vowel of the generic name Hdmatophyllum Metschnikoff, 1887, instead of being indicated by the addition of the letter ““e”’ after the letter “a.” At Copenhagen, however, in 1953 the Fourteenth International Congress of Zoology amended the foregoing Article abolishing the use of diacritic marks over letters in words used as zoological names and pre- scribing in the case of a name in which an umlaut had appeared over a vowel at the time of the first publication of the name that the umlaut is to be indicated by the insertion of the letter “‘e”’ after the vowel concerned (1953, Copenhagen Decisions zool. Nomencl. : 57-58, Decision 101). Accordingly, in the Ruling given in the present Opinion the foregoing generic name has been transcribed as Haematophyllum. 25. At the time of the adoption of the Ruling given in the present Opinion, the expression prescribed for the second portion of the binomen which constitutes the scientific name of a species was the expression “ trivial name ”’ and the Official List reserved for recording such names was styled the Official List of Specific Trivial Names in Zoology, the word “ trivial’ appearing also in the title of the Official Index reserved for recording rejected and invalid names of this category. Under a decision taken by the Fourteenth International Congress of Zoology, Copenhagen, 1953, the expression “ specific name” was substituted for the expression “ trivial name ” and corresponding changes were made in the titles of the Official List and Official Index of such names 46 OPINIONS AND DECLARATIONS (1953, Copenhagen Decisions zool. Nomencl. : 21). The changes in terminology so adopted have been incorporated in the Ruling given in the present Opinion. 26. The prescribed procedures were duly complied with by the International Commission on Zoological Nomenclature in dealing with the present case, and the present Opinion is accordingly hereby rendered in the name of the said International Commission by the under-signed Francis Hemming, Secretary to the Inter- national Commission on Zoological Nomenclature, in virtue of all and every the powers conferred upon him in that behalf. 27. The present Opinion shall be known as Opinion Two Hun- dred and Eighty-Three (283) of the International Commission on Zoological Nomenclature. Done in London, this Fifteenth day of May, Nineteen Hundred and Fifty-Four. Secretary to the International Commission on Zoological Nomenclature FRANCIS HEMMING as ‘ 5 ~*~, » : Vrooen ; SOP ee ; : 4) ‘ = en ; : APPENDIX TO OPINION 283 OPINION 283 49 THE HUMAN MALARIA PARASITES A proposal for the stabilisation of nomenclature on the basis of current usage FRANCIS HEMMING, c.M.c., C.B.E. Secretary to the International Commission on Zoological Nomenclature 50 OPINIONS AND DECLARATIONS TABLE OF CONTENTS Introductory Paragraph No. (a) Purpose and Scope of the present paper ... ss 1 (b) Historical narrative ... ATE nas RA oe 3 (c) Nomenclature adopted for the human malaria parasites by the International Commission on Zoological Nomenclature in Opinion 104 ne 11 (d) Action taken in 1944 preliminary to the use by the International Commission on Zoological Nomen- clature of its Plenary Powers for the purpose of validating the names currently used for the human malaria parasites ... se one ae 13 (e) Terminology employed in the present paper uae 15 (f) Lay-out of the present paper ae ae ae 21 (g) Acknowledgements and Thanks ... ie pe 29 Part 1. The specific trivial names bestowed (or alleged to have been bestowed) upon the Human Malaria Parasites in the period 1881-1897 Paragraph No. (1) The specific trivial name malariae as published in combination with the generic name Oscillaria by Laveran in 1881 for the Malignant Tertian Malaria Parasite ... ae a a sais 34 Note on the misuse at the beginning of the twentieth century of the specific trivial name malariae Laveran, 1881, as published in the combination Oscillaria malariae, followed by the total eclipse of that name bes he if = ae 46 OPINION 283 51 Paragraph No. (2) The specific trivial name malariae as published in combination with the generic name Plasmodium by Marchiafava & Celli in 1885 for the sce eee nant Tertian Malaria Parasite... 53 (3) The specific trivial name malariae as published in combination with the generic name Haemamoeba by Feletti & Grassi in 1889 for the car Malaria Parasite ... 78 (4) The specific trivial name malariae erroneously alleged to have been published as a new name in combination with the generic name Laverania by Feletti & Grassi in 1889 for the secs Aa Tertian Malaria Parasite . “ 91 (5) The specific trivial name vivax as published in combination with the generic name Haemamoeba by Grassi & Feletti in 1890 for the wide Tertian Malaria Parasite . " 94 (6) The specific trivial name praecox as published in combination with the generic name Haemamoeba by Grassi & Feletti in 1890, the name of a parasite of birds which has been erroneously applied to the yamenee Tertian Malaria oS ae Be fe. 96 (7) The specific trivial name falciforme as published in combination with the generic name Ematozoo by Antolisei & Angelini in 1890 for the aoa Tertian Malaria Parasite . 2 102 (8) The specific trivial name immaculata as published in combination with the generic name Haema- moeba by Grassi in 1891, the name of a parasite of birds which has been erroneously applied to the Malignant Tertian Malaria Parasite wet (E20 52 OPINIONS AND DECLARATIONS Paragraph No. (9), (10), (11) The terms “ quartanae,” “* tertianae ”’ and “* quotidianae ” erroneously alleged to have been published by Celli & Sanfelice in 1891 as trivial names for the Quartan Malaria Parasite, the Benign Tertian Malaria Parasite and the Malignant Tertian Malaria Parasite respectively, treated by the above authors as varieties (sub- species) of the nominal species Plasmodium malariae Marchiafava & Celli, 1885... ah Was (12) The term “‘ hominis”? as published in combination with the generic name Laverania by Danilewsky in 1891 Abe ths at 2 ee aD (13), (14), (15) The terms “ quartanae, “ tertianae ’’ and ‘‘ irregularis ’’ erroneously alleged to have been published by Kruse in 1892 as trivial names for the Quartan Malaria Parasite, the Benign Tertian Malaria Parasite and the Malignant Tertian Malaria Parasite respectively, treated by Kruse as varieties (subspecies) of the nominal pi Plasmodium malariae a 144 (16) The cheironym irregularis erroneously sila to have been published in the combination Haemamoeba febris irregularis by Sakharov in 1892 as a name for the ae Tertian Malaria Parasite... on 15] (17), (18), (19) The aie bad trivial name mierda as published in combination with the generic name Haemamoeba by Labbé in 1894 for the Malignant Tertian Malaria Parasite and the trivial names quartana and tertiana published at the same time by that author for the Quartan Malaria Parasite and the Benign Tertian Malaria Parasite respec- tively, then treated as subspecies of the Bare Tertian Malaria Parasite . e 158 (20) The specific trivial name faleiforme a as publiaita in combination with the generic name Haematozoon by Thayer & Hewetson in 1895 for the a ae Tertian Malaria Parasite af 167 OPINION 283 53 Paragraph No. (21), (22), (23), (24), (25) The terms “‘ quartanae,”’ “ ter- tianae, ” “undecimanae,” “ sedecimanae’”’ and ** vigesimo-tertianae ’’ published by Lewkowicz in 1897 in combination with the generic name Haemosporidium Lewkowicz, 1897, and errone- ously alleged by later authors to have been published as specific trivial names for human malaria parasites ... bit ae ba ek Oe (26) The specific trivial name falciparum as published in combination with the generic name Haematozoon by Welch in 1897 for the See Tertian Malaria Parasite ... " 175 Supplementary note on the cheironym tropica erroneously alleged to have been published in the combination Plasmodium tropica by Koch in 1899 for the Malignant Tertian Malaria Parasite 178 Part 2 The names of the nominal genera established (or alleged to have been established) for the Human Malaria Parasites in the period 1881-1897 Paragraph No. (1) The generic name Oscillaria Schrank, 1823, and the generic name Oscillaria erroneously alleged to have been published as a new name by Laveran in 1881... oe ye > Sts a eh (2) The generic name Plasmodium Marchiafava & Celli, 1885 ba: Bs a ag mn S99 (3) The generic name Haematomonas Mitrophanow, : 1883, and the generic name Haematomonas erroneously alleged to have been published as a new name by Osler in 1886 we il et at) The generic name Make ae llum Metschnikoff, 1887 iQ sip - 209 54 OPINIONS AND DECLARATIONS Paragraph (5) The generic name Haemamoeba Feletti & Grassi, Part 3. Synonymy of the specific trivial names bestowed (or alleged 1889 214 (6) The generic name Laverania Feletti & Grassi, 1889 221 (7) The generic name Ematozoo Antolisei & Angelini, 1890 - om 24 225 (8), (9), (10), (11) The generic names Cytamoeba, Cystosporon, Haemocytosporon and Cytozoon published by Danilewsky in 1891 229 (12) The generic name Haemamoeba as used by Labbé } in 1894 232 @ (13) The generic name MHaematozoon Thayer & | Hewetson, 1895 tae we ‘cee aas | (14) The generic name Haemosporidium Lewkowicz, 1897 241 : to have been bestowed) upon the Human Malaria Parasites in the period 1881-1897 and of the names of the nominal genera established (or alleged to have been established) for those species in the same period Paragraph No. (1) Original references for the names cited in the present paper 250 (2) Synonymy of the specific trivial names bestowed (or alleged.to have been bestowed) upon the human malaria parasites in the period 1881-1897 2 (a) The Malignant Tertian Malaria Parasite ... 252 © (b) The Quartan Malaria Parasite 253 # (c) The Benign Tertian Malaria Parasite 254 @ (3) Names of the nominal genera established (or alleged to have been established) for the human malaria parasites in the period 1881-1897 253 OPINION 283 55 Paragraph No. (4) The correct names for the human malaria para- sites under a strict application of the Régles ... 256 (5) Comparison of the names applicable to the human malaria parasites under a strict application of the Régles with the names universally used for those species 543 aa ies ey “a same Zoe Part 4 Proposal for the stabilisation of the names for the Human Malaria Parasites on the basis of current usage Paragraph No. (1) The specific trivial name for the Quartan Malaria Parasite ... -: ee a fe ee (2) The specific trivial name for the Malignant Tertian Malaria Parasite ... od ie a aauirge2 (3) The specific trivial name for the Benign Tertian Malaria Parasite ... # 4) os taoih268 (4) Certain terms consisting of Latin adjectives cited in the genitive case published as technical designa- tions, but not as names, for human malaria parasites... pee pei ct he ON FA i, (5) The generic name Oscillaria as used by Laveran in 1881 ei a het ai Fe ae TG (6) The generic name Plasmodium Marchiafava & Celli, 1885 ne! i. aS ne et (7) The generic name Laverania Feletti & Grassi, 1889 276 Part5 Proposed clarification of Article 14 of the “* Régles ”’ on the question of the status of terms consisting of Latin adjectives cited in the genitive case when published as though constituting names for species or subspecies ae nan £ > 56 OPINIONS AND DECLARATIONS Part 6 Summary of recommendations Paragraph (1) Recommendations for the stabilisation of the names for the human malaria parasites on the basis of current usage (2) Recommendation for the clarification of Article 14 of the Rég/es in relation to the status of terms consisting of Latin adjectives cited in the genitive case when published as though con- stituting names for species or subspecies APPENDICES Appendix 1.—The discovery by Golgi of the Quartan Malaria Parasite. By Colonel Sir Rickard Christo- phers,. C.LE.. O-5.E., FES. = mi Appendix 2.—Reproduction in facsimile of the paper entitled Sui parasiti della malaria by Feletti (R.) & Grassi (B.) as published in the serial publication Riforma medica, Napoli, 1n its issue of 15th January 1890 and of the supplement to the foregoing paper entitled Sui parasiti della malaria: aggiunta alla nota preliminare published in the same serial publica- tion in its issue of Ist March 1890 ... 282 Page 223 225 OPINION 283 57 INTRODUCTORY (a) Purpose and scope of the present paper The purpose of the present paper is to draw attention to the serious discrepancies which exist between the names currently used by malariologists for the human malaria parasites and the names strictly applicable to those species under the Rég/es and, in the light of the evidence so brought forward, to formulate proposals for consideration by the International Commission on Zoological Nomenclature for the use of its Plenary Powers for the purpose of validating existing nomenclatorial practice and, thus, of providing a stable nomenclature for these species. 2. In scope the present paper aims at providing a comprehen- sive survey of all the papers containing new. names—including papers which it has been alleged contain new names—for the human malaria parasites published in the period from 1881, when the first name for any of these parasites was published by Laveran, up to 1897, the year in which the most recent of the names cur- rently in use (the name falciparum Welch) was published. A thorough survey of this kind is necessary in order to unfold the whole of the action under the Commission’s Plenary Powers that will be required if the Commission is to provide an unassail- able juridical position for the names currently in use. Such a Survey is a necessary preliminary for an application to the Commission for the use of its Plenary Powers in relation to any © nomenclatorial problem, but it is all the more necessary in the present case, in view of the great rarity of many of the books and papers involved and of the numerous misstatements on questions of fact with which the literature abounds, as the result, presumably, of authors having accepted at second hand statements published by their predecessors without going back to the original books and papers under discussion. (b) Historical Narrative 3. Consequent upon the decision announced in 1943 by the International Commission on Zoological Nomenclature (1943, Bull. zool. Nomencl. 1 : xxxvi) that the Official List of Generic 58 OPINIONS AND DECLARATIONS Names in Zoology should be published as soon as possible in — book-form, it became necessary for me, as Secretary to the International Commission, to undertake a detailed study of the bibliographical and other problems connected with the names (between 600 and 700 in number) which up to that date had been placed upon the Official List. This was necessary because it is only since 1936 that full references have been included in the Opinions in which such names have been added to the Official — List. 4. Even before I began the detailed study referred to above, I had been aware that the entries in regard to the generic names Laverania and Plasmodium made in the Official List by the Ruling — given in the Commission’s Opinion 104 (1928, Smithson. misc. Coll. 73 (No. 5):25-28) were not only incomplete but in addition contained a number of serious errors. That this was so had come to light in January 1943 on the discovery among certain miscel- laneous documents which had been sent to me by my predecessor Dr. C. W. Stiles of a copy of a very important paper entitled ‘““ The correct names of the malignant tertian parasite’ written © jointly by Colonel Sir Rickard Christophers, C.I.E., O.B.E,~ F.R.S. and Lt.-Colonel (now Brigadier) J. A. Sinton, V.C., M.D., — D.Sc. This paper had been published at the close of 1938 (Brit. — med. J. 1938 (Vol. 2):1130-1134) and from an annexed note by Dr. Stiles had, it appears, been communicated to him by the — first of the two joint authors on some date in 1939. 5. In these circumstances, it appeared to me to be obvious that | the proper course for the Commission was to reconsider the whole question as soon as possible, with a view to correcting the © errors contained in Opinion 104, by the use of its Plenary Powers — for the purpose of preventing the appalling confusion in medical and protozoological nomenclature which would inevitably follow — any material change in the names used for the human malaria — parasites. 6. Before, however, any proposals could be formulated for submission to the International Commission, it was clearly — essential that a thorough examination should be made of the voluminous literature involved, in order to make as sure as OPINION 283 59 possible that the decision now to be taken rested upon firm foundations and covered the whole ground. The literature is very complicated and much of it is extremely scarce, with the result that progress was necessarily slow. Fortunately, however, there existed at the outset the extremely valuable introduction to the subject in the paper by Christophers and Sinton referred to in paragraph 4 above. 7. It was in January 1943 that I began the detailed examination of the literature necessary for the preparation of the present application to the International Commission on Zoological Nomenclature. This task proved much more formidable than even the paper by Christophers & Sinton (1938) had led me to expect, for on several occasions it happened that, when, with the help of Sir Rickard Christophers, I had traced some obscure paper referred to in the literature as having some bearing on the present subject, I found that it contained a reference to some hitherto neglected name or otherwise raised new problems that called for solution. In spite of these difficulties (and others caused by the evacuation of many libraries from London on account of risk of destruction by air attack) the first draft of the paper was com- pleted by the middle of 1944. In preparing it, I had confined myself strictly to a study of the nomenclatorial, bibliographical and similar questions involved, for, not being a specialist in the group concerned, I could have no opinion on any of the associated taxonomic problems. When references to questions of this. character were unavoidable, I based myself upon the views expressed by Christophers and Sinton in 1938. 8. On 22nd August 1944, I received from Dr. Robert L. Usinger (at that time P.A. Sanitarian (R.), i/c Reports Section, U.S. Public Health Services, Malaria Control in War Areas, Atlanta, Georgia, U.S.A.) a letter dated 7th July 1944, with which he enclosed a paper written jointly by himself and Dr. Curtis W. Sabrosky in which those authors applied for the suspension of the Régles in connection with the names of the parasites of human malaria. This paper, which had already been com- municated to Science, was published in that serial publication on Ist September 1944 (Science 100 : 190-192). 60 OPINIONS AND DECLARATIONS 9. The paper by Drs. Sabrosky and Usinger was concerned mainly to point out the errors contained in Opinion 104 and to request the International Commission on Zoological Nomen- clature to use its Plenary Powers to suspend the Rég/es, in order to avoid the confusion that would inevitably follow any attempt © strictly to apply the Régles in this case. It was not designed to enumerate in detail all the names which would need to be sup- | pressed by the International Commission if they were to decide | to accede to the request presented to them. Since it was primarily for the latter purpose that my own paper had been prepared, I came to the conclusion that it would be useful for me to proceed ~ with its submission to the International Commission, notwith-— standing the receipt of the paper directed towards similar ends | prepared by Dr. Sabrosky and Dr. Usinger. It was accordingly arranged between the foregoing specialists and myself that the two papers should be submitted to the International Commission ~ as independent (or rather complementary) applications. 10. Although the line of argument and the conclusions of the two papers on the nomenclatorial issues were substantially identical, there was one important difference on a taxonomic question on which, having no opinion of my own, J had accepted in my paper the views expressed by Christophers & Sinton in — their paper published in 1938 (see paragraph 6 above). This — difference related to the identity of the single nominal species | which in 1885 Marchiafava & Celli had placed in the genus | Plasmodium. On this question Christophers and Sinton had in 1938 taken the view that the species involved was the Quartan Malaria Parasite, whereas in their paper Sabrosky and Usinger | had identified Marchiafava and Celli’s species with the Malignant — Tertian Malaria Parasite. Immediately therefore upon receiving — the Sabrosky /Usinger paper, I informed Sir Rickard Christophers — of this development and asked him to be so good as to re-examine — this particular question. Sir Rickard very kindly complied with — this request and in due course informed me that he now accepted, ~ and concurred in, the conclusions reached by Drs. Sabrosky and Usinger. In a later paper published in 1946, Dr. Sabrosky drew attention to the fact that Marchiafava & Celli, when publishing ~ the generic name Plasmodium, had published also a new name Plasmodium malariae, for the sole species which they included in ~ OPINION 283 61 that genus; at the same time Dr. Sabrosky noted that the material described and figured by Marchiafava & Celli, though pre- dominantly composed of the Malignant Tertian Malaria Parasite, comprised examples also of the Benign Tertian Malaria Parasite. (c) Nomenclature adopted for the human malaria parasites by the International Commission on Zoological Nomenclature in ** Opinion ” 104 11. The generic names Plasmodium and Laverania were placed on the Official List of Generic Names in Zoology by the Inter- national Commission on Zoological Nomenclature in its Opinion 104 published in 1928. That Opinion was not concerned solely with these names but consisted essentially of a list of fifty-seven generic names, of which those with which we are concerned were two, which were then placed on the Official List as being available names of nominal genera, the type species of which had been duly determined in accordance with the provisions of Article 30. The “Statement of the Case” contained no reference to the long and complicated history of the names published for the human malaria parasites. The information furnished in regard to these names in this Opinion was meagre in the extreme. The following is an extract of the particulars regarding these names given in the *“ Summary ” (i.e. in the operative portion of the Opinion, setting out the decision reached by the Commission) :— SUMMARY .—The following............... generic names, with type species cited, are hereby placed on the Official List of Generic Names :— | ee 62 OPINIONS AND DECLARATIONS 12. The foregoing entries were supplemented by the following passages in the section of the Opinion entitled ‘‘ Presentation of the Case ”’:— PRESENTATION OF THE CASE:— ee a) Cee OEE EET EET SEE HEHEHE HEHEHE EH HEHEHE EHH HEHEHE EEE HEHEHE HEHEHE HEHEHE HEHEHE EEE EEE EEE Laverania Grassi & Feletti, 1890a, 60, mt. falcipara Welch, 1897, 36, 47, type host Homo. [For authors who consider the — parasite of aestivo-autumnal malaria generically distinct from that of quartan malaria.] Not Laverania Labbé, 1899a, 82, type ranarum, type host Rana esculenta. Plasmodium Marchiafava & Celli, 1885d, 791, mt. tsd. malariae — (as restricted to quartan fever), type host Homo. (d) Action taken in 1944 preliminary to the use by the Inter- national Commission on Zoological Nomenclature of its — Plenary Powers for the purpose of validating the names — currently used for the human malaria parasites 13. The errors in Opinion 104 relating to the names of the human malaria parasites detected in 1943 and 1944 were so serious that it was immediately evident that only by the use by the Commission of its Plenary Powers could the nomenclature of the © human malaria parasites be placed upon a satisfactory basis. The Plenary Powers Resolution adopted by the International Congress of Zoology at Monaco in 1913 (see Declaration 5, ~ published in 1943, Opinions and Declarations rendered by the International Commission on Zoological Nomenclature 1 : 31—40) — requires that, before the International Commission may use its — Plenary Powers to suspend the Rég/es, it shall give Public Notice of the proposal in question in any two of five serial publications — specified in the Resolution of 1913. Accordingly, in November — 1944 such notices were given to Science and to Nature, two of the — | OPINION 283 63 serial publications specified in the Plenary Powers Resolution and these notices appeared shortly afterwards in those publications. Owing to the European War, it was not possible at that time to give similar notices to the three other serial publications specified in the foregoing Resolution but this action has since been taken!. 14. The formalities required by the Plenary Powers Resolution have thus been complied with and the International Commission is accordingly in a position to use its Plenary Powers forthwith in the present case, if (as I hope) it agrees that this is the proper course to adopt. (e) Terminology employed in the present paper 15. Every specialist who has ever had occasion to revise the nomenclature of a group of closely allied species has experienced the difficulty, when making that revision, of deciding by what name to refer to each species until such time as, by examining the original descriptions of each of the nominal species concerned and, if possible, their type specimens, he has ascertained what is the oldest available name for each of the collective groups which on taxonomic grounds he accepts as constituting separate specific entities. These difficulties are greatly enhanced where, as with the human malaria parasites, one specific trivial name (in the present case, the specific trivial name malariae) has been bestowed independently upon two closely allied species and further where (as also in the present case) that name when published for one of the species concerned has been continuously misapplied to the other species (through a misunderstanding of, or a failure to read, the original description). 1 The procedure to be adopted in dealing with cases involving the possible use of the Commission’s Plenary Powers here described was modified by the Thirteenth International Congress of Zoology, Paris 1948, which decided that in future Public Notice should be given in the Bulletin of Zoological Nomenclature but that, apart from this, the compulsory giving of Public Notice should be limited to two serial publications, one published in Europe, the other in America, the serials in question to be selected from time to time by the Secretary to the Commission (1950, Bull. zool. Nomencl., 4 : 51\—56). The serials then so selected were Nature and Science, and this arrangement is still in force (intl’d. F.H., 10th January 1954). 64 OPINIONS AND DECLARATIONS 16. An indispensable preliminary to any revision of this kind is | the adoption of precise definitions for the expressions used to — denote the various species in the portion of the work which ~ precedes the identification of each of the named forms concerned. — Normally, the safest—and often indeed the only practicable— course is to define each of the expressions accepted as representing — a specific entity by reference to some work in which that organism ~ is described (and, if possible, figured) in such a way as to preclude — the possibility of subsequent misidentification, and, having done — so, to allot to that specific entity a number or letter by which © alone to refer to it until it has been possible clearly to establish — what, under the Régles, are the oldest available generic and © trivial names for that species. | 17. Where the identity of the organisms representing specific — entities is generally accepted by specialists in the group concerned | and where in any given language there exists for each of the species in question a vernacular name, the meaning of which is | clearly understood by those specialists, it may be more convenient | in the early stages of the revision of a group to refer to the various | species by their vernacular names rather than by numbers or by © letters. 18. Vernacular names of this kind exist in the English language | for each of the human malaria parasites discussed in the present — paper and these names have accordingly been adopted in Parts | 1 and 2 of the present paper as the means by which to refer to | the separate specific entities now recognised by malariologists on © taxonomic grounds. | OPINION 283 65 19. In the present paper the three parasite species discussed -are therefore referred to in the following manner :— Name used in the present Identification of taxonomic paper to indicate species unit referred to under the name cited in column (1) (1) (2) Malignant Tertian Malaria | The parasite with crescent- Parasite shaped gametocytes described by Laveran as Oscillaria malariae in 1881 (Nature parasit. Accid. Impaludisme : 87) Quartan Malaria Parasite The parasite of Quartan Fever described and figured by Golgi in 1885 (G. Accad. Med. Torino 33 : 734) Benign Tertian Malaria | The parasite described by Parasite Grassi & Feletti as Haema- moeba vivax in March 1890 (Arch. ital. Biol. 13 : 300) 20. For the purpose of the present paper the vernacular names “Specified in the first column of the foregoing table are used - exclusively in the sense attributed to them in the second column of that table. Wherever therefore it is stated that a given name is a synonym of another name previously published for the same species, that statement represents a subjective taxonomic judgment by whoever is cited in the passage concerned as the authority for the decision to identify with one another the two nominal species in question. Where, in the opinion of competent specialists, that taxonomic judgment is well-founded, the name sunk as a synonym is correctly so sunk for taxonomic purposes. It must be realised however that, while such a judgment implies that the name in question is not required for taxonomic purposes, it does not and cannot affect the nomenclatorial availability of the name con- cerned. A name so sunk is a subjective synonym and, as such, is not required, so long as the subjective judgment which led to Cc 66 OPINIONS AND DECLARATIONS its being sunk as a synonym is accepted as correct. But im- mediately that judgment is reversed on taxonomic grounds, the name in question can once more be brought into use. The only names, rejected as synonyms, which can never be brought to life again in this way are names belonging to the relatively rare classes of case where a rejected name is an objective synonym of a previously published name There are two classes of such synonyms, namely: (1) the name of a genus the type species of — which was cited by the author of the generic name under the — same specific name as was similarly cited for the type species of a — previously established nominal genus having a different name, ~ and (2) the name of a species based upon the same specimen as that on which was based a previously established nominal species — having a different name (f) Lay-out of the present paper 21. Of the names cited in Opinion 104 for the human malaria parasites, the oldest name is “‘ Plasmodium Marchiafava & Celli, 1885” and the most recent is the specific trivial name “ falcipara — Welch, 1897 ” (which was originally published by the binominal — combination Haematozoon falciparum) There is thus a period ~ of twelve years between the dates of publication of these two — names. In actual fact, as we shall see, the Malignant Tertian ~ Malaria Parasite was first named in 1881, when Laveran described ~ it under the name Oscillaria malariae. There is therefore a period of sixteen years between the date on which the Malignant — Tertian Malaria Parasite was first given a scientific name (1881) © and the date when the same species received the specific trivial name falciparum (1897). Accordingly it is the literature which © appeared during these sixteen years which has to be examined in order to determine the nature and scope of the errors contained in Opinion 104. 22. The first Part of the present paper is designed to provide a list in historical sequence of all the specific trivial names — bestowed (or alleged to have been bestowed) between 1881 and 1897 upon the Malignant Tertian Malaria Parasite and upon the Quartan Malaria Parasite. In order to establish the status of | every name which may need to be dealt with by the International — Commission for the purpose of validating the erroneous entries — ——— j | 4 ‘ OPINION 283 67 in Opinion 104, I have included in this list also all the names bestowed during the same period upon the Benign Tertian Malaria Parasite and also upon certain parasites of birds which some authors have claimed to identify with human malaria parasites. 23. Part 2 is devoted to a corresponding examination of every generic name bestowed (or alleged to have been bestowed) upon human malaria parasites during the period 1881-1897. 24. In Part 3 of the paper synonymies are compiled for the Malignant Tertian Malaria Parasite, the Quartan Malaria Parasite and the Benign Tertian Malaria Parasite on the basis of the material assembled in Parts 1 and 2. In the light of these synony- mies, attention is drawn to the extent of the errors embodied in Opinion 104 in regard to the generic names Plasmodium and Laverania and the names of the type species of those genera. 25. Part 4 contains proposals for the stabilisation of the names, both generic and specific, for the human malaria parasites on the basis of current nomenclatorial usage. In Part 5 I submit a proposal for the clarification of certain aspects of Article 14, this being, in my opinion, necessary, in view of certain doubts as to the proper interpretation of that Article which were encountered in the examination of certain terms alleged to have been published as specific or subspecific trivial names for one or other of the human malaria parasites. 26. A summary of the recommendations submitted in Parts 4 — and 5 is given in the concluding Part (Part 6). 27. Two appendices are annexed to the present paper, for the first of which I have to thank Sir Rickard Christophers. Appendix | gives particulars of the early papers by Golgi in which the Quartan Malaria Parasite was first definitely distinguished from the Malignant Tertian Malaria Parasite but in which no name was given to the species so determined. Appendix 2 con- tains reproductions in facsimile of the paper by Feletti & Grassi published first in December 1889 as a preprint (see paragraph 78), and a fortnight later (in January 1890) in the Riforma medica which forms the basis of a large part of the historical background of the nomenclature of the human malaria parasites. I feel that 68 OPINIONS AND DECLARATIONS the reproduction of this paper forms a necessary part of a com-— prehensive review of the nomenclature of these parasites, not only — because of its intrinsic importance but also because, owing to its extreme rarity, it has been seen by very few living malariologists, a fact which has, no doubt, been responsible for a large part of © the misconceptions which have gathered around this paper. The paper consists of two portions, which are reproduced in Appendix — 2 as Document No. 1 and Document No. 2 respectively. Document No. 1 consists of a reproduction of pages 62 to 64 of Issue No. 11 of volume 6 of the Riforma medica which appeared ~ on 15th January 1890. This Document contains the paper entitled “‘ Sui parasiti della malaria’? in which Feletti & Grassi gave the name Haemamoeba malariae to the Quartan Malaria Parasite, which, though recognised as distinct from the Malignant Tertian Malaria Parasite by Golgi some five years earlier (see — Appendix 1), had till then remained without a scientific name. In — the same paper Feletti & Grassi established the nominal genera ~ Laverania and Haemamoeba for the Malignant Tertian Malaria — Parasite and the Quartan Malaria Parasite respectively. This is the paper which, as explained in paragraph 84(1), and (2)(a) below, © was republished in a French translation four months later (in May ~ 1890). It is this French version, in which the foregoing names - again appeared as new names, which, owing to the rarity of © the Italian original, has been relied upon almost exclusively by subsequent workers. Document No. 2 in Appendix 2 consists of © a reproduction of pages 296 and 298 of Issue No. 50 of the same ~ volume of the Riforma medica which appeared on 1st March 1890. These pages contain a further paper by Feletti & Grassi entitled * Sui parasiti della malaria. Aggiunta alla nota preliminare”~ which, as its title indicates, is a supplement to the paper of 15th” January 1890, reproduced as Document No. | referred to above. This second paper, like the first, was republished in a French ™ version in May 1890, and it is from this latter version that this paper is generally known (see paragraph 84(2)(b) below). 28. One of the great difficulties encountered in the preparation — of the present paper has been the extreme rarity of many of the” books and papers in which new names were published for the ~ human malaria parasites, and the conflicting accounts of the contents of those papers given by subsequent authors. It is therefore worth recording that, with the help of the Institutions | OPINION 283 69 and individual specialists enumerated in the immediately following Section, I have been able to examine personally every one of source books and source papers discussed in the present paper, with the single exception of the pre-print issued in December 1889 of the paper in which Feletti & Grassi first established the nominal genus Haemamoeba and gave the name Haemamoeba malariae to the Quartan Malaria Parasite. The issue of the Riforma medica of 15th January 1890 in which this paper was issued in identical form is, however, reproduced in facsimile in Appendix 2 to the present paper. Thus, subject to the foregoing qualification, all the discussions of earlier publications given in the present paper are based upon a first-hand examination of the original sources. (g) Acknowledgments and Thanks 29. Having now completed my review of the literature of the human malaria parasites, I wish to record my grateful thanks to all the institutions and individual! specialists who have assisted me in this task and without whose help it would have been impossible to compile the present paper. First, I owe an altogether special debt of gratitude to Colonel Sir Rickard Christophers, C.I.E., O.B.E., F.R.S. (late Indian Army Medical Service, and now of the Department of Zoology, Cambridge University). Not only does the paper written by Sir Rickard jointly with Brigadier J. A. Sinton, V.C., M.D., D.Sc. in 1938 (paragraph 6 above) form the essential foundation and starting point for any study of the nomenclature of the human malaria parasites, but in addition lam personally peculiarly indebted to Sir Rickard for the unsparing pains which he has been kind enough to take in tracing obscure publications and in unravelling obscure synonymies and also for the immediate response which he has always made to requests for advice on many different problems encountered in the preparation of the present paper. The wide range and scope of the assistance so rendered will be seen from the numerous extracts which I give from letters which I received from Sir Rickard at various stages of the investigation?. 2 Some of the conclusions which emerged from this correspondence have since been published by Sir Rickard Christophers in a paper entitled *‘ Some Remarks on the Nomenclature of the Malaria Parasites of Man” (Christophers, 1945, Rev. Inst. Salubridad y Enfermedades trop. 6(4) : 213—227). 70 OPINIONS AND DECLARATIONS 30. I desire also to express my thanks to the following specialists and bibliographers, each of whom has given the most valuable — assistance, either in the form of advice on particular questions or by furnishing additional data on individual points on which further elucidation was essential or by supplying photostat copies — of rare papers: Miss I. M. Bellis (formerly Librarian to the — Wellcome Bureau of Scientific Research, London); Dr. G. Robert — Coatney (United States Public Health Service, National Institute of Health, Bethesda, Maryland, U.S.A.); Dr. Edward Hindle, — Sc.D., F.R.S. (Zoological Society of London); Dr. C. A. Hoare D.Sc., F.R.S. (Wellcome Laboratories of Tropical Medicine, Lon-— don); Dr. C. F. W. Muesebeck (United States Department of Agriculture, Bureau of Entomology and Plant Quarantine, Washing- ton, D.C., U.S.A.); Brigadier J. A. Sinton, V.C.) Mia agise (War Office, London); Professor Robert L. Usinger (University of — California, Division of Entomology and Parasitology, Berkeley, California, U.S.A); Dr. C. M. Wenyon, C.M.G., C.B.E., F.R-S: (The Wellcome Foundation, London). 31. I have also to express my grateful thanks to the following Institutions for the grant of facilities to study rare works in their ~ libraries and to the Librarians of these Institutions for assistance granted when using these libraries: British Museum (Natural History); London School of Hygiene and Tropical Diseases; Royal College of Surgeons; Royal Entomological Society of London; Royal Society of Medicine; University College, London University; Zoological Society of London. I wish also to take © the present opportunity to thank Mrs. Cornelia Rosner for — locating rare works in the foregoing libraries, for making tran- scripts from those works and generally for the help given by her in this field. 32. Finally, I wish to thank my old friend and colleague Dr. — Karl Jordan, Ph. D., F.R.S. (British Museum (Natural History), Zoological Museum, Tring, Herts., England), President of the — International Commission on Zoological Nomenclature, for the encouragement which he has at all times given in the present — investigation and for the interest which he has shown at every stage of its course. OPINION 283 71 PART 1 THE SPECIFIC TRIVIAL NAMES BESTOWED (OR ALLEGED TO HAVE BEEN BESTOWED) UPON THE HUMAN MALARIA PARASITES IN THE PERIOD 1881-1897 33. In the revision of any group it is necessary, after having determined on taxonomic genus how many species are involved, firmly to establish the identity of each of the nominal species concerned (that is, the identity of each unit which has been treated by any author either as a distinct species to which a new specific name (binominal combination) has been given) or as a new nominal subspecies to which a new subspecific name (trinominal combination) has been given, before it is possible to determine the identity of the type species of the genera which have been established for species of the group concerned. It is for this reason that in the present paper the discussion of the specific trivial names bestowed (or alleged to have been bestowed) upon human malaria parasites precedes the discussion of the generic names similarly bestowed. (1) The specific trivial name ‘ malariae” as published in combination with the generic name “ Oscillaria ” by Laveran in 1881 for the Malignant Tertian Malaria Parasite 34. In 1880 Laveran reported for the first time the discovery in the blood of human patients suffering from malaria of elements which he thought to be parasitic (paper entitled ‘““ Un nouveau parasite trouve dans le sang des malades atteints de fiévre palustre. Origine parasitaire des accidents de limpaludisme,’” 24th December 1880, Bull. Soc. med. Hép., Paris, (2) 17 : 158—164). In this preliminary report Laveran described three forms of the organism which he had discovered, namely (1) crescentic or ovoid bodies which were transparent or colourless, except for certain rounded pigment granules near the centre, (2) bodies which in repose were spherical, transparent and contained a ring of rounded pigment granules of about the same size, but which, when in movement, were surrounded by three of four fine fila- ments which had an active worm-like movement, (3) spherical bodies, slightly granular, and non-motile, which Laveran believed were cadaveric forms of body (2). Laveran did not in this paper give a scientific name to the parasite which he had discovered. 72 OPINIONS AND DECLARATIONS 35. In May 1881 Laveran followed up this preliminary report in a monograph of 104 pages published by J. B. Bailliére et fils, Paris, under the title “‘ Nature parasitaire des Accidents de l’Impaludisme.” In this monograph Laveran gave a fuller des- cription of the parasite which he had discovered. He gave particulars of the three types of body which he had described in his paper the previous year and again came to the conclusion that all three were referable to a single species. On the relationship of that species to other species, Laveran expressed himself as being in doubt, but he finally decided to treat it as a new species and to give it a scientific name. The passage concerned, which is on pages 86 and 87 of his monograph, reads as follows :— [86] A quelle espéce appartient ce nouvel hématozoaire? Ici j’avoue mon embarras et je fais appel aux naturalistes. J’avais d’abord pensé qu’il s’agissait d’un amibe, qui, a l’état de développement complete, était muni de filaments mobiles; nous avons vu en effet, que les corps no. 2, en dehors des mouvements oscillatoires qui leur sont imprimés par les filaments mobiles, présentent assez souvent des mouvements lents, analogues a ceux des amibes; mais j’ai di abandonner cette idée lorsque j’ai eu constaté que les filaments mobiles puvaient se détacher des corps no. 2 et vivre a l’état de liberté dans le sang. Je suppose aujourd’hui que les filaments mobiles renflés légerement a une de leurs extrémités représentent l’état parfait du parasite du sang, et que les corps no. 1, no. 2 et no. 3 ne sont que des especes de poches dans lesquelles ces parasites vivent pendant un certain temps a |’état d’agglomération, d’enkystement. [87] Les corpuscules brillants, arrondis, mobiles que l’on trouv presque toujours dans les préparations de sang provenant des malades atteints de fi¢vre palustre, représentent peut-étre la premiere phase de developpement des élements parasitaires. Ces corpuscules n’ont pas, du reste, de caractere specifique. | Le filaments mobiles des corps no. 2 ont une grande analogie avec desoscillariées. TIlest a noter, que plusieurs observa- teurs, qui ne soupconnaient pas l’existence de ces animalcules dans le sang des malades atteints de fiévre palustre, ont attribué déja un grand role aux oscillariées dans la pathogénie de l’impaludisme. Hallier, le premier je crois, a émis cette opinion, toute théorique d’aillieurs. Le Dr. Schurtz de (Zwickau) cite le fait d’un homme pris de fievre intermittente dans des conditions de salubrité trés bonnes en apparance; ce malade se livrait 4 l’étude des crytogrames et il OPINION 283 73 avait dans sa chambre a coucher vingt-quatre soucoupes renfer- mant des oscillariées (Arch. d. Heilk. 1868, p. 69). Sil était demontre que les filaments mobiles des corps no. 2 sont bien réellement de l’espéce des oscillariées le nom @Oscillaria malariae conviendrait bien au nouvel hematozoaire. “species to which Laveran applied the name Oscillaria malariae, there is a preliminary point of a purely nomenclatorial character which we must pause to examine. In this connection, it will be “noted that the name Oscillaria malariae was not given in an “unequivocal manner to the parasite which Laveran had dis- covered but was bestowed conditionally, Laveran not going further than noting that, if it were to be established that the _ mobile filaments seen in “‘ Body No. 2” (i.e. in the second of the three forms described in his paper) were really a species of * oscillarian,” then the name Oscillaria malariae would be a suitable name for the new Haematozoon. The question which we have to ask ourselves is whether the name Oscillaria malariae is invalidated by reason of having been given in this conditional Manner or whether, notwithstanding the method by which this name was bestowed, it possesses rights under the Law of Priority (Article 25). If we had only the Rég/es to guide us, this question might be one of considerable difficulty, but fortunately the International Commission on Zoological Nomenclature has given a definite Ruling in an exactly similar case. _ 36. Before we consider the question of the identity of the 37. The Ruling given by the International Commission on this question is embodied in its Opinion 49 (1912, Smithson. Publ. 2060 : 112—113). In the case dealt with in this Opinion, an author (Thomas) had described a species which he doubtfully identified with a previously established nominal species (Aphis asclepiadis Fitch) but added that, should the species which he so described prove distinct from Fitch’s species (asclepiadis), “‘ it may be named Siphonophora asclepiadifolii.”” The question put to the Commission was whether the specific trivial name asclepia- difolii Thomas was an available name or whether it was invalidated by the conditional manner in which it had been bestowed. In the latter event, the next oldest specific trivial name for Thomas’s species was, it was stated in the application, “* asclepiadis Cowen, 1895” (that name not being a homonym because originally 74 OPINIONS AND DECLARATIONS described in a different genus and Thomas’s species and Fitch’s species not being regarded as congeneric). The answer to this question, as given by the Commission in the “‘ summary” to Opinion 49 reads: ‘‘ On basis of the data submitted, asclepiadifolii Thomas, 1879, stands in preference to asclepiadis Cowen, 1895.” This Opinion, therefore, lays down the general principle that the availability of a scientific name is not impaired if that name is bestowed conditionally, provided that in other respects the name in question is an available name. We may conclude, therefore, that the name Oscillaria malariae Laveran, 1881, is not invalidated by reason of having been bestowed conditionally?. 38. When we turn to the question of the identity of the parasite to which in May 1881 Laveran applied the specific name Oscillaria malariae, we come up against the first of the main sources of confusion in the nomenclature of the human malaria parasites. Laveran himself was attempting to determine the cause of malaria fever in Man and believed that he had succeeded in so doing, when, as he thought, he established the presence in the blood of malaria subjects of the parasite to which he gave the name Oscillaria malariae (in the passage quoted in paragraph 35 above). He described three forms of the parasite but he considered that — these were all referable to a single species. Further, he certainly did not envisage the possibility that the various forms of malaria might be due to the existence of more than one species of parasite. In view of the fact that since 1881 malariologists have established the existence of a number of distinct species of parasite, each responsible for a particular form of malaria, it is necessary to determine which of these species it was that Laveran in 1881 described under the name Oscillaria malariae. It is necessary also to examine the possibility that what are now regarded as two or more species were included by Laveran in his single nominal ~ species Oscillaria malariae at the time when he published that © name in 1881. 39. Until the publication of Christophers’ and Sinton’s paper — in 1938 (paragraph 6 above) the specific trivial name malariae 3 In 1948 the Thirteenth International Congress of Zoology, on the recom- mendation of the International Commission on Zoological Nomenclature, inserted in the Régles a provision embodying the Ruling given in Opinion 49 (1950, Bull. zool. Nomencl. 4: 144—145). ' ; i 4 . j a 4 OPINION 283 75 Laveran, 1881, as published in the combination Oscillaria malariae, had for more than a generation been universally accepted as applying to the Quartan Malaria Parasite. This view was based upon what were thought to be authoritative determinations by earlier malariologists and was not supported by any re-examina- tion of Laveran’s original description. A critical review was, however, undertaken by Christophers and Sinton who reported (1938 : 1131) as follows: “ After a careful examination of the original publications we have convinced ourselves that Laveran’s. description, given at the time of naming, relates with absolute certainty to the sexual forms of the malignant tertian parasites and to these only.”” An exactly similar conclusion was reached by Sabrosky & Usinger as the result of an independent study carried out by them during the war, when serving in the Malaria Control in War Areas Unit of the United States Public Health Service, these authors summing up their conctusions in the follow- ing words: “ Malariologists are now agreed that Laveran had the sexual forms of this species [i.e. “the parasite of malignant tertian malaria ’’], only, in his first paper cited above ’ (Sabrosky & Usinger, 1944 : 190). 40. These authorities are thus agreed not only that, when the name Oscillaria malariae was first published, it was applied by Laveran to the Malignant Tertian Malaria Parasite, but also that the whole of the material on which this name was based consisted of this parasite and of no other. This latter conclusion is of importance from the nomenclatorial standpoint, for, if it could be shown that, as originally published, the nominal species Oscillaria malariae Laveran, 1881, was a composite species which, though based largely (or almost exclusively) upon the Malignant Tertian Malaria Parasite, was based in part also upon the Quartan Malaria Parasite, an entirely different situation would arise. In that event, it would be possible legitimately to attach the name Oscillaria malariae Laveran, 1881, to the Quartan Malaria Parasite (notwithstanding the fact that Laveran clearly based his descrip- tion mainly upon the Malignant Tertian Malaria Parasite), if it could be shown that the first author to note that Oscillaria malariae Laveran, 1881, was a composite species, had, in his capacity as “ first reviser ’’ (under Article 31), selected the Quartan Malaria Parasite as the species to which the specific trivial name malariae Laveran, 1881, should adhere. It will be convenient 76 OPINIONS AND DECLARATIONS here briefly to summarise the evidence on this subject which leaves no loophole for the belief that, as originally published, the nominal species Oscillaria malariae was a composite species. 41. It is interesting in this connection to compare Laveran’s treatment of the subject in his “‘ Nature parasitaire des Accidents de Il’ Impadudisme ” in which in 1881 he first published the specific name Oscillaria malariae, with his treatment of the subject in his book ‘‘ Du Paludisme et de son Hématozaire ’’ which was pub- lished ten years later (1891) and in which he reviewed the whole subject in the light of the knowledge gained during the intervening period. In the first of these works he recognised, as we have seen (paragraphs 34-35 above), three types of body, all of which he regarded as referable to his Oscillaria malariae. In the second of these books he recognised four types of body. The three types of body recognised in 1881 which are the same as bodies Nos. 1 to 3 recognised in 1891 are, it is now agreed, all forms of the Malignant Tertian Malaria Parasite. 42. It is only in regard to Body No. 4 that any question can ~ arise. Here we have to consider (1) to what species does this type of body belong and (2) was Laveran aware of the existence of this type of body when he wrote his “ Nature parasitaire des Accidents de l’Impaludisme ”’ in the early part of 1881. 43. In their paper of 1938, Christophers and Sinton stated (1938 : 1132, footnote) that the descriptions and figures relating to Body No. 4 given by Laveran in 1891 (in his “ Du Paludisme et de son Hématozaire’’) “ strongly suggest segmenting forms ” of the Quartan Malaria Parasite. More recently, however, Sir Rickard Christophers has somewhat modified this view (in Jitt., 24th March 1944) :— Even in his later treatises it would be very difficult to say that Laveran had definitely singled out any form as quartan. In fact his forms en rosace might from his figures fit much more with segmenting forms of the malignant tertian parasite. This is all the more probable from the fact, noted by Sinton and myself, that Laveran was dealing largely with the parasite as seen in the organs _ in post-mortem material. OPINION 283 77 44. But, whatever the nature of these No. 4 Bodies which _Laveran in 1891 regarded as referable to the species described by him in 1881 as Oscillaria malariae, they can have no relevance to the identity of that nominal species itself, since there is no evidence whatever that Laveran had before him, or indeed had ever seen, any bodies of this type when he published the name Oscillaria malariae in 1881. It is true, as Christophers and Sinton have pointed out (1938 : 1132, footnote) that Laveran himself stated in 1891 (Joc. cit.) that he had seen these No. 4 Bodies in the blood of a case examined in September 1881. This observation was immediately reported in a paper entitled “‘ De la Nature para- sitaire de l’Accident de l’Impaludisme,” which was published in the following month (October 1881, C. R. Acad. Sci., Paris 93 : 627—630). Thus, we have Laveran’s own word that he saw these No. 4 Bodies in September 1881 and we have also his paper reporting this observation in October. But Laveran’s book * Nature parasitaire des Accidents de ’Impaludisme ”’ (in which the name Oscillaria malariae first appeared) was actually published in 1881 as early as the month of May, for we know (Christophers & Sinton, 1938 : 1132, footnote) that on 3rd May 1881 Laveran. presented this book to the Academie de Medicine. In that book Laveran set out to describe the whole of the results so far achieved by him in his investigations of the causes of malaria in Man; he ‘described fully the three types of body which he had found and it is inconceivable that, if he had been aware of a fourth type of © body, he would have omitted to record its existence. From the evidence summarised above, we may conclude (1) that, when Laveran wrote the book in which he first published the name Oscillaria malariae, he had never seen the type of body which he later called Body No. 4, (2) that it was not until September 1881 that he observed these No. 4 Bodies for the first time, and, there- fore (3) that Laveran’s discovery of the No. 4 Bodies took place five months subsequent to the publication (in May 1881) of the book containing the name Oscillaria malariae. ‘We may also conclude that the paper referred to above which appeared in the issue of October 1881 of the Comptes rendus des Séances de l’ Academie des Sciences of Paris was the first occasion on which Laveran recorded the existence of these No. 4 Bodies. 78 OPINIONS AND DECLARATIONS 45. It is perfectly clear from the foregoing evidence that these ~ No. 4 bodies were unknown to Laveran at the time when he wrote — his original description of Oscillaria malariae (which must have been on some date prior to May 1881, by which time the book ~ containing that name is known to have been actually published) — and that the only forms of malaria parasite which were known to — Laveran in the early part of 1881 were the three forms described — in his book as forms of Oscillaria malariae. All these forms, it is now agreed, were forms of the Malignant Tertian Malaria © Parasite. Accordingly, this is the species to which the specific — trivial name malariae Laveran, 1881 as published in the combina- — tion Oscillaria malariae, applies. Being both a nomenclatorially ~ available name (1.e. not a homonym of any previously published — name) and at the same time the first specific trivial name to be © bestowed upon the Malignant Tertian Malaria Parasite, the name malariae Laveran, 1881, is under the Reégles, the valid specific trivial name for this species. For reasons which, apart from the — extraordinary events described in paragraphs 46-51 below, would — be quite inexplicable, the name malariae Laveran, 1881, although — the oldest available name for the Malignant Tertian Malaria — Parasite and, indeed, for any of the human malaria parasites, has completely dropped out of use. Its resuscitation at this late date for the Malignant Tertian Malaria Parasite would, however, — lead to the most serious confusion and there is the strongest possible case for the use by the Internation Commission of its Plenary Powers to prevent this from happening. Note on the misuse at the beginning of the twentieth century of — the specific trivial name ‘‘ malariae’’ Laveran, 1881, as published in the combination ‘‘ Oscillaria malariae ”’, followed by the total eclipse of that name 46. Before we leave the name malariae Laveran, 1881, as” published in the combination Oscillaria malariae, we must pause ~ to examine the causes of the extraordinary misuse to which this ~ name was put some twenty years after it was published and to note ~ how, after this misuse had served its turn, this name was quietly — allowed to drop out of use altogether. For this episode is not — OPINION 283 79 only of interest by reason of its unique character in the history of nomenclature but in addition an understanding of it is necessary in order to follow the history of the other names for the human malaria parasites. 47. In approaching this problem, we have first to note that the specific trivial name malariae was given independently to two different human malaria parasites: first by Laveran in 1881 in the combination Oscillaria malariae to the Malignant Tertian Malaria Parasite (paragraph 45) and, second, by Feletti & Grassi in 1889 in the combination Haemamoeba malariae to the Quartan Malaria Parasite (paragraphs 53-77 below), following the discovery by Golgi in 1885 of the existence of this second species. For reasons on which no light is thrown by the literature the name malariae (as published by Feletti & Grassi) quickly won acceptance for the Quartan Malaria Parasite, while there was a correspondingly rapid decline in the use of the name malariae (as published by Laveran) for the Malignant Tertian Malaria Parasite, a variety of later names being applied to that species, e.g. praecox Grassi & Feletti, 1890, immaculata Grassi, 1891, Javerani Labbé, 1894, and, finally to the exclusion of these names, the name falciparum Welch, 1897. During the same decade the name Laverania Feletti & Grassi, 1889 (usually attributed to Grassi & Feletti and dated 1890) came into general use as the generic name for the Malignant Tertian Malaria Parasite by those malariologists—at that time numerous—who accepted the view advanced by Feletti & Grassi in 1889 that this species was generically distinct from the Quartan Malaria Parasite. A serious problem remained for those specialists who regarded both the toregoing species as belonging to a single genus, for it was generally agreed by this time that the name Oscillaria used by Laveran in 1881 (paragraphs 187-198) was not appropriate and therefore that the name to be used must be Plasmodium Marchiafava & Celli, 1885. The diffi- culty which arose here was that this name was published four years before Golgi first detected the Quartan Malaria Parasite as being distinct from the Malignant Tertian Malaria Parasite and it would therefore clearly be difficult to sustain an argument that the name Plasmodium could be used for the Quartan Malaria Parasite. Nevertheless, if no such argument could be devised, the commonly accepted name Laverania Feletti & Grassi, 1889, 80 OPINIONS AND DECLARATIONS would fall as a junior synonym of Plasmodium Marchiafava & Celli, 1885, which would thus become the generic name for the © Malignant Tertian Malaria Parasite, while it would be necessary to use for the Quartan Malaria Parasite the name Haemamoeba Feletti & Grassi, 1889. The present Régles were not in existence at the time that we are considering but the unwritten code by — which zoologists of that day guided themselves provided that the relative status both of generic names and of specific trivial names should be regulated in accordance with the Law of Priority. — Thus, malariologists who desired to retain (a) the word “ malariae”’ — as the specific trivial name for the Quartan Malaria Parasite and (b) the generic name Laverania Feletti & Grassi, 1889, for the Malignant Tertian Malaria Parasite were confronted with in- superable nomenclatorial difficulties. 48. In these circumstances we have to inquire how it was that © within a few years of the opening of the present century malario- — logists had come to believe (1) that the name malariae Feletti & Grassi, 1889, could properly be used for the Quartan Malaria Parasite by workers who accepted only one genus for the human ~ malaria parasites, notwithstanding the fact that the word malariae — (as from Laveran, 1881) was the oldest available name for another species (the Malignant Tertian Malaria Parasite) in the same | genus (Plasmodium) and (2) that the generic name Laverania Feletti & Grassi, 1889, could be used, in preference to the name © Plasmodium Marchiafava & Celli, 1885, for the Malignant Tertian Malaria Parasite by those specialists who regarded that species as generically distinct from the Quartan Malaria Parasite. Christophers & Sinton (1938 : 1132-1133) have shown that this change of attitude took place in about the year 1900, in which year © Liihe published his work Ergebnisse der neuren Sporozoenforschung and also two papers on this subject in the Zentralblatt fiir Bakterio- ~ logie (1900, Zbl. Bakt. 27 : 367 and 28 : 205), in which he cut the | Gordian Knot by boldly asserting that the type species of the genus — Plasmodium Marchiafava & Celli, i885, was ‘“‘ Plasmodium — malariae Lav., s. str.” and that this species was the Quartan | Malaria Parasite. Two years later (1902, Arb. Gesundhampt. ~ Berl. 19 : 169) this erroneous identification was reinforced by | its adoption by the great authority Schaudinn. ‘ OPINION 283 : 81 _ 49. It is quite impossible to believe that two such eminent authorities as Liihe and Schaudinn could possibly have fallen into such gross errors as those described above. There must therefore be some entirely different explanation of their action in this matter. Nor is it difficult to guess what is the true explanation of their ‘action. In the serious dilemma in which they found themselves they were faced with the choice between on the one hand applying the international rules of zoological nomenclature, thereby losing both the generic name Plasmodium as the generic name ‘for the Quartan Malaria Parasite and also the specific trivial ame malariae for that species and on the other hand dis- ‘regarding the rules of nomenclature. In this serious situation they must have asked themselves the question: Is there no way out of these difficulties? Clearly there was one way and one “way only. This was to claim that Laveran in 1881, like Feletti & Grassi in 1889, had applied the specific trivial name malariae to the Quartan Malaria Parasite, for, if acceptance for such a claim could be secured, that species would then retain its specific trivial name malariae, although in future it would be necessary to _ attribute that name to Laveran, 1881, instead of to Feletti & Grassi _ 1889 (or 1890). The only possible means by which such a trans- _ formation of the scene could be effected was either (1) to claim _ that Laveran’s original description of Oscillaria malariae in 1881 _ showed that that species was a composite species which, in addition to the Malignant Tertian Malaria Parasite, had included _ also the Quartan Malaria Parasite, which, therefore, any subse- _ quent author, acting as the “ first reviser,”’ would be entitled to _ select as the species to which the name malariae Laveran, 1881, should attach, or (2) to claim that, in publishing the name - Oscillaria malariae, Laveran had published not a description of a particular species of malaria parasite but a general description of _~ the parasite’ responsible for malaria in Man, in which case also it could be claimed (though not with any real validity) that a later author, acting as “ first reviser,” could restrict the specific : trivial name malariae Laveran, 1881, to some one of the known _ Malaria parasites, say, the Quartan Malaria Parasite. 50. When we turn back to the papers published by Luhe in 1900, we find that, without a word of explanation, he stated that ‘the species which he called ‘‘ Plasmodium malariae Lav., s. str.’ i was the Quartan Malaria Parasite. This was the statement which 82 OPINIONS AND DECLARATIONS was endorsed (again without comment) by Schaudinn in 1902. — This action by these two authors is undoubtedly the root cause of the misappropriation for the Quartan Malaria Parasite of the name properly belonging to the Malignant Tertian Malaria © Parasite which was to persist unchallenged for the next forty — years. The only conclusion which can be drawn from this episode is that first Lithe and later Schaudinn were more impressed by the | need for (as they thought) stabilising the nomenclature of the | human malaria parasites on the basis of current practice then by the need for the strict application of the international rules of | zoological nomenclature. | 51. The bold plan conceived and carried out by Lithe and Schaudinn was completely successful. The immense prestige of — these authorities at once secured an uncritical acceptance from ~ other malariologists, who were, it may be assumed, only too glad © to adopt without question such highly acceptable conclusions — backed, as they were, with the sanction of the two greatest experts — of the day. In one small respect only did the solution devised — by Lithe and Schaudinn fail to secure an apparently valid basis | for the then-existing nomenclatorial practice; this was that under | that solution it was necessary to attribute the trivial name © malariae, as applied to the Quartan Malaria Parasite, to Laveran, | 1881, instead of to Feletti & Grassi, 1889. It was not long © however, before even this disadvantage was overcome, for, once q the attribution to Laveran of the name malariae as the name for — the foregoing species had served its turn, it was quietly dropped, — Feletti & Grassi (or rather—though incorrectly—Grassi & — Feletti) were restored to their position as the authors of the name | malariae as applied to the Quartan Malaria Parasite, and the — name malariae Laveran, 1881, though a valid name and, indeed, © the oldest name for any of the human malaria parasites, dis-— appeared altogether from current use. : 52. It is interesting to reflect that this conflict between the requirements of stability in nomenclature and strict adherence to | the rules of zoological nomenclature (in regard to the elimination ~ of secondary homonyms) could only be resolved at that time by | an almost certainly deliberate misidentification of one of the nominal species concerned. It was to meet such embarrassing — OPINION 283 83 dilemmas (and, incidentally, to eliminate dangerous temptations of this kind) that at its meeting held at Monaco in 1913 the Ninth International Congress of Zoology conferred upon the Inter- national Commission on Zoological Nomenclature Plenary Powers to suspend the Rég/les (which had been adopted at Berlin in 1901) in any case where, in the judgment of the Commission, the strict application of the Rég/es would clearly result in greater confusion than uniformity. Had these Powers existed in 1900, the difficulty which then confronted Liihe and Schaudinn could have been met by the Commission using these Powers* in such a way as would have legitimately secured the objective sought by those authors, namely the validation for the Quartan Malaria Parasite of the specific trivial name malariae, which could then have continued to be attributed to Feletti & Grassi, its rightful authors. (2) The specific trivial name ‘‘ malariae” as published in combination with the generic name ‘ Plasmodium” by Marchiafava and Celli in 1885 for the Malignant Tertian Malaria Parasite 53. It was only when I was in course of preparing the present paper that I came across for the first time the hitherto entirely overlooked publication of a new specific trivial name as applied to a human malaria parasite, namely malariae Marchiafava & Celli, 1885, as published in the combination Plasmodium malariae. This appeared in the paper entitled “‘ Weitere Untersuchungen uber die Malariainfection ”’ in which Marchiafava & Celli first published the name Plasmodium (1885, Fortschr. Med., 3(24) : 791). “ At its meeting held in Paris in 1948, the Thirteenth International Congress of Zoology, on the recommendation of the International Commission on Zoology, decided to insert a provision in the Rég/es incorporating the Resolution adopted by the Ninth International Congress, Monaco, 1913, granting Plenary Powers to the International Commission to suspend the normal operation of the Rég/es in cases where it was satisfied that “* the strict application of the Rég/es would clearly result in greater confusion than uniformity ” (1950, Bull. zool. Nomencl. 4: 64—66). At Copenhagen in 1953 the Fourteenth International Congress of Zoology, on the recommendation of the International Commission, deleted the restrictive phrase quoted above and in its place inserted a provision prescribing that the Plenary Powers had been granted “for the purpose of preventing confusion and of pro- moting a stable and universally accepted nomenclature’ (1953, Copenhagen Decisions zool. Nomencl. : 23). 84 OPINIONS AND DECLARATIONS 54. I was led indirectly to this discovery by following up the important contribution by Drs. Sabrosky & Usinger (in the paper submitted by them to the International Commission on Zoological Nomenclature in August 1944) on the question of the identity of the species on which in 1885 Marchiafava & Celli had based their genus Plasmodium. Whereas Christophers & Sinton in their paper in 1938 had accepted the view universally held at that time— a view which resulted from the misidentification by Lihe and Schaudinn referred to in paragraphs 46-51 above—that the Quartan Malaria Parasite was the type species of the genus Plasmodium Marchiafava & Celli, Sabrosky & Usinger had reached the conclusion that the type species of that genus was not the Quartan Malaria Parasite but the Malignant Tertian Malaria Parasite. 55. While, as I realised, the taxonomic interpretation of what- ever Marchiafava & Celli might have written about the unit which they placed in their new genus Plasmodium was a matter upon which only malariologists could pronounce, I considered it desirable, as a first step, myself to examine the passage in the — foregoing authors’ paper of 1885, in order that I might satisfy — myself on any purely nomenclatorial issues which I might find to be involved. The results of this examination were so unexpected that I quote in full the passage in which Marchiafava & Celli introduced the generic name Plasmodium:— Extract from page 791 of the paper published in 1885 in which Marchiafava & Celli established the nominal genus “‘ Plasmodium” Aus dem gesagten geht hervor, dass die beschriebenen K6rper- chen nicht verwechselt werden diirfen mit irgendwelchen zufalligen oder pathologischen Dingen die man bisher in den roten Blutscheiben bemerkt hat Hier handelt es sich um KGrperchen, welche frei im Protoplasma der roten Blutscheibe befindlich sind, die sie, wie wir unten nadher sehen werden, verlassen k6nnen, um KOrperchen welche aus Teilchen des Protoplasmas ohne ausges- prochene Structur bestehen, und die mit lebhafter amdboider Bewegung begabt sind. Und wenn wir diese Constitution, diese Lebensdusserungen bedenken, wenn wir an den innigen Zusammenhang denken, welcher zwischen ihren Wirkungen und der Pathologie der Malariainfection (Melanaemie, Oligaemie, etc.) besteht und an ihre ausschliessliche und constante Gegenwart im OPINION 283 85 Malariablut, welches unzweifelhaft die Fahigkeil besitzt, die Krankheit zu iibertragen, so scheint es uns nicht fernliegend, sie als parasitare Organismen anzusprechen und ihnen den Namen Plasmodium Malariae zu geben. Wir glauben nicht, dass dieser Name vollkommen geeignet ware, und keiner K ritik unterworfen werden kénnte, aber wir haben, wie wir gestehen miissen, keinen besseren dafiir finden k6nnen. Es ist gesagt worden dass diese einfachsten Organismen keine ausgepragte Structur besitzen, wir miissen noch hinzufiigen dass dieselben in gewissen Phasen ihrer Bewegungen und wenn sie unbeweglich sind, eine periphere dicke und glanzende Partie zeigen und eine innere weniger glainzende, welche bisweilen feinkérnig erscheint. Die periphere glanzende Partie ist die, von der die Fortsatze oder Pseudofiisse ausgehen; die letzteren erscheinen in der Tat stets sehr gl4nzend und farben sich intensiver, wahrend die centrale Partie entweder so diinn ist, dass sie das Protoplasma der roten Blutscheibe durchscheinen lasst, oder den Eindruck eines hellen oder leichtgefarbten Kernes von deutlicher Begrenzung darbietet. 56. The above passage brought to light an entirely new fact, hitherto overlooked by malariologists, namely that, when Marchiafava & Celli originally described their new genus Plas- modium, they at the same time applied a new scientific name, Plasmodium malariae, to the taxonomic unit which they placed in that genus. It had always been assumed by malariologists that, in establishing the new genus Plasmodium, Marchiafava & Celli had not used a scientific name for the species which they placed in it. Since, prior to Sabrosky & Usinger’s paper of 1944, the | genus Plasmodium Marchiafava & Celli, 1885, was assumed to be a monotypical genus based upon the Quartan Malaria Parasite, its type species had commonly been cited either as Oscillaria malariae Laveran, 1881, or as Haemamoeba malariae Feletti & Grassi, 1889. Sabrosky & Usinger, who treated the species con- cerned as being the Malignant Tertian Malaria Parasite and not the Quartan Malaria Parasite, also assumed that no scientific name had been used by Marchiafava & Celli for the sole species included by them in Plasmodium and referred to that species as Oscillaria malariae Laveran, 1881, citing this, the oldest name for the Malignant Tertian Malaria Parasite, as the name of the type Species of the genus Plasmodium. Later on, however, this question was re-examined by these authors, and in another paper of October 1946 (Science 104 (No. 2704) : 401—2), Dr. Sabrosky 86 OPINIONS AND DECLARATIONS himself drew attention to the publication by Marchiafava & Celli of the specific name Plasmodium malariae Marchiafava & Celli, 1885. He quoted an extract from the passage (reproduced in paragraph 55 of the present paper), in which Marchiafava & Celli — introduced the name Plasmodium malariae, and added: ‘“ The name was italicised and unquestionably intended as a formal © scientific name. The proposal of Plasmodium as a new generic name has been accepted, but the new specific name has apparently ~ been quite generally overlooked.”’ 57. It was important to discover that, contrary to the univer- sally held belief, Marchiafava & Celli actually applied a scientific name to the species on which their new genus Plasmodium was — based. We have, however, still to consider the question whether the specific trivial name malariae in the combination Plasmodium — malariae was regarded by Marchiafava & Celli as a new specific trivial name appropriated to an entirely new nominal species, or whether they were using the trivial name malariae Laveran, 1881, because they regarded themselves as describing the same species” as that which Laveran had named Oscillaria malariae in 1881, and were merely transferring it to their new genus Plasmodium. In the first case the specific name Plasmodium malariae as used by Marchiafava & Celli would be Plasmodium malariae Marchiafava & Celli, 1885, whereas in the second it would be Plasmodium — malariae (Laveran, 1881). ; F 4 . d a , { 58. This is not an easy question to answer because we know ‘ (a) that Marchiafava & Celli, like Laveran, never considered that | there teem be more than one human malaria parasite, and, like Plasmodium malariae was, in fact, referable to the same species as that previously described by Laveran, namely the Maa Marchiafava & Celli were certainly familiar with the paper if which Laveran had described this species and had given it the name Oscillaria malariae. OPINION 283 87 59. On the other hand, as Sir Rickard Christophers has pointed out to me (in Jitt., 6th October 1944), the bodies described by Marchiafava and Celli in 1885 as Plasmodium malariae were the small amoeboid intracellular asexual forms of the Malignant Tertian Malaria Parasite and were therefore very different from the large flagellating sexual forms of the same species, to which Laveran in 1881 had given the name Oscillaria malariae. It would have been natural therefore from this point of view if Marchiafava & Celli had regarded the new organism which they had discovered as representing an entity specifically distinct from that previously described by Laveran. Moreover, no one, of course, at this time had any idea of the existance of distinct sexual and asexual cycles in this species. 60. It seems clear, therefore, that Marchiafava & Celli must have thought that they were describing an entirely new species. They knew that Laveran thought that he had discovered the parasite responsible for human malaria. They themselves des- cribed what seemed to them to be quite another organism, which they, in their turn, regarded as the organism responsible for malaria. In other words, they must have thought that Laveran was in error in believing that the organism described by him was the organism responsible for malaria. 61. For the new species which they regarded as the organism responsible for malaria, Marchiafava & Celli selected the new specific name Plasmodium malariae. For that new name, they adopted, as the generic name, the word ‘“‘ Plasmodium,” which appeared to them to be suitable in view of the amoeboid character of the organism in question; for the specific trivial name of this organism, they selected the word “ malariae,” believing, as they did, that they had at last discovered the organism really respon- sible for the disease malaria. It is no matter for surprise in such circumstances that the obvious appropriateness of this name outweighed any objections arising on grounds of nomenclatorial priority (if, in fact, such questions as this were ever considered by them at all). 88 OPINIONS AND DECLARATIONS 62. The conclusion that Marchiafava & Celli, in using the name Plasmodium malariae for the species placed by them in their genus” Plasmodium, were definitely introducing a new specific name as well as a new generic name, has been reached independently both by Sir Rickard Christophers and by Dr. Sabrosky as well as by myself. In a letter dated 16th October 1944, Sir Rickard Christophers wrote: ‘I think that Plasmodium malariae M. & C.— must be regarded as probably a new name and not merely the identification by these authors of Oscillaria malariae Laveran’”’;~ and Dr. Sabrosky in his paper of 1946 concluded: ‘“ Careful reading of Marchiafava & Celli (Fortschr. Med., 1885, 3, 791—797) shows that they too were proposing what they regarded as a distinct new form, quite unlike any previously described.” 63. We may take it as established, therefore, that, in introduc- © ing the name Plasmodium malariae, Marchiafava & Celli definitely © intended this to be regarded as a new specific name and not as a | specific name, of which the trivial component (malariae) was the” trivial name malariae Laveran, 1881, brought over into the genus Plasmodium, consequent upon the decision taken by Marchiafava & Celli on taxonomic grounds that this species should be trans-_ ferred to that genus from the genus Oscillaria, in which it had originally been described by Laveran. . 64. We must now revert to the question from which we started the investigation which led to the discovery of the new specific name Plasmodium malariae Marchiafava & Celli, 1885, namely that of the identity of the species on which the new genus Plas- modium had been based. As explained in paragraph 54 above, until 1944 it had been generally assumed that the species in question was the Quartan Malaria Parasite. When I received in | August 1944 the paper by Drs. Sabrosky & Usinger, in which they expressed the view that the type species of the genus Plasmodium Marchiafava & Celli was the Malignant Tertian Malaria Parasite, I immediately wrote to Sir Rickard Christophers (who, in his paper written jointly with Brigadier Sinton in 1938 had treated the Quartan Malaria Parasite as the type species of this genus), and asked him to re-examine Marchiafava & Celli’s original description of Plasmodium in the light of the views expressed by Drs. Sabrosky & Usinger. OPINION 283 89 - 65. In a letter dated 14th September 1944 Sir Rickard hristophers stated that in the paper in which Marchiafava & Celli had established the genus Plasmodium: “ They were des- cribing especially certain small amoeboid rings, some without pigment, and from the plate these might well be Malignant Tertian rings. Also they were describing parasite finds in some 40 cases seen by them in the Pontine Marshes, some of which were cerebral pernicious cases.” Sir Rickard added: ‘* Altogether ‘the impression left with me is that Marchiafava in this paper was mainly dealing with Malignant Tertian.” 66. The use of the word “ mainly ” by Sir Rickard Christophers a this context suggested the idea that, although Marchiafava & Celli themselves were unaware of the existence of more than one human malaria parasite, it was possible that, in fact, they had _more than one species before them when they were establishing their new genus Plasmodium; in other words, that the genus Plasmodium was not monotypical and the nominal species Plasmodium malariae Marchiafava & Celli was a composite - species. 67. This point was brought out specifically by Dr. Sabrosky in his paper of 1946°. He there quotes an analysis by Dr. Martin Young (United States Public Health Service, National Institute of _ Health, Bethesda, Maryland) of Marchiafava & Celli’s detailed descriptions and figures, in which Dr. Young says: “It seems to me definitely that most of the infections that they saw were falciparum. However, they seem occasionally to have run across “a vivax infection. Some of the descriptions are definitely of “wivax segmenters, while others of the descriptions, especially Where they mention the finding of crescents, are definitely falci- parum.” Dr. Sabrosky himself concludes: ‘‘ There seems little doubt, therefore, that Plasmodium malariae M. & C. was based mainly on the malignant tertian parasite (falciparum). The _ benign tertian parasite (vivax) was seen, but there is no evidence _ of quartan.”’ 5 See paragraph 56. 90 OPINIONS AND DECLARATIONS 68. It appears therefore that the nominal species Plasmodium malariae Marchiafava & Celli, 1885, must be regarded as a composite species made up mainly of the Malignant Tertian Malaria Parasite but composed in part also of the Benign Tertian Malaria Parasite. Because the specific name has for so long been overlooked, this nominal species has up till now not been restricted to one or other of the two species of which it is composed (the Malignant Tertian Malaria Parasite and the Benign Tertian — Malaria Parasite), despite the fact these two species have long been clearly distinguished from one another. 69. In order that the nominal species Plasmodium malariae Marchiafava & Celli, 1885, shall become determinate, one of the two species of which it was originally composed must be selected in accordance with the provisions of Article 31 of the Régles®, to be the sole species to which the specific name malariae Marchia- fava & Celli shall adhere. We may now consider what would happen in case the species so selected was (a) the Malignant Tertian Malaria Parasite or (b) the Benign Tertian Malaria Parasite respectively. 70. Taking the first of these possibilities, we find that, if the nominal species Plasmodium malariae Marchiafava & Celli, 1885, were to be restricted to the Malignant Tertian Malaria Parasite, the specific trivial name malariae Marchiafava & Celli, 1885, would fall as a junior synonym, and as a junior secondary homonym, of the specific trivial name malariae Laveran, 1881, 6 At the time when this passage was written, the provisions in the Régles for the selection of a lectotype for a nominal species, the type material of which was con- sidered by later authors to consist of examples of more than one taxonomic species were inadequate in the extreme, the Article concerned (Article 31) reading (in the. then currently accepted English translation of the substantive French text) as follows: “‘ The division of a species into two or more restricted species is subject to the same rules as the division of a genus.”’ A first step towards the improvement of this Article was taken by the Thirteenth International Congress of Zoology, Paris, 1948, when it substituted for the provisions quoted above a new Article setting out in a form appropriate to the designation and indication of holotypes and to the selection of lectotypes for species the Rules given in Article 30 for the designation, indication and selection of type species of genera given in Article 30 (1950, Bull. zool. Nomencl. 4: 74—76). The revision of Article 31 was completed by the Fourteenth International Congress of Zoology, Copenhagen, 1953, when it recast the form of this Article and amended its provisions in certain minor respects (1953, Copenhagen Decisions zool. Nomencl. : 72—78). aa en 9] as published in the combination Oscillaria malariae Laveran, 1881, since the taxonomic units to which these names were re- spectively applied by their original authors have been identified as being referable to ths same species, namely the Malignant Tertian Malaria Parasite. 71. When we come to the second possibility, namely that there might be made a selection under Article 31, by which the name Plasmodium malariae Marchiafava & Celli, 1885, would apply exclusively to the Benign Tertian Malaria Parasite, we are con- fronted with a difficulty arising from the doubts which exist at the present time as to the meaning of Articles 35 and 36 relating to the rejection of junior secondary homonyms and the fact that these Articles are due to be reviewed by the next International Congress of Zoology’. The main question at issue is whether under those Articles the rejection of a specific name on the ground that it is a junior secondary homonym is in all circumstances permanent—as it certainly is in the case of a rejected junior primary homonym—or whether such rejection is permanent only for so long as the condition of secondary homonymy endures, that is, only for so long as two species bearing identical trivial names are judged on taxonomic grounds to belong to the same genus. If the latter view is the correct one or if these Articles were to be so amended by the next International Congress as make that view the correct one, the availability of the name malariae Marchiafava & Celli, 1885, as published in the combina- tion Plasmodium malariae, would, if restricted to apply only to the Benign Tertian Malaria Parasite, depend on the question . of whether the Benign Tertian Malaria Parasite and the Malignant Tertian Malaria Parasite are regarded on taxonomic grounds as being congeneric with one another. In fact, most malariologists today place these two species in the same genus, but we have to consider the nomenclatorial situation also for any malariologists, present or future, who might prefer to treat these two species as belonging to separate genera. ? The doubts as to the meaning to be attached to Articles 35 and 36 in relation to the rejection of secondary homomyms here referred to were completely removed by the series of decisions in relation to the Law of Homonymy taken by the Thirteenth International Congress of Zoology, Paris, 1948, when, after defining the conditions in which any pair of specific names are to be regarded as secondary homonyms of one another, it expressly provided that, once the junior of two second- ary homonyms has been rejected on the ground of that homonymy, the rejection so made shall be permanent (1950, Bull. zool. Nomencl. 4 : 118—125). — 92 OPINIONS AND DECLARATIONS 72. For the majority of malariologists who do regard the Benign Tertian Malaria Parasite as congeneric with the Malignant — Tertian Malaria Parasite, the restriction of the nominal species — Plasmodium malariae Marchiafava & Celli, 1885, to the Benign | Tertian Malaria Parasite would lead to no change in current — practice, for, although the name malariae Marchiafava & Celli, 1885, would then become the oldest specific trivial name for the — Benign Tertian Malaria Parasite, it would fall as a junior second- — ary homonym of malariae Laveran, 1881, the oldest available, and, — therefore the valid specific trivial name of the Malignant Tertian — Malaria Parasite. In consequence, the Benign Tertian Malaria — Parasite would retain its currently accepted specific trivial name, vivax Grassi & Feletti, 1890 (see paragraphs 94-95 below). 73. When we turn to consider the effect of restricting the nominal species Plasmodium malariae Marchiafava & Celli, 1885, to the Benign Tertian Malaria Parasite for any malariologist, present or future, who does not accept the taxonomic judgment under which the Benign Tertian Malaria Parasite has been transferred to the same genus as the Malignant Tertian Malaria Parasite, we at once find ourselves confronted by the doubts — which exist as to the interpretation of Articles 35 and 36 of the Régles to which reference has already been made (paragraph 71)8. Accordingly, in existing conditions it is not possible to be sure — what would be the nomenclatorial effect of the selection, under Article 31, of the Benign Tertian Malaria Parasite to be the species ‘ to which alone the name Plasmodium malariae Marchiafava & Celli, 1885, shall refer. This is rendered all the more doubtful by reason of the fact that this nominal species has been so com- pletely overlooked that it cannot be said as yet that it has been formally rejected as a junior secondary homonym of the name Oscillaria malariae Laveran, 1881, consequent upon the species ~ bearing this latter name having been transferred on taxonomic grounds to the genus Plasmodium. All that has happened so far is that Sabrosky (1944) has expressed the view that the nominal species bearing these names represent the same taxonomic species, an opinion which, as explained in paragraph 67 above, he has 8 As explained in Footnote 7 the doubts here referred to were completely dis- pelled by the action taken in 1948 by the Thirteenth International Congress of Zoology. a a i Dt ace ei a se er ee = OPINION 283 93 ‘since (1946) modified, in the light of the detailed review of the _ paper by Marchiafava & Celli, carried out by Dr. Martin Young, to the extent of agreeing that, while Plasmodium malariae Mar- _chiafava & Celli was based primarily upon asexual forms of the Malignant Tertian Malaria Parasite, it was also—at least to some small extent—based upon specimens of the Benign Tertian Malaria Parasite. 74. It is necessary therefore to take note that, if under any amendment or clarification of the provisions in the Rég/es relating to the rejection of junior secondary homonyms, the name malariae Marchiafava & Celli, 1885, if restricted to the Benign Tertian Malaria Parasite, were to become an available name for that species for any malariologist who did not regard that species as -congeneric with the Malignant Tertian Malaria Parasite, the situation so created would be open to very strong objection®. For _ alone of the specific trivial names for the human malaria parasites, _ the name vivax Grassi & Feletti, 1890, as published in the com- bination Haemamoeba vivax, as applied to the Benign Tertian _ Malaria Parasite, has succeeded in maintaining itself in continuous _ use for nearly sixty years and its supercession at this late stage for _ purely nomenclatorial reasons, while comprehensible to the small number of specialists who have made a study of the nomenclature of the human malaria parasites, would be incomprehensible to, and strongly resented by, the great body of malariologists for whom the names applied to these species are of interest only in so far as they are not subject to change and provide therefore a reliable means by which to refer to these species and to distinguish one from another. The objections to the supercession of the “name vivax Grassi & Feletti, 1890, as the trivial name for the Benign Tertian Malaria Parasite would be a thousand times reinforced, if, as would be the case, the name which replaced vivax consisted of the word malariae (malariae Marchiafava & Celli, 1885), for it is the indiscriminate use of this ill-starred name in the past for both the Malignant Tertian Malaria Parasite and for the Quartan Malaria Parasite that has been the chief of the causes which has led to the present state of confusion in the names of the human malaria parasites. * For the reasons explained in Footnote 7 the contingency here discussed is one which since 1948 could no longer arise. 94 OPINIONS AND DECLARATIONS 75. It should, further, be noted that, if, as the result of the restriction of the nominal species Plasmodium malariae Marchia- fava & Celli, 1885, to the Benign Tertian Malaria Parasite, the name malariae (malariae Marchiafava & Celli, 1885) became the oldest available specific trivial name for that species for malario- logists who do not regard it as congeneric with the Malgnant Tertian Malaria Parasite, a new cycle of confusion would certainly open. For not only would the Benign Tertian Malaria Parasite have to be known by the name malariae (malariae Marchiafava & Celli, 1885) but in addition the name malariae (malariae Feletti & Grassi, 1890) for the Quartan Malaria Parasite would fall as a junior secondary homonym for the entire body of malariologists who regard that species and the Benign Tertian Malaria Parasite as congeneric with one another. 76. In order to avoid the risk, however remote, of such catas- trophic results, I hereby myself, acting as first reviser in accordance with the provisions of Article 31 of the Régles, restrict the name Plasmodium malariae Marchiafava & Celli, 1885, bestowed a composite species consisting of the Malignant Tertian Malaria ~ Parasite and the Benign Tertian Malaria Parasite, to apply solely to the former of these species, namely the Malignant Tertian Malaria Parasite. By this selection under Article 31, the speci- mens of the Malignant Tertian Malaria Parasite described and figured by Marchiafava & Celli, 1885, under the name Plasmodium malariae Marchiafava & Celli, 1885, as noted by Dr. Martin Young (paragraph 67 above) become the sole syntypes of the nominal species so named. | 77. In view of the above selection under Article 31, the specific name Plasmodium malariae Marchiafava & Celli, 1885, applies exclusively to the Malignant Tertian Malaria Parasite and the specific trivial name malariae Marchiafava & Celli, 1885, becomes a junior subjective synonym of malariae Laveran, 1881 (originally published as Oscillaria malariae), which is the oldest available specific trivial name for this species. OPINION 283 95 (3) The specific trivial name “ malariae” as published in com- bination with the generic name “‘ Haemamoeba”’ by Feletti and Grassi in December 1889 for the Quartan Malaria Parasite 78. The specific trivial name malariae was published as a new name in the combination Haemamoeba malariae in a paper by Feletti & Grassi which appeared under the title ‘“‘ Sui parassiti della malaria ” on page 63 in Part 11 of volume 6 of the Riforma medica of Naples. This issue of the Riforma medica is dated 15th January 1890. The above is the place where the name Haemamoeba malariae (and also the generic names Haemamoeba and Laverania) are usually treated as having been first published but it appears that, in fact, the paper by Feletti & Grassi containing these names was published in pamphlet form.in December 1889, i.e. about a fortnight before it appeared in the Riforma medica. This pamphlet is clearly very rare, for in spite of the most careful search Christophers and Sinton (1938 : 1133 note) had to report that they had been unable to locate a copy. Existing information in regard to this pamphlet is derived solely from the statements regarding it made by Blanchard (1905, Les Moustiques : 443, 449 nota). In this work Blanchard gave the following note regarding the first publication of the generic name Laverania, one of the names first published in the foregoing pamphlet by Feletti & Grassi: “R. Feletti et B. Grassi. Swi parassiti della malaria; 8° de 11 p., 22 dec. 1889.—La brochure est datée du 22 décembre, » mais n’a été remise a la poste que la 30 décembre.” Elsewhere Blanchard spoke of this pamphlet as having been “ distribuée le 30 décembre.”’ The specific name Haemamoeba malariae Feletti & Grassi thus ranks for priority as from 30th December 1889. 79. The publication of separates of papers in advance of the appearance of the serial publication in which those papers are destined to be published (as was done by Feletti & Grassi in the present case) is liable to cause serious and unnecessary biblio- graphical difficulties at a later date. For this reason the practice has been severely condemned by the International Commission on Zoological Nomenclature in Declaration 2 (1943, Opinions and 96 OPINIONS AND DECLARATIONS Declarations rendered by the International Commission on Zoo- logical Nomenclature, 1: 7—14)'°. In the present case the fact that no copy of the original issue is available makes it impossible to quote a page reference for the original publication of the name — Haemamoeba malariae (and the generic names Haemamoeba and — Laverania) but otherwise no harm has been done, for the exact date of publication of the original pamphlet is known, thanks to the careful note published by Blanchard. 80. In the present case the position is complicated, however, by the fact that the early volumes of the Riforma medica are also extremely scarce (there being no copy in the United Kingdom) and most workers have had to content themselves with a French mai 2) ay translation published by Feletti & Grassi in volume 13 of the © Archives italiennes de Biologie. This translation appeared on 26th May, 1890. Fortunately, through the kindness of Dr. G. Robert Coatney in furnishing me with a photostat copy, I have been able in the following paragraphs to rely upon the original — text as published in Italian in the Riforma medica on 15th January 1890 and therefore (as explained above) as originally published in — pamphlet form in December 1889. In view of the great import- ance of this paper and the fact that (for the reasons given) it is so little known, I attach a facsimile copy of it as Document 1 in Appendix 2 to the present paper. 81. In this paper Feletti & Grassi reviewed the work of Laveran, — Marchiafava, Celli, Golgi, Guarneri and other contemporary — workers and in the light of this review reached the conclusion ~ that the pigmented bodies characteristic of cases of what are known to be Benign Tertian Malaria and Quartan Malaria (referred to by Feletti & Grassi as “ terzana’’ and “‘ quartana ” respectively) differed from the crescentic (“‘ semiluna’’) type to © an extent which required the establishment for the former of a — 10 At its Session held in Paris in 1948 the International Commission on Zoological — Nomenclature obtained the approval of the Thirteenth International Congress of Zoology for the insertion in the Régles of a Recommendation deprecating the distribution of pre-prints of papers printed for publication in serial publications (1950, Bull. zool. Nomencl. 4 : 168). ee “ OPINION 283 97 new genus to which they applied the name Haemamoeba (loc. cit.: 63, column 2). Having thus formed the view that two species were involved and not one only (as hitherto supposed) and having established a new nominal genus for the newly separated species, it was necessary for Feletti & Grassi to find a specific trivial name for that species. This they did in the concluding paragraph of their paper, which appears on pages 63 and 64 and reads as follows!! :— [63, column 3] In conclusione, sono certamente paras- [64, column 1] siti della malaria le forme descritte da Laveran, Marchiafava, Celli ecc.; secondo ogni verosimiglianza appartengono ai Rizopodi e particolarmente agli Amoebiformes, e ne esistono per lo meno due specie: la Haemamoeba malariae (delle febbri regolari) e la Laverania malariae (delle febbri irregolari). . 82. It will be seen from the foregoing passage that Feletti & Grassi regarded their genus Haemamoeba as containing but a single species, to which they applied the new name Haemamoeba malariae. From their standpoint, therefore, the genus Haema- moeba Feletti & Grassi is monotypical with the above nominal species as type species. 83. It is equally clear, however, from the passage in which Feletti & Grassi established the genus Haemamoeba (quoted in . paragraph 81 above) that their species Haemamoeba malariae was a composite species consisting partly of the Benign Tertian Malaria Parasite (“‘ terzana ”’) and partly of the Quartan Malaria Parasite (“ quartana ”’). 84. It is necessary next to examine the circumstances in which the specific trivial name malariae Feletti & Grassi, December 1889, came to be restricted to one of the two species included in it at the time of its first publication. This restriction was effected in the year 1890 in a paper by the same authors published in volume 13 of the Archives italiennes de Biologie. There has been so 11 For a facsimile of the paper here quoted see Appendix 2, Document 1. D 98 OPINIONS AND DECLARATIONS much misconception regarding the date of publication of this — paper, its authorship and its contents that it requires particularly — careful study. The following are the points which require to — be noted :— (1) The paper which begins on page 287 and ends on page 300 of volume 13 of the Arch. ital. Biol. has as its title: “‘ Sur _ les parasites de la malaria. Notes préliminaires des Prof. R. Feletti et B. Grassi.” This paper appeared in Part 2 of the volume cited above and was published on 26th May 1890, the wrapper to this Part bearing the words: “ Paru le 26 mai 1890.” (2) The foregoing paper consists of four parts, namely :— (a) a portion headed “ Premiére Note” (: 287—293). The ‘‘ Premiére Note” is described as “ Ex- ~ traite de la Riforma medica, No. 11, 15 janvier 1890.” This is a translation into French of the paper originally published in Italian by Feletti & © Grassi, which (as explained in paragraphs 78 and — 79 above) is identical with the pamphlet published — in December 1889 in which those authors first pub- — lished the specific name Haemamoeba malariae. The Italian original of this paper is reproduced as ~ Document No. | in Appendix 2. ! (b) a portion headed “‘ Deuxiéme Note” (: 293—296). The “‘ Deuxiéme Note” is described as ‘“‘ Ex- — traite de la Riforma medica, No. 50, ler mars 1890.” This paper, of which, like the paper published in January 1890, I possess a photostat — copy through the kindness of Dr. Robert Coatney, — was published on pages 296 and 297 of Part 50 of © Volume 6 of the Riforma medica. This paper is reproduced in facsimile as Document 2 in- Appendix 2 to the present paper. It was pub-— lished in Italian under the title “‘ Sui parassiti della malaria. Aggiunta alla nota preliminare.” The authors were cited as “ Prof. R. Feletti e B. OPINION 283 99 Grassi.” In this paper Feletti & Grassi con- tinued the discussion of the characters which they looked upon as distinguishing the genera Laverania and Haemamoeba from one another but did not name any new forms. (c) aportion bearing the title “* Parasites malariques chez les oiseaux. Note préliminaire des Prof. B. Grassi et R. Feletti”’’ (: 297—300). Attached to the title of this portion is a reference to a footnote, which reads as follows: “‘ Bollettino mensile dell’ Accademia Gioenia di scienze naturali in Catania. Fasc. XIII. séance du 23 mars 1890.” This portion also is thus seen to be a translation into French of a paper previously published in Italian by these authors. It must be noted that, unlike the two previous papers, of which the authors were cited as “Feletti & Grassi,” the authors of this third paper were cited as “‘ Grassi & Feletti.” (d) a-single paragraph headed by the letters ““ N.B.”’ but without any title or attribution of authorship (:300). This paragraph, which is quoted in full in paragraph 85 below, is essentially concerned to record a new observation (“‘ Dans ces derniers jours *’) made by the authors concerning the repro- duction of crescent forms even in birds. Since bird parasites are discussed only in the paper entitled (in the French translation) “ Parasites malariques chez les oiseaux’ reproduced as the third portion (portion (c) above) of the composite paper published in the Arch. ital. Biol., it may be concluded that this paragraph, which was here published for the first time, is in the nature of a postscript to the paper, the French translation of which immediately precedes it. It follows, there- fore, that the authorship of this paragraph should 100 OPINIONS AND DECLARATIONS be attributed to Grassi & Feletti, that being the order in which the names of these authors were cited in the paper “* Parasites malariques chez les oiseaux.” 85. It was in the postscript by Grassi & Feletti discussed in sub-paragraph (2)(d) of the preceding paragraph that those ~ authors restricted the specific trivial name malariae Feletti & — Grassi, 1889, as published in the combination Haemamoeba — malariae, to the Quartan Malaria Parasite by distinguishing as a — separate species, and giving the name Haemamoeba vivax to, the second (taxonomic) species which earlier (in December 1889) they — had confounded with the Quartan Malaria Parasite and had ~ included with that species under the name Haemamoeba malariae. — The paragraph by Grassi & Feletti containing this restriction reads as follows :— N.B.—Dans ces derniers jours nous avons constaté que les corps — en croissant se reproduicent méme chez les oiseaux, et que les | amibes, ci-dessus décrites, n’ont rien de commun avec les corps | croissant; elles appartiennent au genre Haemamoeba et nous les | appelons Haemamoeba praecox n.sp. A cette espece se rapportent ~ trés vraissemblablement les Haemamoeba de la fiévre quotidienne ~ avec courtes intermittences. C’est pour cela que nous distinguons — dans le genre Haemamoeba, trois espéces (Haemamoeba malariae — de la fiévre quarte, Haemamoeba vivax de la fiévre tierce et | Haemamoeba praecox de la fiévre quotidienne avec courtes | intermittences, etc.). 3 86. The present Régles were not in existence at the time when ~ Grassi & Feletti published the foregoing note, but in taking this — action as the “ first reviser ’’ of the nominal species Haemamoeba — malariae Feletti & Grassi, 1889, these authors proceeded in strict — accordance with the procedure laid down in Article 31 of the © present Régles. The restriction by Grassi & Feletti of Haema- — moeba malariae Feletti & Grassi, 1889, to the Quartan Malaria © Parasite is therefore perfectly valid under the Régles. , OPINION 283 101 87. It should be noted that, although Feletti & Grassi were the first authors to publish a scientific name for the Quartan Malaria Parasite, this parasite had already been described and figured by Golgi. This important discovery is usually dated from 1889, the year in which Golgi published a paper (1889, Arch. Sci. med. 13 : 173), in which he described in detail the characters which distinguish the Quartan Malaria Parasite from the Malignant Tertian Malaria Parasite. In fact, however (as pointed out by Sir Rickard Christophers in. /itt., 24th March 1944) the first paper in which Golgi established the characters of the Quartan Malaria Parasite was published four years earlier (in 1885, G. Accad. Med. Torino 33 : 734). A note on the rather complicated bibliographical problems raised by these early papers by Golgi, which has been kindly prepared for the present paper by Sir Rickard Christophers, is given in Appendix 1. 88. In none of his papers did Golgi give a scientific name to the Quartan Malaria Parasite. The first authors to give this species such a name were Feletti & Grassi in their pamphlet published in December 1889 (see paragraph 81 above). The specific trivial name so given, malariae Feletti & Grassi, December 1889, as restricted in 1890 by Grassi & Feletti (see paragraph 86 above) is thus the oldest specific trivial name for the Quartan Malaria Parasite. 89. At the time when the specific trivial name malariae Feletti & Grassi, December 1889, was published for the Quartan Malaria Parasite, it was the valid specific trivial name for that species for not only had that species itself not previously been named but the name then given to it (Haemamoeba malariae) was not a homonym of any previously published name. When, however, at a later date the Quartan Malaria Parasite was transferred on taxonomic grounds to the same genus as the Malignant Tertian Malaria Parasite (originally described as Oscillaria malariae Laveran, 1881), the specific trivial name malariae Feletti & Grassi, December 1889, ceased to be available for the Quartan Malaria Parasite, for it became a junior secondary homonym of the specific trivial name malariae Laveran, 1881 (the Malignant Tertian Malaria Parasite) from the point of view of those authors who 102 OPINIONS AND DECLARATIONS accepted the taxonomic judgment that these two species were congeneric. When, however, we turn to consider the position of those malariologists, present or future, who do not accept that judgment and for whom in consequence the specific trivial name malariae Feletti & Grassi, 1889, is not a junior secondary homo- nym of the name malariae Laveran, 1881, as published in the combination Oscillaria malariae, we find ourselves confronted with the same problem as that which met us when we had to consider what would be the relationship between the names malariae Marchiafava & Celli, 1885, as published in the com- bination Plasmodium malariae, and malariae Laveran, 1881, as published in the combination Oscillaria malariae, if an author, acting as a “ First Reviser”’ under Article 31 were to select the Benign Tertian Malaria Parasite to be the species to which alone the name malariae Marchiafava & Celli, 1885, should apply (paragraphs 71-73 above). The question so raised was the inter- pretation to be placed on Articles 35 and 36 of the Régles in relation to the rejection of names on the ground of secondary homonymy. On that occasion it was not necessary to express a view on this admittedly controversial question, for, by then myself acting as a “‘ First Reviser’’ and selecting—under Article | 31—those parts of the material described and figured by Mar- chiafava & Celli which were undoubtedly referable to the Malignant Tertian Malaria Parasite to be the sole specimens by which the name Plasmodium malariae Marchiafava & Celli, 1885, shall be interpreted, that name became a subjective junior synonym of Oscillaria malariae Laveran, 1881. In consequence, in that particular case the difficulties arising from the uncertainty as to the proper interpretation of Articles 35 and 36 ceased to be relevant and it was unnecessary therefore to adopt a working hypothesis on this subject. In the present case, the adoption of such a hypothesis cannot be avoided,'* for, in order to determine the status of the name malariae Feletti & Grassi, 1889, as pub- lished in the combination Haemamoeba malariae, as the name for the Quartan Malaria Parasite, it is essential to take a view on the question whether under the Rég/es the rejection of that name as a 12 The doubts here discussed regarding the meaning to be attached to Articles 35 and 36 of the Rég/es in relation to the rejection of names on the ground of secondary homonymy were, as has been explained in Footnote 7, completely set at rest in 1948, by the revision of these Articles by the Thirteenth International Congress of Zoology. Under that revision, a junior secondary homonym, when once rejected and replaced, can never again validly be used for the species in question. OPINION 283 103 junior secondary homonym of malariae Laveran, 1881, by those specialists who regard these two species as congeneric with one another and who place them both in the genus Plasmodium is permanent in the sense that it is binding also upon other specialists who regard these species as being referable to separate genera or whether, on the contrary, the name malariae Feletti & Grassi, 1889, can properly be used for the Quartan Malaria Parasite by those specialists who regard that Parasite and the Malignant Tertian Malaria Parasite as being generically distinct from one another. While, pending an authoritative clarification of Articles 35 and 36, no view can be held to be correct to the exclusion of every other view, I have myself always been of the opinion that the unqualified nature of the provision in Article 36, coupled with the total absence in Article 35 of any reference to a distinction between primary and secondary homonyms, ‘is such that on any natural construction of the wording of these Articles, the only conclusion to be drawn is that any junior specific homonym, whether primary or secondary, is rendered permanently unavila- able, once it has been rejected and replaced. Owing to the extraordinary misidentification of the nominal species Oscillaria malariae Laveran, 1881, with the Quartan Malaria Parasite (paragraphs 46-51 above), the name malariae Feletti & Grassi, 1889, has customarily been rejected not as a junior secondary homonym of malariae Laveran, 1881, but as a junior subjective synonym of that name. It is likely therefore that the name malariae Feletti & Grassi, 1998 has never as yet been formally rejected as a junior secondary homonym and in consequence, replaced by some other name. Certainly the only authors who in recent times have recognised that the name malariae Laveran, 1881, applies to the Malignant Tertian Parasite and not to the Quartan Malaria Parasite (Christophers & Sinton (1938); Usinger & Sabrosky (1944)), while recognising the existance of a condition of secondary homonymy in the genus Plasmodium between the names malariae Laveran, 1881 (for the Malignant Tertian Malaria Parasite) and malariae Feletti & Grassi, 1889 (for the Quartan Malaria Parasite) have refrained from formally rejecting the latter as an invalid junior secondary homonym of the former, the 104 OPINIONS AND DECLARATIONS reason in each case being the same, namely a reluctance to take any action which would lead to still further confusion in the literature of human malariology. Such action has its value as a temporary measure, pending the use by the International Com- mission of its Plenary Powers for the purpose of preserving existing nomenclatorial practice. It cannot however alter in any way the fact that under the Rég/es the name malariae Feletti & — Grassi, 1889, will cease to be an available name for the Quartan Malaria Parasite for any specialist who regards that species as congeneric with the Malignant Tertian Malaria Parasite, immedi- ately any author (a) accepts the name malariae Laveran, 1881, as the name of the Malignant Tertian Parasite, (b) regards that species and the Quartan Malaria Parasite as congeneric and (c) in consequence rejects the name malariae Feletti & Grassi, 1889, as a junior secondary homonym of malariae Laveran, 1881, and replaces it with the next available name for the Quartan Parasite. Whether or not this rejection would render this name permanently unavailable for use even by specialists who do not regard these two species as being congeneric with one another, is, as we have seen, at present a matter of dispute, though for the reasons which I have explained, I hold that such action would have this effect.!* 90. To sum up, the name malariae Feletti & Grassi, 1889, as published in the combination Haemamoeba malariae, cannot be used by any specialist who regards the species so named (the Quartan Malaria Parasite) as congeneric with the Malignant Tertian Malaria Parasite by reason of the fact that, if this name were to be used in this way, it would immediately become a junior secondary homonym of the name malariae Laveran, 1881, as published in the combination Oscillaria malariae, and in con-— sequence would need to be rejected and replaced by some other ~ name. The position of the name malariae Feletti & Grassi, 1889, — for those specialists who do not regard the species so named (the © Quartan Malaria Parasite) as congeneric with the Malignant Tertian Malaria Parasite is not clear owing to the doubts which ‘8 The assumption here adopted later proved to have been justified, for in Paris ~ in 1948 the Thirteenth International Congress of Zoology when revising Articles 35 and 36 of the Régles, definitely prescribed that a name, once rejected as a secondary | homonym and replaced by another name, is to be treated as being permanently — rejected and therefore incapable of being validly brought into use again by any — later author. See Footnote ’. } OPINION 283 105 exist as to the proper interpretation of Articles 35 and 36 relating to the rejection of secondary homonyms. It is necessary for the purposes of the present study to adopt some working hypothesis on this subject and the conclusion which I have reached—which coincides with the view which I have always held on the question of interpretation referred to above—is that the rejection and replacement of the name malariae Feletti & Grassi, 1889, as a junior secondary homonym in the genus Plasmodium renders that name permanently invalid, not only for those specialists who regard the Quartan Malaria Parasite and the Malignant Tertian Malaria Parasite as congeneric with one another but also for those who do not hold this view!*. For the remainder of the present paper I therefore treat the name malariae Feletti & Grassi, 1889, as published in the combination Haemamoeba malariae, as a permanently invalid name. On this view it is necessary under the Régles to find some later name for the Quartan Malaria Parasite. It must be noted at this point, however, that the name malariae Feletti & Grassi, 1889, has been used continuously for the Quartan Malaria Parasite for over fifty years and that its displacement at this date on purely nomenclatorial grounds would be open to the strongest possible objection. (4) The specific trivial name ‘“ malariae ” erroneously alleged to have been published as a new name in combination with the generic name “ Laverania ” by Feletti and Grassi in 1889 for the Malignant Tertian Malaria Parasite 91. As we have seen in paragraph 81 above, Feletti & Grassi in their paper “‘ Sui parassiti della malaria” (published, first, as a pamphlet on 30th December 1889 and, second, as a paper, in the issue of the Riforma medica of 15th January 1890) recognised two species of human malaria parasite, one of which they referred to as Haemamoeba malariae and the other as Laverania malariae. 92. The generic names Haemamoeba and Laverania were both published in this paper for the first time. So also was the specific ** As will be seen from Footnote 12, the decision taken by the Thirteenth Inter- national Congress of Zoology, Paris, 1948, gave the seal of approval to the working hypothesis here adopted. 106 OPINIONS AND DECLARATIONS trivial name malariae as placed by Feletti & Grassi in their new genus Haemamoeba. On the other hand the choice of the word ** Laverania’”’ for the name of their new genus makes it clear that the purpose of Feletti & Grassi was to give honour to Laveran, the discoverer of the first of the human malaria parasites to be detected (the Malignant Tertian Malaria Parasite). Further, their use for that species of the trivial name (malariae) published for that species by Laveran at the time of his discovery, establishes beyond doubt that the trivial name malariae, as used by Feletti & Grassi in 1889 in combination with the generic name Laverania was not a new name of their own but on the contrary was the name malariae as published by Laveran himself in 1881 in the combination Oscillaria malariae. 93. In a number of synonymies of the names of the human malaria parasites (of which the most recent instance is afforded by Christophers, 1945, Rev. Inst. Salubridad y Enfermedades trop., 6(4) : 220) the specific name (binominal combination) Laverania malariae has been cited as though it were a new name published by Feletti & Grassi for the Malignant Tertian Malaria Parasite in 1889, i.e. as though the specific trivial name malariae as there used is attributable to Feletti & Grassi, 1889, whereas it is in fact attributable (as shown above) to Laveran, 1881. In this way there has grown up the erroneous belief that in 1889 (or in 1890, in the case of those authors who believed that the paper “ Sui parassiti della malaria’ was not published until that year) Feletti & Grassi published the specific trivial name malariae as a new name in combination with the generic name Laverania. (5) The specific trivial name “ vivax ” as published in combination with the generic name ‘“ Haemamoeba” by Grassi and Feletti in 1890 for the Benign Tertian Malaria Parasite 94. The specific trivial name vivax was first published by Grassi — & Feletti (in the combination Haemamoeba vivax) on 26th May 1890 (see paragraph 84(1) above) (Arch. ital. Biol. 13 : 300) in the short note quoted in paragraph 85 above. This was the place where those authors first recognised that the Benign Tertian Malaria Parasite and the Quartan Malaria Parasite were distinct species and gave a specific trivial name (vivax) to the first of these species. As explained in paragraph 83 above, these authors had OPINION 283 107 previously (1889, Sui parassiti della malaria and 15th January 1890, Riforma medica 6(11) : 64) included both these species in their (at that time) composite species Haemamoeba malariae Feletti & Grassi, 1889. 95. The specific trivial name vivax Grassi & Feletti, 1890, as published in the combination Haemamoeba vivax, is the first such name to have bestowed upon the Benign Tertian Malaria Parasite and, as it is a nomenclatorially available name, it is the valid specific trivial name for that species. (6) The specific trivial name “‘ praecox ” as published in combina- tion with the generic name ‘‘ Haemamoeba ” by Grassi and Feletti in 1890, the name of a parasite of birds which has been erroneously applied to the Malignant Tertian Malaria Parasite 96. The specific trivial name praecox was published by Grassi & Feletti in 1890 in the combination Haemamoeba praecox. This name appeared in a brief postscript (written in French) attached by Grassi & Felletti to a translation into French (with the title ** Parasites malariques chez les oiseaux ’’) of a paper (written in Italian) which those authors had presented to the “‘ Accademia Gioenia di scienze naturali in Catania” at the meeting of that body held on 23rd March 1890 and which had just appeared in Fascicule 13 of its Bollettino mensile. The French translation of this paper, together with the brief postscript then published for the first time (on page 300), appeared in Part 2 of volume 13 of the Archives italiennes de Biologie. The exact date of publication is known, for fortunately the wrapper of this Part bears the legend ** Paru le 26 mai 1890." The original reference for the nominal species Haemamoeba praecox Grassi & Feletti is thus ““ May 1890, Arch. ital. Biol. 13(2) : 300.” 97. The text of the passage containing the original publication of the name Haemamoeba praecox Grassi & Feletti, 1890, is reproduced in full in paragraph 85 of the present paper. In order to understand the description given by Grassi & Feletti for this species, it is necessary to be aware of the substance of the paper to which (as already explained) the passage containing the original. 108 OPINIONS AND DECLARATIONS description of Haemamoeba praecox was a postscript. The description there given leaves no room for doubt that the fore- going name was introduced for the express purpose of denoting certain parasites found in the blood of birds which had been described in the paper entitled ‘* Parasites malariques chez les oiseaux,’ to which the note containing the description of Haema- moeba praecox was, as has been explained, added as a postscript to the French version of the foregoing paper. It was only after the name Haemamoeba praecox had been given in this way to the bird parasite that in the following sentence Grassi & Feletti added, by way of, what they regarded as an important, comment, their view that the bird parasite to which they had given the foregoing name was the same as that which in Man was responsible for ‘la fiévre quotidienne avec courtes intermittences.”” The human parasites identified in this way by Grassi & Feletti were, it is now generally agreed, asexual forms of the Malignant Tertian Malaria Parasite. As has been pointed out by Christophers & Sinton (1938 : 1132), Grassi & Feletti, when they wrote the paper now under discussion, were unaware that these asexual forms of the human parasite were referable to the same species (the Malignant Tertian Malaria Parasite) as were the sexual forms to which nine years earlier Laveran had (in 1881) given the name Oscillaria malariae. Grassi & Feletti were familiar with these asexual forms in human cases and it was, no doubt, because they recognised the resemblance between these forms and the bird parasite which they had just named Haemamoeba praecox that, as we have seen, they added the comment that, in their view these asexual forms, found in human cases belonged to the same species as the bird parasite to which they given the name Haemamoeba praecox. 98. The specific trivial name praecox Grassi & Feletti was later extensively used as the specific trivial name of the Malignant - Tertian Malaria Parasite by French workers who uncritically followed the erroneous determination by Lihe (1900) (discussed in paragraph 49 above) of Oscillaria malariae Laveran, 1881, as the Quartan Malaria Parasite. Workers in France who accepted Liihe’s determination of Laveran’s species, being thus forced to find a specific trivial name in place of malariae for the Malignant Tertian Malaria Parasite, adopted for this species the specific trivial name praecox Grassi & Feletti, 1890. The French workers OPINION 283 109 who adopted this course argued, first, that the nominal species - Haemamoeba praecox, as established by Grassi & Feletti in 1890, "was a composite species, made up partly of a new bird parasite and partly of asexual forms of the Malignant Tertian Malaria Parasite of Man. Second, it was argued that this nominal species had ceased to be a composite species in 1891 when Grassi & Feletti had removed the bird parasite from it, on renaming the bird parasite Haemamoeba relicta (Grassi & Feletti, 1891, Bull. mens. _ Accad. Gioenia Sci. nat. Catania (n.s.) 1 (No. 16) : 19). Accord- ingly, under this argument, the Malignant Tertian Malaria Para- site had been left in undisputed possession of the trivial name _ praecox Grassi & Feleiti, 1890. This argument would have been valid only (i) if, in fact, examples of the Malignant Tertian Malaria Parasite had been included among the syntypes of the “nominal species Haemamoeba praecox Grassi & Feletti, 1890, and (ii) if in 1891 Grassi & Feletti had effectively selected the foregoing syntypes to form the basis for the nominal species _ Haemamoeba praecox to the total exlusion of the syntypes of the bird parasite by removing the latter to the new nominal species _ Haemamoeba relicta. 99. We may conveniently examine first the second of these claims. On this subject, I applied for assistance to Dr. C. A. Hoare, F.R.S. (The Wellcome Laboratories of Tropical Medicine, London). After examining the whole of the papers concerned, Dr. Hoare replied as follows :— | I shall now deal with the points raised by you in chronological order. (a) Grassi, B. & Feletti, F. ‘* Parassiti malarici negli uccelli. Nota preliminare.”” Bulletino mensile della Accademia Gioenia di Scienze naturali in Catania. Marzo 1890, (N.S.), Fascicolo XIII : 3—6. Here they give the name Laverania danilewsky [sic] n.sp. (: 4) for the crescentic parasite of sparrows; they also found an amoeboid parasite, which they left un-named, but suggested it was a Haemamoeba. (b) ID. ‘* Ancora sui parassiti malarici degli uccelli. Seconda nota preliminare.”” Aprile 1890, ibid. Fasc. XIV : 2—7. _ In this paper the amoeboid parasite of sparrows, mentioned in (a) is named Haemamoeba praecox (: 2). Lye 110 OPINIONS AND DECLARATIONS (c) IID. ‘* Nuova contribuzione allo studio della malaria. Nota — preliminare.” Jbid. Dicembre 1890 e Gennaio 1891, Fasc. XVI : 16—20. The year at the foot of the title page is 1891. In this paper they record the findings in sparrows of a new form of — Haemamoeba, which they name Haemamoeba relicta n.sp. (: 19). It — is evident from the text that they regard this as a distinct species, not merely a new name for H. praecox. This is also obvious from the next paper (paper (d) below). (d) IID. ‘‘Inoculazione dei parassiti malarici da uccello ad uccello.—Parassiti dei globuli rossi delle rane.” JIbid., Marzo, Aprile 1891, Fasc. XVII—XIX : 6—12. Here it is stated that the authors recognise in sparrows three species of malaria parasites: Haemamoeba praecox, H. relicta and Laverania danilewsky [sic]. This confirms the view of the authors as to the independence of the two Haemamoeba. Subsequently, some authors (but not Grassi & Feletti: at least not up to 1891 inclusive) regarded Plasmodium relictum as a synonym of P. praecox, while others retained it as a distinct species. I think it is clear from the above excerpts that Grassi & Feletti did not rename H. praecox but recognised two species, namely: H. praecox - and H. relicta (in addition to Laverania danilewsky). 100. From the information quoted above, we see at once that — there is no substance in the claim that the name Haemamoeba — relicta Grassi & Feletti, 1891, was published as a substitute name (nom. nov.) for the name Haemamoeba praecox Grassi & Feletti, 1890, as applied by those authors to the bird parasite which they then so named. It is perfectly clear indeed that the foregoing authors regarded the name praecox as applying to one bird parasite and the name relicta as applying to another bird parasite. Thus, the main part of the argument used by the French authors who sought to justify the use of the name praecox Grassi & Feletti for the Malignant Tertian Malaria Parasite is seen to be without any foundation. We have already seen (paragraph 97 above) that the other part of the French workers’ argument, namely that the nominal species Haemamoeba praecox Grassi & Feletti, 1890, contained among its syntypes examples not only of the bird — OPINION 283 111 parasite then so named but also of the Human Malignant Tertian Malaria Parasite is also completely devoid of foundation. Accord- ingly, under the Rég/es the specific name praecox Grassi & Feletti adheres exclusively to the bird parasite and has, moreover, done so from the moment at which it was first published. 101. The contention that the trivial name praecox Grassi & Feletti applied to the Malignant Tertian Malaria Parasite and was the oldest available such name for that species and therefore its valid name never won any general acceptance, and, in particular, was vigorously rejected by Schaudinn (1902, Arb. kaiserl, Ges. Berlin, 19 : 169), by Blanchard (1905, Les Moustiques, Paris) and by Stiles (1929, in Sergent Ed.) et. al. in Arch. Inst. Pasteur Alger. 7: 232). These authors all took the view that the text of the original description of Haemamoeba praecox Grassi & Feletti showed beyond question that this name was definitely bestowed by its authors upon the bird parasite there described and that it must therefore adhere only to that species. In the most recent of the papers cited above (Stiles, etc. 1929) Sergent (Ed.) and others re-examined this question and, while formally adhering to the traditional French view that under the Rég/es the trivial name praecox Grassi & Feletti was the correct specific trivial name for the Malignant Tertian Malaria Parasite, agreed that in existing conditions it was neither practicable nor desirable to apply the name praecox to that species. We may, therefore, now leave this subject, contenting ourselves with noting that, for the reasons explained above, the specific trivial name praecox Grassi & Feletti, 1890, has no place in any list of the names bestowed upon the human malaria parasites. At most, this name can be included in such a list only as a name which has been erroneously applied to a human malaria parasite (the Malignant Tertian Malaria Parasite) through a misreading by subsequent authors of the description published by Grassi & Feletti. 112 OPINIONS AND DECLARATIONS (7) The specific trivial name “ falciforme ” as published in com- bination with the generic name ‘‘ Ematozoo ” by Antolisei and Angelini in 1890 for the Malignant Tertian Malaria Parasite 102. When the bibliographical reference to Maematozoon — falciparum Welch, 1897 (in Loomis’s Syst. pract. Med. 1 : 36) was — being checked in the course of the preparation of the present paper, the following footnote given by Welch immediately after his description of the above species suggested the existance of a hitherto ignored specific name for the Malignant Tertian Malaria © Parasite, namely Haematozoon falciforme Antolisei & Angelini:— — The name Haematozoon falciforme suggested by Antolisei & — Angelini is objectionable, as it implies that the shape is always falciform, and is applicable only to the crescentic forms. The ~ adjective “‘ falciparum ” ( falx, “‘ sickle,” parire,“‘ to bring forth,” ~ “to produce ’’), on the other hand, indicates that the property of forming crescents is a distinctive character of the organism, and © it is therefore applicable to the variety of the parasite which ~ possesses exclusively this property. ; 103. Welch did not give a reference to the place where Antolisei — & Angelini published the name Haematozoon falciforme but he — did state that a full bibliography of works on malaria was given | by Thayer (W. S.) and Hewetson (J). in their ‘‘ Malarial Fevers of © Baltimore ” published in 1895. 104. Thayer & Hewetson’s paper, which occupies the whole — of one volume of the “Johns Hopkins Hospital Reports” — (Thayer & Hewetson, 1895, Johns Hopkins Hosp. Rep. 5: 1\—218, — 2 pls.) contains a detailed survey of the previous literature of ~ malariology. This survey includes copious references to the annexed bibliograpy (: 189—208) which includes 359 titles. In ~ their survey, Thayer & Hewetson described at length (: 25—27) © the paper ‘“‘ Sulle febbri malariche predominanti nell’estate e © autumno a Roma ” (1890, Atti. Accad. Med. Roma 16) in which — } ; . ‘ ] OPINION 283 113 Marchiafava & Celli had recorded their observations on “‘ ‘aestivo- ~ autumnal’ Roman fever.”’ After noting (: 27) that these observa- tions had been confirmed in their main features by a large number of observers, among the first of whom being Gualdi & Antolisei (who demonstrated the existence of this type of organism by inoculation experiments), Thayer & Hewetson proceeded as follows :— Antolisei & Angelini (119, 131) refer to this variety as the *‘ haematozoon falciforme.”’ 105. The following are the entries given by Thayer & Hewetson in respect of the papers by Antolisei & Angelini to which they had referred under the numbers “ 119 ” and * 131 ”:— (119). ANTOLISEI & ANGELINI. Osservacioni sopra alcuni casi d’infezione malarica con forme semilunari. Arch. Ital. di clinica medica, 1890, 1. (131). ANTOLISEI & ANGELINI. Nota sul ciclo biologico dell’ ematozoa falciforme. Riforma medica, 1890, nos. 54, 55, 56, March 6, 7, 8, pp. 320; 326; 332. 106. it is clear from the way in which Thayer & Hewetson cited these references that it was upon one or other or both of these papers that they relied for their statement that Antolise1 & Angelini had given the specific name (binominal combination) Haematozoon falciforme to the aestivo-autumnal Roman fever. It was necessary therefore to examine each of these papers care- fully. The results are given in the immediately following paragraphs. 107. The first of these papers (i.e. that numbered (119) by Thayer & Hewetson), which was published on some date in 1890, in the Archivio italiano de clinica Medica, Milan, Naples, Turin 29 : 1—23, has been kindly examined for me by Sir Rickard 114 OPINIONS AND DECLARATIONS Christophers who has been good enough to furnish the following report on it (in /itt., dated 26th April 1944):— It does not apparently give the first naming but refers several times to “‘ l’ematozoo falciforme’”’ in each case in italics. I give the passages below. Whether this naming is above all suspicion as to its being a correct zoological designation is, I think, just doubtful. Even supposing the genus name were accepted as Haematozoon it seems doubtful whether the name ematozoo falciforme is quite the zoologist’s idea of a correct binominal designation. However one cannot say what validity there may be until one has seen the passage giving the name. That others have accepted “ falciforme’”’ as a specific name definitely given meantime carries considerable weight. The passages. giving the name “ l’ematozoo falciforme”’ are:— :2 Avertiamo che tanto i casi, che pubblichiama quanto le osservazioni, che seguono, provengono soltanto da studii fatti dacche, abbiamo acquistato la convinzione, che lematozoo falciforme sia un parassita a se, nettamente diverso dagli altri, e dacche abbiamo imparato a distinguere le diverse fasi della sua vita da quelle degli altri. :8 Dalla lettura dei casi riferiti ognuno si sara_persuaso quanto sia vero, che non basta l’esame del sangue di un solo malarico per conoscere la biologia dell ’ematozoo falciforme. :9 Dicevamo che dall’esame di alcuni dei nostri casi si deume che nell’inizio delle infezioni malariche provocati dall’ematozoo falciforme, nel sangue circolante non si mostrano che le amebe. Of course all this applies to the crescentic gametocytes of falciparum which puzzled the authors because they could get no life cycle such as they now expected in any parasite on the analogy of the quartan and tertian parasites. They were not merely out, so to speak, to describe some definite zoological form. 108. There is no copy in the United Kingdom of the second of the two papers by Antolisei & Angelini cited by Thayer & Hewet- son (i.e. the paper numbered (131) in their bibliography). I accordingly applied to Dr. Robert L. Usinger in the hope that it might be possible to trace a copy of this paper in some library in the United States. Dr. Usinger very kindly consulted Dr. G. Robert Coatney, who (on 13th September 1945) was so kind as to furnish the following very interesting report :— I find that Antolisei, E. & Angelini 1889, ‘“‘ Due altri casi di febbre malarica spermentale,”’ La Riforma medica 2 : 1352—1353, —_—~—-— = OPINION 283 115 1358—1359, do, on page 1359, name falciforme but only as a type of fever, like quartan fever, and, therefore, the name is not a zoological synonym of falciparum. In 1890 the same authors in ** Nota sul cicio biologico dell’ematozoo falciforme,’” La Riforma medica 6 : 320-—321, 326—327, 332—334, on the last page (: 334) name falciforme n.sp., preceding the name falciforme with ematozoo in italics. This does not, in my opinion, constitute a generic name but rather a Japsus calami. 109. Leaving aside for the moment the question whether a specific term consisting of the words Ematozoo falciforme (i.e. the name published by Antolisei & Angelini, but with the first letter of the generic name written with a capital letter) constitutes a name which complies with the requirements of the Rég/es and is therefore, other things being equal, an available name, we have to note:— (1) that in each of the two papers cited by Thayer & Hewetson, Antolisei & Angelini did publish a new specific trivial name falciforme but that in each of these papers the generic name was spelt “‘ ematozoo ”’ and not “ haemato- zoon’”’ as stated by Thayer & Hewetson; and, therefore, that it must be assumed that the latter spelling is a deliberate emendation by Thayer & Hewetson of the original spelling “‘ ematozoo”’ ; (2) that, as no particulars are available to show the date in 1890 on which was published the Part of volume 29 of the Arch. ital. clin. Med., containing Antolisei & Angelini’s paper “‘ Osservazioni sopra alcuni casi d’infezione malarica con forme semilunari,”’ whereas the Parts of volume 6 of the Riforma medica containing those authors’ paper “ Nota sul cicio biologico dell’ematozoa falciforme ’’ are known to have been published on the 6th, 7th and 8th March 1890, it must be concluded (at least until further information is available) that the last- named paper was the first to be published and therefore that it was in this paper that the name ematozoo falci- forme was first published (Antolisei & Angelini, 8th March 1890, Riforma medica 6(56) : 334) as a name for the malaria parasite of semilunar form. (The question 116 OPINIONS AND DECLARATIONS as to which of these papers was the first to be published is of theoretical importance only, for in both papers the specific trivial name falciforme was published by Antolisei & Angelini in the same genus (ematozoo) and applied to the same species of parasite.) 110. We must turn now to consider whether the term “ematozoo falciforme”’ as published by Antolisei & Angelini constitutes a name which can be regarded as complying with the requirements of the Régles. Two problems are involved: (i) Does this name comply with the requirements of Article 3, which relates to the form of scientific names for animals ? (ii) Apart from any answer which may be given to the foregoing question, is the name pub- lished by Antolisei & Angelini invalidated either wholly as a specific name (binominal combination) or as regards the first of its component elements (the generic name) by reason of the fact that, when this specific name was published, the generic name was printed with a small, instead of a capital, initial letter, i.e. as ematozoo and not as Ematozoo? For the first of these questions, the relevant Article in the Régles is Article 3. For the second of these questions the relevant Article is Article 8 (form of generic names). In the present context this latter Article should be read in conjunction with the parallel Article (Article 13), which relates to the form of specific trivial names. 111. The following is the substantive French text of Article 3 of the Régles:— 3. Les noms scientifiques des animaux sont des mots latins ou latinisés, ou considerés et traités comme tels, au cas ou ils ne seraient pas d’origine classique. 112. The generic name Ematozoo is certainly not a “ mot latin” nor can it be claimed that it is a “‘ mot latinisé ’’ except in so far as the fact that it was printed in italics suggests that in the opinion of Antolisei & Angelini, it was a Latinised word and therefore one which, on being printed, should be distinguished in this way from the Italian words which formed the remainder of the sentence in which it occurred. The word Ematozoo is not a “ mot d@origine classique.’ Article 3 provides that a word not of classical origin ~ ae So.) oe j > 3 OPINION 283 117 ‘is to be accepted if it was considered and treated as a Latin or -Latinised word by its author. The fact that Antolisei & Angelini gave to the parasite in question a binominal name and printed that name in italics to distinguish it from the adjoining Italian . words does, in my view, create a presumption that they were con- sidering and treating this non-classical word Ematozoo as a Latin or Latinised word. This presumption is greatly strengthened by the fact that the second part of the specific name introduced by Antolisei & Angelini, namely the specific trivial name falciforme is undoubtedly a Latin word. I conclude therefore that the 1orm in which Antolisei & Angelini published the name Ematozoo falciforme satisfies the requirements of Article 3 of the Régles. 113. On turning to the second of the questions raised in paragraph 110 above, we have to consider the.substantive French texts of Articles 8 and 13. These read as follows :— 8. Le nom générique consiste en un mot unique, simple ou composé, écrit par un premiere lettre capitale et employé comme substantif au nominatif singulier. 13. Les noms de personnes employés comme noms spécifiques peuvent étre écrits par une premiere lettre capitale; tous les autres noms spécifiques s’écrivent par une petite lettre initiale. 114. It is perfectly clear that the form in which Antolisei & Angelini published the specific name here under consideration (ematozoo falciforme) does not comply with all the provisions of Article 8 (since the generic name was not written with a capital initial letter) but that it does comply with those of Article 13 (since the specific trivial name was written with a small initial letter, as is required by that Article in all cases except where the word chosen for a specific trivial name is the name of a person, in which case the author is left to choose between the use of a small initial letter and the use of a capital initial letter).1° 15 At its Session held in Paris in 1948 the International Commission on Zoological Nomenclature obtained the approval of the Thirteenth International Congress of Zoology for the redrafting of Article 13 so as to require that every specific name shall be written with a small initial letter (1950, Bull. zool. Nomencl. 4 : 173). 118 OPINIONS AND DECLARATIONS 115. The question now arises as to whether non-compliance — with Article 8 in the original publication of a new generic name | should be regarded as invalidating that name. No authoritative — ruling has been given by the International Commission on | Zoological Nomenclature on this matter, but practical considera- — tions militate strongly against any such interpretation. If Article 8 were to be read in this sense, Article 13 would similarly have to be regarded as invalidating any specific trivial name (other than that consisting of the name of a person) originally published with a capital initial letter. Now every systematist knows that count- — less thousands of specific trivial names were originally published ~ in this way, particularly a very large number already in common use at the time when the present Rég/es were adopted in 1901. It is inconceivable that those responsible for the introduction of these Régles contemplated the inclusion of a provision which would have had the effect of immediately invalidating a large proportion of the specific trivial names in common use at that time. My conclusion therefore is that Articles 8 and 13 should be read in another sense, namely: as indicating the correct forms in ~ which generic and specific trivial names should be published, ~ implying that any departure from those forms should call for automatic correction but should not be regarded as rendering the names invalid. 116. My interpretation of Article 8 is therefore that where a generic name is first published with a small initial letter, that form of printing the generic name, although constituting a contraven- tion of Article 8, does not invalidate the generic name in question for the purposes of Article 25 (Law of Priority)'®. At the same time, however, where a generic name is first published in this defective manner, that defect does call for an automatic correction of the name by the substitution by later authors of a capital initial letter for the small initial letter (erroneously) used in the original publication. 16 In 1948 at its meeting held in Paris the Thirteenth International Congress of — Zoology inserted in the Régles a provision that infringements of Article 14 (and of © certain other Articles) should be subject to automatic correction (1950, Bull. zool. — Nomencl., 4 : 67—68) and this provision was expressly extended to similar infringe- — ments of Article 8 by the Fourteenth International Congress of Zoology, Copenhagen — 1953 (1953, Copenhagen Decisions zool. Nomencl., :47—48). In the light of this clarification of the Rég/es, the assumption here made as to the status of the generic name ematozoo, as published with a small, instead of with a capital, initial letter is seen to have been correct. —— _-s OPINION 283 119 117. Applying these conclusions to the case of the generic name published by Antolisei & Angelini in 1890 for the Malignant Tertian Malaria Parasites, we may conclude that the fact that that name was originally printed with a small initial letter (as ematozoo) instead of with a capital initial letter (Ematozoo) constituted a defective method of publication, but that the defect involved does not invalidate this generic name, though it does call automatically for the substitution by later writers of a capital initial letter for the small initial letter (incorrectly) used in the original publication, the name thus becoming Ematozoo in place of ematozoo. 118. In the light of the foregoing considerations I conclude that the specific name Ematozoo (correction of ematozoo) falci- forme Antolisei & Angelini satisfies the requirements of the Rég/es and accordingly possesses rights under the Law of Priority (Article 25) as from 8th March 1890, the date of publication of the portion of Antolisei & Angelini’s paper in which this name was first published. 119. The species described by Antolisei & Angelini in 1890 as Ematozoo falciforme is the Malignant Tertian Malaria Parasite (paragraph 104), for which, as shown in paragraph 45 above, the oldest available specific trivial name, and accordingly the valid name, is malariae Laveran, 1881, of which therefore the specific trivial name falciforme Antolisei & Angelini, 1890, as published in the combination Ematozoo (correction of ematozoo) falciforme, is a junior subjective synonym. (8) The specific trivial name “ immaculata” as published in combination with the generic name ‘‘ Haemamoeba” by Grassi in 1891, the name of a parasite of birds which has been erroneously applied to the Malignant Tertian Malaria Parasite 120. In 1891 a paper entitled “‘ Weiteres zur Malariafrage ” by Grassi & Feletti was published in the Centralblatt fiir Bakteriologie und Parasitenkunde (Grassi & Feletti, 1891, Zb/. Bakt. 10: 449454, 481—488, 517—521). This paper was divided into three Sections, of which the first (: 449—454, 481—488) and the 120 OPINIONS AND DECLARATIONS third (: 519—521) were written jointly by Grassi & Feletti. The © second Section (: 517—519), however, was written by Grassi — alone. In this Section, to which was applied the sub-title © ‘“* Verschiedene Untersuchungen,” Grassi first published (: 517) — the specific trivial name immaculata (in the binominal combination Haemamoeba immaculata). 121. Grassi began by referring to a paper in which Celli & © Guarnieri had recorded the occurrence in Man of parasites which ~ segmented at a very early stage, that is to say, even before pig- mentation had occurred; those authors had associated this — parasite with another parasite which also segmented at an early stage, though not before pigmentation had occurred. Grassi had suspected that this second parasite might be a separate species, distinct from Haemamoeba praecox Grassi & Feletti, 1890, but at that time he had not found this form in any bird which he had examined. Quite recently, however, he had found a parasite — which he identified with this form in an immature specimen of — the Kestrel (Cherchneis tinnunculus). The material so obtained — was then described by Grassi and given the name Haemamoeba — immaculata. : in the immediately preceding paragraph :— A. Schon in einer vorhergehenden Note findet man angedeutet, dass Celle und Guarnieri eine Parasitenform im Menschen ~ beschrieben haben, welche sich sehr friihzeitig, bevor auch nut die i geringste Spur von Pigment existirt, segmentirt. Wahrend nun © obige Verfasser diese Form mit einer anderen, welche sich eben- falls friihzeitig, aber erst nachdem sie bereits Pigment erworben, — zusammenwerfen, stieg in mir der Zweifel auf, ob sie nicht eine Art fiir sich, verschieden von der Haemamoeba praecox, reprasen- tiren kénne. Ich hatte seiner Zeit bereits bemerkt, dass in den ~ von mir bis dahin untersuchten Végeln obige Form gewiss fehlte. Kiirzlich nun ereignete es sich, dass ich eine gleiche Form in einem, ~ sich noch im Dunenkleide befindlichen Thurmfalken (Cherchneis | tinnunculus) vorfand. Die anderen drei kleinen Thurmfalken welche sich mit ihm zusammen im Nest befanden, waren nicht von dieser Parasitenform infizirt, wahrend der erste es derartig war, dass man nur mit grosser Mihe ein freies rothes Blutk6r- ~ perchen vorfinden konnte. Diese Falke lebte acht Tage und bot |} mir stets den gleichen Befund. OPINION 283 121 Stets, in jedem Augenblick, fand sich in den rothen Blutk6r- perchen des Blutes eine Haemamdbe, welche, wie die anderen Haemamdoben der Végel, durchaus keine améboidische Bewegun- gen zeigte. Fast in jedem Praparat fand man zu jeder Stunde Segmentationsfiguren. Die Segmentation fand statt, wenn ein grosser Theil des Blutk6rperchens noch unversehrt war und ohne einen Nucleus de reliquat zu hinterlassen. Auch diese Haemamébe hat einen Kern, Keine kontraktilen Vakuolen etc. Ich schlage vor, derselben den Namen Haemamoeba immaculata zu geben. 123. In view of the unfortunate subsequent history of this name, it is particularly necessary to establish beyond possibility of question the identity of the species to which it applies. It is for this reason that I have quoted in full the original description given by Grassi for his Haemamoeba immaculata. We must note, therefore, that the material to which Grassi applied the new name Haemamoeba immaculata consisted exclusively of material obtained from the blood of an immature Kestrel. The parasite to which this name was given was thus an avian parasite. It is true, however, that in the passage immediately preceding that in which he described this avian parasite Grassi subjectively identified the _ parasite which he had found in the blood of the Kestrel with a parasite with which he was not familiar in nature but which had been described in a paper published by Celli & Guarnieri as having been found in the blood of Man. The specific name Haemamoeba immaculata, being a name based solely upon material obtained from the Kestrel, applies objectively only to that material, for it was that material alone which was before Grassi when he drew up his description. Accordingly, the identification by Grassi of the bird parasite which he had before him (and to which he gave the new name Haemamoeba immaculata) with a published description of a parasite found in Man repre- sents no more than a subjective taxonomic judgement which, whether correct or incorrect, can have no possible bearing on the status of the name Haemamoeba immaculata, which objectively was attached for all time to the bird parasite from which it was described and to no other. 122 OPINIONS AND DECLARATIONS 124. On page 518 of his paper (the page following that on which the name Haemamoeba immaculata was first published and on which the species so named was described) Grassi turned to review the existing state of knowledge regarding the human malaria parasites. He summarised his views on this subject in a table in which he recognised five such parasites (four in the genus Haemamoeba and one in the genus Laverania). The fourth of the ‘“‘ species’ recognised by Grassi as belonging to the genus Haemamoeba figured in his list as ‘““Haemamoeba immaculata.” The human malaria parasite so referred to (together with the parasite which immediately preceded it and to which Grassi applied the name Haemamoeba praecox Grassi & Feletti, 1890, erroneously as has been shown in paragraph 101 above) was characterised by Grassi as follows: ‘“‘ verursacht pernicidse, quotidiane, subkontinuirliche und kontinuirliche Fieber.” A little lower down the same page, Grassi, clearly quoting from the observations published by Celli & Guarnieri, added the note: ‘** Die Haemamoeba immaculata wurde im Menschen bis jetzt nur in Rom angetroffen.’’ Thus in his survey of the human malaria parasites Grassi repeated and amplified his earlier subjective taxonomic judgment that the bird parasite Haemamoeba immacu- lata occurred also in the blood of Man and was responsible for ~ one of the kinds of human malaria fever. For the reasons already explained, this subjective judgment on the part of Grassi is entirely irrelevant, so far as concerns the question of the identity of the species to which the name Haemamoeba immaculata applies, that being a question settled objectively once and for all by the fact that it was the bird material which alone was used by Grassi when he drew up his description of that species. 125. It is now generally agreed that Grassi was (taxonomically) in error when he subjectively identified the bird parasite Haema- moeba immaculata with a human malaria parasite. Malariologists are generally agreed also that the human malaria parasite which Grassi misidentified with his bird parasite and to which he there- fore erroneously applied the specific name Haemamoeba immacu- lata was the Malignant Tertian Malaria Parasite. OPINION 283 123 126. So far as human malariology is concerned, the subsequent history of the specific trivial name immaculata Grassi, 1891, is very similar to that of the specific trivial name praecox Grassi & Feletti, 1890 (discussed in paragraphs 98-99 above). Both names have been used for the Malignant Tertian Malaria Parasite and both for the same reason. In each case the widespread acceptance of the erroneous determination by Liihe (1900) (discussed in paragraphs 46-51 above) of Oscillaria malariae Laveran, 1881, as the Quartan Malaria Parasite left (as it was believed) the Malig- nant Tertian Malaria Parasite without an available specific trivial name. The difference between the later development of these two cases lies in the fact that, while workers in France accepted (erroneously as is now seen) the specific trivial name praecox Grassi & Feletti, 1890, as the next available specific trivial name for the Malignant Tertian Malaria Parasite (paragraph 98 above), German workers such as Schaudinn (1902, Arb. Gesundh. Amt., Berlin 19 : 169) rejected the name praecox (quite correctly) on the ground that it had been originally applied to a bird parasite and not to a human malaria parasite. These and later German authors accordingly adopted the specific trivial name immaculata Grassi, 1891, for the Malignant Tertian Malaria Parasite, over- looking the fact that the use of this name in this sense was open to precisely the same objection as that advanced against the use of the name praecox Grassi & Feletti. 127. The position is therefore that from a nomenclatorial point of view the specific trivial name immaculata Grassi, 1891, as published in the combination Haemamoeba immaculata, is a name published for a parasite of birds and not for a malaria parasite of Man and that it was only through the erroneous subjective identification of the bird parasite with a human malaria parasite that this trivial name ever found its way into the literature of human malariology. The specific trivial name immaculata Grassi, 1891, has, therefore no place in the synonymy of the names bestowed upon human malaria parasites, though (like praecox Grassi & Feletti, 1890), it must be included in any list of names at one time or another erroneously believed to have been so bestowed which may be drawn up in connection with such a synonymy. 124 OPINIONS AND DECLARATIONS (9—11) The terms “ quartanae,” “ tertianae ” and “ quotidianae ” _ erroneously alleged to have been published by Celli and Sanfelice in 1891 as trivial names for the Quartan Malaria | Parasite, the Benign Tertian Malaria Parasite and the © Malignant Tertian Malaria Parasite respectively, treated by | the above authors as varieties (subspecies) of the nominal © species ‘Plasmodium malariae” Marchiafava and Celli, — 1885 | 128. In 1891 Celli & Sanfelice published a long paper entitled “Sui Parasiti del Globulo Rosso nell’?Uomo e negli Animali ” (1891, Ann. Ist. Igiene sper. Univ. Roma (n.s.) 1 : 33—63, 3 pls.). This paper is divided into a number of Sections of which the third is entitled ““ Uomo malarico.”’ In this Section (: 54) human malaria fever was stated to consist of three forms, This passage reads as follows :— . .. . Secondo poi il tipo del febbricitare si pud svolgere sotto tre forme :— (a) quartana: ne sono combinazioni la quartana doppia, la — quartana triplice 0 pseudoquotidiana; (b) terzana: n’é una combinazione la terzana doppia o pseudo- — quotidiana; (c) quotidiana: combinazioni ne sono le febbri ad accessi prolungati, le subentranti, le subcontinue, le perniciose. 129. Throughout the remainder of Section III of their paper Celli & Sanfelici discussed human malaria fever (not the human malaria parasites) under these names. Quartan Malaria was discussed under the title ‘‘ Nella quartana”’ (: 54—55), Benign Tertian Malaria under the title ‘‘ Nella terzana” (: 55) and Malignant Tertian Malaria under the title ‘‘ Nella quotidiana ”’. (: 55—58). In a concluding Section (which is unnumbered and lacks a title), commencing on page 59 and ending on page 61, Celli & Sanfelice brought together the conclusions reached in the OPINION 283 125 _ earlier part of their paper and on page 61 they set out as shown in the following table (reproduced in facsimile in Plate I) the views _ which they had formed in regard to the parasites responsible for | the types of fever discussed in their paper :— Per distinguere le specie adottiammo anche noi il criterio della _ sede, al quale va talora congiunto quello morfologico e sempre quello sperimentale, per cui finora da una specio all’altra un parasita non fu inoculabile. Alcune specie hanno della varieta, che distinguiamo secondo la rapidita di sviluppo, cioé: A) lento; B) accelerato; C) rapido. 0 aS ae a oo Tenendo conto per ora soltanto delle specie meglio studiate, la loro classificazione sarebbe dunque come segue: a es ee ee Generi Species Varieta . ranarum R. Lank . testudinis Dan. . lacertae Dan. columbae Gras. . Danilewsky Kru. aluci n.sp. . bubonis n.sp. 1° Haemogregarina (Danilewsky) Soe TT eT ; H. passeris Gras. A 2° Haemoproteus & (Kruse) | | H. noctuae n.sp. A C A H. alaudae n.sp. B C A—quartanae 3° Plasmodium Pl. malariae M.¢C. B—tertianae (Marchiafava e C—quotidianae Celli) 130. The serial publication in which the foregoing paper by Celli & Sanfelice was published is very scarce and it is doubtless for this reason that the nomenclature used in it for the human malaria parasites has not before now been critically examined. Even Christophers had not in 1945 (Rev. Inst. Salubridad y Enfermedades trop. 6(4) : 220—221) seen this paper and, misled by earlier copyists, fell into the error of stating that Celli & 126 OPINIONS AND DECLARATIONS Sanfelice had published ‘‘ quotidianae”’ and “‘ tertianae’”’ as the ‘* names ”’ of varieties of a species to which they had applied the specific name ‘‘ Haemamoeba malariae” but that they had published ‘‘ quartanae”’ as the name of a variety of a species to which they had applied the specific name “‘ Plasmodium malariae.” Actually, as can be seen from the table reproduced in the immediately preceding paragraph, all three of these so-called ‘** names ”’ were published by Celli & Sanfelice in connection with varieties attributed by those authors to Plasmodium malariae Marchiafava & Celli. 131. It will be seen from the table reproduced in paragraph 129 above that Celli & Sanfelice:— (1) treated all the then known forms of human malaria parasite as being referable to a single species which they placed in the genus Plasmodium Marchiafava & Celli, 1885, and to which they applied the specific trivial name malariae Marchiafava & Celli, 1885; (2) distinguished the three forms of human malaria parasite as varieties of Plasmodium malariae Marchiafava & Celli, 1885, under the letters “A,” “ BL” gaa exactly the same way as, for the three preceding species, each placed in the genus Haemoproteus Kruse, 1890, they had recognised two forms (designated as varieties “A” and “B”) of H. passeris Gras., two forms _ (similarly designated) of the new species H. noctuae Celli & Sanfelice, and three forms (designated as varieties “A,” “B” and “C’’) of the new species H. alaudae Celli & Sanfelice; (3) added, in the case of Plasmodium malariae Marchiafava & Celli, the word “ quartanae ” against “‘ Varieta A’, the word “tertianae’”’ against “ Varieta B”’ and the word ‘* quotidianae ’’ against “‘ Varieta C.”’ OPINION 283 12d 132. When we come to interpret the new terms introduced by Celli & Sanfelice in the paper under discussion, the first point which we have especially to observe is that it is only in the table (reproduced in paragraph 129 above) on page 61 of their paper that they used the Latin words “ quartanae,” “ tertianae’’ and ** quotidianae ” respectively in connection with “ the”? human malaria parasite. It is only by reference back to the Italian words used for the types of fever concerned, namely “‘ quartana,”’ *terzana’”’ and “ quotidiana,” used on pages 54 and 55 of Celli & Sanfelice’s paper that it is possible to identify ‘‘ Varieta A— quartanae ’’ with the Quartan Malaria Parasite, “‘ Varieta B— tertianae’’ with the Benign Tertian Malaria Parasite, and “ Varieta C—dquotidianae ” with the Malignant Tertian Malaria Parasite. Article 25 of the Rég/es requires that a name, to acquire rights under the Law of Priority, must be accompanied by an indication, definition or description (“‘ accompagné d’une indica- tion, d’une définition ou d’une déscription ’’): The word selected as the trivial name for a taxonomic unit, even if (as in the present case) it conveys a clue to the identity of the organism so named, cannot be regarded as constituting an “ indication,” unless it is accompanied by at least a few descriptive words. In the present case the three alleged names (quartanae, tertianae and quotidianae) were not so accompanied in the only place where they appear in Celli & Sanfelice’s paper. We must conclude therefore that, even if those authors intended to use these words as scientific zoological names (a question which is discussed in paragraphs 134 to 137 below) they published them without even the minimum “indication ’’ required by proviso (a) to Article 25 and therefore that these names (if they were names) possess no rights under the Law of Priority as from the date of the publication of Celli & Sanfelice’s paper. 133. We must note here, in passing, that, if Celli & Sanfelice had intended that the terms “ quartanae,” “ tertianae”’ and ** quotidianae ” should be regarded as scientific zoological names and if each of those names had been duly accompanied by an indication, one of them would nevertheless automatically have been invalid. For under the Rég/es the nominotypical subgenus of a genus automatically bears the same name as the genus itself (Article 9), and by universal practice the same principle is observed 128 OPINIONS AND DECLARATIONS by zoologists in relation to the name to be used for the nomino- typical subspecies of a species. It would be well however if the International Congress of Zoology were to take an early oppor- tunity to insert in the portion of the Rég/es relating to the names of species and subspecies an express provision similar to that already prescribed by Article 9 in relation to the names of nominotypical subgenera, even though such a provision would do no more than give official sanction to the present practice of zoologists!”. In the present case Celli & Sanfelice stated that they recognised only one species of human malaria parasite (Plasmodium malariae Marchiafava & Celli), which, in their view, comprised three varieties (““ A,” ““B” and “C’’). If, as we have assumed above for the purpose of the present argument, a name had been given by Celli & Sanfelice to each of these varieties, one of them would have been given to the nominotypical subspecies of Plasmodium malariae Marchiafava & Celli, 1885. Celli & Sanfelice themselves both clearly and correctly indicated (: 55) that, in their view, the name Plasmodium malariae, as published by Marchiafava & Celli in 1885, applied to the Malignant Tertian Malaria Parasite. The passage in which Celli & Sanfelice gave this indication reads as follows: ‘“‘ Nella quotidiana vera (fig. 20—38) la varieta para- sitaria (Marchiafava & Celli)...” . It is clear, therefore, that Celli & Sanfelice considered that their “‘ Varieta C—quotidianae ” (as so cited in the table given on page 61 of their paper) was the parasite responsible for the fever which (: 55) they had discussed under the Italian term “ quotidiana” and which, being the Malignant Tertian Malaria Parasite, was the species to which Marchiafava & Celli had given the name Plasmodium malariae. Thus, if Celli & Sanfelice had intended that the Latin adjective 17 At Paris in 1948 the International Commission on Zoological Nomenclature remedied the deficiency in the Régles here referred to by securing the approval of the Thirteenth International Congress of Zoology to the insertion in the Rég/es of an Article providing that, where a nominal species has two or more nominal sub- species, the subspecific name for the subspecies on which the original description of the species itself was based shall have as its subspecific name the same name as that borne by the collective species itself, thereby giving valid force to the existing practice of systematists by introducing a provision at the specific and subspecific name level exactly parallel to the provision which had always existed in Article 9 of the Régles at the generic and subgeneric name level (1950, Bull. zool. Nomencl. 4:191). At the same time the Paris Congress decided that the subspecies on which a nominal species is based should be known as the “‘ nominotypical ”’ sub- species of the species concerned. At Copenhagen in 1953 the Fourteenth Inter- national Congress of Zoology substituted the expression “ nominate” for the expression “‘ nominotypical ”’ (1953, Copenhagen Decisions zool. Nomencl. : 21). OPINION 283 129 quotidianae applied by them to their “ Varieta C” should be regarded as a scientific zoological name for a “ variety’ of the nominal species Plasmodium malariae Marchiafava & Celli, 1885, the name quotidianae Celli & Sanfelice, 1891, would have been a subjective junior synonym of the name malariae Marchiafava & Celli, 1885, since, as is agreed, the former as well as the latter name applies to the Malignant Tertian Malaria Parasite. Since, as has been explained above, the giving of a separate subspecific trivial name to the nominotypical subspecies of a polytypic species contravenes the Régles, the name quotidianae Celli & Sanfelice;, 1891, would have been invalid, even if (as we have seen was not the case) it had been duly published with an “ indication ’’ as required by proviso (a) to Article 25. 134. We have next to consider whether the Latin words *“ quartanae,” “‘ tertianae’’ and “ quotidianae,” as published by Celli & Sanfelice in the table on page 61 of their paper were intended to be scientific zoological names at all. For this purpose we have first to consider the status of these alleged names in relation to the provisions of paragraph (1) of Article 14 of the Régles, which specifies the various permissible types of specific trivial name. The position of such names, when adjectives, is dealt with in sub-paragraph (a) of paragraph (1), which provides that such names must agree in gender with the generic name. Attached to this sub-paragraph is the example “ Felis marmorata.” (The substantive French text of this portion of Article 14 reads: *“ Les noms spécifiques sont: (a) des adjectifs s’accordant en genre avec le nom générique; Exemple: Felis marmorata.’) The generic name “ Felis’ (in the example cited) is in the nominative singular as required by Article 8. Accordingly, as the generic _ name Felis is a feminine noun, the adjectival specific trivial name is in the feminine gender and in the nominative singular case, appearing in the form “‘ marmorata.” 135. Applying these principles to the present case, we find that an adjectival trivial name of a subspecies (variety) of Plasmodium malariae Marchiafava & Celli would need to be (1) in the nomina- tive singular and (2) in the neuter gender, that being the gender of the word Plasmodium (the generic name). Thus, if (to take, E 130 OPINIONS AND DECLARATIONS as an example, the first of the Latin words used by Celli & Sanfelice) it were desired to use the adjective guartanus, -a, -um, as the trivial name of a subspecies of a species of Plasmodium, the subspecific trivial name would appear in the form “ quarta- num.” When we turn to Celli & Sanfelice’s paper, what do we find? Instead of being in the form “* guartanum” (as required by the Rég/es) this word is in the form “‘ quartanae.” Thus this word was used by those authors in the genitive singular (instead of in the nominative singular) and in the feminine gender (instead of in the neuter gender). The other two Latin words used by Celli & Sanfelice, namely “ tertianae’’ and “‘ quotidianae,” were published in exactly the same form. 136. Why was it that Celli & Sanfelice placed these three Latin words in the genitive singular case and treated them as applying to some feminine noun? No explanation can be found for their action, so long as it assumed that these authors, when using these Latin adjectives, were intending to give scientific zoological names to the three “varieties”? of the single nominal species Plasmodium malariae Marchiafava & Celli, recognised by them in | the paper under consideration. The action of Celli & Sanfelice is only explicable on the assumption that they never intended these adjectives to be treated as scientific zoological names and were quite content to leave unnamed the three “ varieties ” designated by them as varieties “‘ A,’”’ ““ B”’ and “ C,”’ in the same way as they designated by letters, but did not give names to, the ‘* varieties ’’ which they recognised for the three species of the genus Haemoproteus Kruse enumerated in the same table (repro- duced in paragraph 129 above). Once we discard the erroneous notion that these Latin adjectives were intended to be scientific names, it becomes apparent that the object of Celli & Sanfelice in using these words was to denote the type of malaria for which each of the so-called “varieties” of Plasmodium malariae Marchiafava & Celli was responsible. We may therefore conclude that Celli & Sanfelice employed these words as adjectives in agreement with the noun malariae (as used for the trivial name of the species), regarding them not as names but as descriptive epithets. To take, for example, their use of the word “‘ quartanae” in connection with their “‘ Varieta A,’”’ they may ‘have used this word in agreement with the word “‘ malariae ”’ (the trivial name of a ae ———— OPINION 283 131 the species), which is a feminine noun and, in the way there used, is in the genitive singular case; the full phrase understood in that case would be ‘“‘ Plasmodium malariae quartanae”’ or, as one might say, the ‘‘ Plasmodium of (or responsible for) Quartan Malaria.” It is possible however, that the word understood but not expressed with which the adjective “‘ quartanae,” as used by Celli & Sanfelice was in grammatical agreement, was not the word “ malariae’ but the word “‘ febris,” in which case it would be even clearer that the word “ quartanae ”’ was used by these authors as a technical designation and not as a subspecific trivial name, the concept intended to be conveyed by this designation being ** The Plasmodium of the Quartan Fever.” That this is the true explanation of the use of these descriptive adjectives in the genitive case with no noun expressed is lent some colour by the fact that technical designations of this kind were certainly used by some malariologists at about this time. - In this connection, we may refer to the cheironym Haemamoeba febris irregularis attributed to Sakharov (paragraphs 151—157 below) and the use by Koch of such expressions as febris tropica (paragraphs 178—185 below). 137. While, therefore, there may be some doubt as to the precise manner in which Celli & Sanfelice intended that their use of the terms “ quartanae,” “tertianae’’ and “ quotidianae ” should be interpreted, it seems clear that those authors did not look upon these terms as names in the sense understood in zoological nomenclature. 138. To sum up, if the terms “‘ quartanae,” “ tertianae’”’ and *“ quotidianae”’ had been published by Celli & Sanfelice as scientific zoological names, the names consisting of these adjectives would have been invalid, because they were published without an indication, definition or description (see paragraph 132 above). In fact, however, it was never the intention of Celli & Sanfelice that these adjectives should be regarded as scientific zoological names, for these words were employed by those authors not as subspecific trivial names for the parasites concerned but as Latin adjectives to designate types of the fever to which those parasites 132 OPINIONS AND DECLARATIONS give rise. The so-called varietal (i.e. subspecific) trivial names quartanae, tertianae and quotidianae of Celli & Sanfelice, 1891, are thus seen not to be names at all. It is likely, indeed, that these terms would never have found their way into the literature if Celli & Sanfelice’s paper had been readily available to sub- sequent students. 139. While for the reasons which I have explained there appears no reasonable doubt that the interpretation of Article 14 suggested in the present Section is correct, it is admittedly a weakness in — the Régles that they should contain no express provision relating — to the status of terms such as quartanae as published by Celli & Sanfelice in 1891, and in a later portion of the present paper (paragraphs 279—280 below) I therefore submit a recommenda- tion that words should be inserted in Article 14 clarifying its meaning in the sense suggested above?®. (12) The term ‘‘ hominis ’’ as published in combination with the - generic name ‘‘ Laverania ’’ by Danilewsky in 1891 140. In 1891 (Ann. Inst. Pasteur 5(12) : 758—782), Danilewsky published a paper entitled “‘ Contribution a l’étude de la Micro- biose malarique,’’ in which he discussed the relationship of the malaria parasites of Man and birds. Towards the close of his paper Danilewsky gave (: 780) the following schematic picture of his ideas of classification, although (it will be noted) he expressly _ stated that he reserved for another occasion the discussion of the place to be allotted in his scheme in the zoological system :— Au point de vue de l’hypothése unitaire de l’infection malarique on pourrait proposer le rapprochement suivant des diverses 18 The recommendation here referred to was approved by the International Com- mission on Zoological Nomenclature at its Session held at Paris in 1948, and on its recommendation, words clarifying the meaning of Article 14 in this matter were added to the Régles by the Thirteenth International Congress of Zoology (1950, Bull. zool. Nomencl. 4 : 624—625). OPINION 283 133 formes du parasite, sans entrer pour cela dans la discussion de sa place dans le systeme zoologique :— Cytozoon malariae Cytozoon praecox (a) Haemamoeba-Cytamoeba («) hominis s. Cytosporon (b) Cyosporon avium (B) avium Polymitus (c) (c) Haemogregarina avium Laverania (e) Laverania hominis 141. The only specific trivial name first published in the above table in connection with the human malaria parasites is the word “hominis ’ which appears in the first column as a sub-unit of the comprehensive term “* Cytozoon malariae’’ and in the third column in combination with the generic name Laverania (as Laverania hominis). 142. Later authors devoted a considerable amount of time and trouble in trying to elucidate the meaning to be attached to the new names first published in the foregoing table. These efforts were, however, completely fruitless owing to the hopeless over- lapping and confusion which characterised the nomenclature employed by Danilewsky. Accordingly, in order to put an end to discussion of this question, Dr. Ernest Hartman (School of Hygiene and Public Health, Johns Hopkins University, Baltimore, Maryland, U.S.A), invited the International Commission on Zoological Nomenclature to give a definite ruling on the status - (if any) of the names used by Danilewsky in his 1891 paper. The decision of the Commission is embodied in its Opinion 101 (published in 1928) (Smithson. misc. Coll. 73 (No. 5) : 13—15), the “summary” of which reads as follows: ‘“‘ The technical Latin designations used by Danilewsky, 1891, Annales de I’ Institut Pasteur, Vol. 5 (12), pp. 758—782, are not in harmony with the International Rules of Zoological Nomenclature and are therefore not subject to citation or the Law of Priority on basis of said publication.” 143. The so-called specific trivial name hominis as published in the combination Laverania hominis by Danilewsky in 1891 is thus seen to have no status in zoological nomenclature. 134 OPINIONS AND DECLARATIONS (13—15) The terms ‘‘ quartanae,”’ “‘ tertianae ” and “ irregularis ” erroneously alleged to have been published by Kruse in 1892 as trivial names for the Quartan Malaria Parasite, the Benign Tertian Malaria Parasite and the Malignant Tertian Malaria Parasite respectively, treated by Kruse as varieties (subspecies) of the nominal species ‘“‘ Plasmodium malariae ” 144. In 1892 (Hyg. Rdsch. 2 : 357—380, 453—485) Walther Kruse published a paper entitled ‘“‘ Der gegenw4rtige Stamd unserer Kenntnisse von den Parasitaren Protozoen,”’ in which he treated the three known forms of human malaria parasite as belonging to a single polytypic species to which he applied the name Plasmodium malariae. He gave descriptions of each of these parasites, referring to each by a term consisting of a com- bination of three Latin words, the first two being “‘ Plasmodium malariae” and the third an adjective which, on first inspection, resembles a subspecific trivial name. 145. The following is an extract from Kruse’s paper showing how he used these names if that was what they were intended to be:— [466] Wir unterscheiden jetzt: 1. Das Plasmodium malariae quartanae. WDasselbe vollendet seinen Cyclus in drei Tagen, von dem jiingsten unpigmentirten bis zu dem das Blutk6érperchen ganz ausfiillenden melaninhaltigen erwachsenen Stadium, das entweder in Form einer Margerite 8—12 Sporen entwickelt, oder ohne sol che zu bilden, degenerirt. [Here follows detailed description. ] 2. Das Plasmodium malariae tertianae, vollendet seinen Cyclus in zwei Tagen. Wenn es erwachsen ist, fiillt es gleichfalls das Blutkérperchen vd6llig aus, bildet aber dann entweder 14—20 Sporen in Form einer Rosette, oder degenerirt, sei es nach Art der entsprechenden Quartanaparasiten, sei es unter Verwandlung in einen mit Geisseln versehenen K6rper OPINION 283 135 [467] — ein Prozess der beim Haemoproteus beschriebenen genau gleicht. [Here follows detailed description.] 3. Dar Plasmodium malariae irregularis erscheint in zwei Entwickelungsreihen, die sich scharfer wie bei den andern Varietaten von einander differenziren lassen. [Here follows detailed description. ] 146. We have now to consider whether the words “* quartanae,”’ “tertianae”’ and “irregularis”’ were intended by Kruse to be varietal names in the sense in which names are given in zoological nomenclature. We have first to note that Kruse gave no indica- tion whatever that, in using these words, he was introducing new scientific names for these parasites. If that was his intention, this is a curious and unusual way in which to publish new names. Second, we have to note that, if it was Kruse’s intention to intro- duce new subspecific names, he did so in correct trinominal form but in a manner which would nevertheless have been very strange at the date when he published his paper, for at that time the universal method of introducing a new “varietal” (i.e. sub- specific) name was to insert the word “‘var.’”’ [=varietas] or some equivalent expression between the specific trivial name (in this case the name “ malariae’) and the new varietal (i.e. subspecific trivial) name. For both these reasons it must be admitted that, if Kruse intended to bestow new varietal (subspecific) trivial names on the three human malaria parasites known to him, he — did so in a very peculiar manner. 147. We have further to note that, if the words “ quartanae,”’ * tertianae ” and “ irregularis ’’ were intended to be names in the zoological sense :— (a) each of these “‘ names’? would have been incorrectly formed, for, contrary to the provisions of Article 14 (taken in conjunction with Article 8) of the Régles (of which the latter requires that a generic name shall be a noun substantive in the nominative singular and the former that a specific or subspecific trivial name, when consisting of an adjective, must agree in case and gender 136 OPINIONS AND DECLARATIONS with the generic name), the adjectives used by Kruse in the present context were placed by him in the genitive case and the feminine gender, although the generic name used (Plasmodium) was correctly cited in the nominative singular and is a neuter noun; (b) the specific name Plasmodium malariae would need to be looked upon as having been used by Kruse as a kind of collective name with no definite content, the taxonomic unit to which the specific trivial name malariae was first given (i.e. the Malignant Tertian Malaria Parasite) by Laveran in 1881 in the combination Oscillaria malariae (which is also the same species as that to which the same specific trivial name was given by Marchiafava & Celli in 1885 in the combination Plasmodium malariae), being given by Kruse a separate varietal (subspecific trivial) name, “ irregularis,’ contrary to the provisions of the Régles that, where it is necessary to refer to the nomino- typical subspecies of a polytypic species having two or more subspecies, the trivial name of the species is to be applied to the nominotypical subspecies, no new sub-_ specific trivial name being employed. 148. We are, therefore, confronted with the question: “‘ Why, if Kruse intended the words “ quartanae,” “ tertianae”’ and ‘* irregularis ’’ to be subspecific trivial names, did he not write the first of these words as “‘ quartanum,” the second as “ tertianum ” and the third as “ irregulare,”’ as he would have done if he had formed those “‘ names” in accordance with the practice which then and for many years previously had already been universally recognised as correct (and which has since been enshrined in Articles 8 and 14 of the Régles)?” The correct answer to this question is, I am convinced, that Kruse never intended these three adjectives to be regarded as subspecific trivial names in the scientific zoological sense. 149. The reasons which have led me to this conclusion are identical with those which led me to a similar conclusion, when considering the use by Celli & Sanfelice in 1891 of three adjectives OPINION 283 137 used also in the genitive singular and in the feminine gender (“‘ tertianae,” “‘ quartanae”” and “ quotidianae’’) in connection with the human malaria parasites which those authors (like Kruse a year later) regarded as belonging to a single polytypic species, to which they (again like Kruse) applied the specific name Plas- modium malariae. The interpretation which I have given of this use by Celli & Sanfelice of these three adjectives is (as will be seen from paragraphs 134—136 above) that those adjectives were deliberately placed by those authors in the genitive singular and in the feminine gender because they were using those adjectives not as names in the zoological sense but in their ordinary Latin sense as descriptive epithets applied to the specific trivial name malariae which is a noun substantive here cited in the genitive singular and is of the feminine gender. Exactly, the same answer should, in my view, be given to the questions raised by Kruse’s use of the three adjectives here under discussion. 150. If the terms “‘ quartanae,” “ tertianae ’’ and “ irregularis ”’ as published by Kruse in 1892 in combination with the specific name Plasmodium malariae had been acceptable as validly pub- lished zoological names, the first—as Kruse’s description shows— would have applied to the Quartan Malaria Parasite, the second, to the Benign Tertian Malaria Parasite, the third, to the Malignant Tertian Malaria Parasite. For the reasons explained in the immediately preceding paragraph it is clear however that the foregoing terms were not published by Kruse as scientific names. and therefore that there are, in fact, no such names as the alleged names quartanae Kruse, 1892, tertianae Kruse, 1892, and irregu- laris Kruse, 1892. (16) The cheironym ‘“‘ irregularis ’’ erroneously alleged to have been published in the combination ‘‘ Haemamoeba febris. irregularis ®’ by Sakharov in 1892 as a name for the Malignant Tertian Malaria Parasite 151. Sir Rickard Christophers has informed me (in Jitt., 14th September 1944) that he has a note that the name irregularis has been alleged to have been published by Sakharov (or Sakharoff) as a varietal (i.e. subspecific) name in 1892 in the trinominal combination Haemamoeba febris irregularis and to have been applied by Sakharov to the Malignant Tertian Malaria Parasite. 138 OPINIONS AND DECLARATIONS At the same time, Sir Rickard added: “ I cannot find any definite naming by Sakharoff in proper form and unless there is some other paper than I have so far seen in which this is done, the name may, I think, be safely omitted.” 152. In a later letter, dated 16th October 1944, Sir Rickard furnished the following supplementary report: “I have not succeeded in finding any definite zoological naming by Sakharoff of irregularis. I have looked up all the papers by him that I can find but without result. He refers to his parasite constantly as ‘ le parasite des fiévres paludéennes irreguliéres ’ and as “les parasites des fiévres irreguliéres,’’ but nowhere have I found a name in proper zoological form.” 153. Dr. C. M. Wenyon, writing on 18th December 1944, forwarded a note prepared by Miss I. M. Bellis in the light of a ~ search of the literature which she had made both for this name and also for the cheironym tropica. Miss Bellis pointed out, by way of negative evidence, that, as observed by Ziemann (1906, in Mense’s Handb. Tropenkrankh. 3 : 281), Luhe (1906, in Mense’s ~ ibid. 3. : 220) expressed the opinion that there was so much con- — fusion in the nomenclature of the human malaria parasites that it was much better to refer to those species by their vernacular names. In spite of this, Liihe then proceeded to discuss all the scientific names which to his knowledge had been applied to these species. In this survey however he made no mention of a specific trivial name irregularis. | 154. Miss Bellis drew attention also to the fact that Koch © (1898, Reise-Berichte iiber Rinderpest, Bubonenpest :95) men-_ tioned “‘ irregulare Malaria ” (and also “* tropische Malaria ’’) but did not introduce any new specific trivial name for the parasite that he was discussing (the Malignant Tertian Malaria Parasite). OPINION 283 139 155. Dr. G. Robert Coatney, in a letter dated 13th September 1945, reported as follows :— I cannot verify Haemamoeba febris irregularis Sakharov, 1892. However, Sakharoff (same man but different spelling), 1891, ** Recherches sur le parasite des fieévres paludéennes irreguliéres ”’ (Ann. Inst. Pasteur 5: 445—449) says that he thinks those malarias that exhibit irregular fevers are distinct species from those that do not but he does not name a n.sp. irregularis. 156. It seems to me possible that the explanation of this problem may be found in the (alleged) use (by Sakharov) of the term irregularis not as a specific trivial name in a binominal combination (as required for the scientific name of a species in zoological nomenclature) but as the third portion of a term consisting of a trinominal combination of Latin words, of which the second word was the word “ febris.”’ For it is possible that, if not Sakharov himself, at least some later author referring to Sakharov’s papers, may have used a trinominal expression of this kind not as a subspecific name in the sense in which such names are used in zoological nomenclature but rather as a convenient technical designation of the “Haemamoeba of the Irregular Fever,” i.e. the Malignant Tertian Malaria Parasite. The use by one author of a technical designation of this kind might readily be misinterpreted by a later author as indicating that the author by whom this designation had first been used had thereby intro- duced the name irregu/aris as a new specific trivial name for the species of Haemamoeba responsible for irregular fevers. 157. Whether or not the above is the correct explanation of the Origin in the literature of malaria of the alleged name irregularis Sakharov, the possibility that that name was ever published as a specific trivial name in conditions which satisfy Article 25 of the Régles may be regarded as being very remote. The chance that this name was duly published as a specific trivial name in proper zoological form cannot however be entirely ruled out, but in the circumstances the proper course is, I think, to treat this name as a cheironym. It is desirable however, that, when the International Commission on Zoological Nomenclature comes to deal with the problem of the human malaria parasites, it should take appropriate steps to guard against the contingency that the above name was ever validly published. 140 OPINIONS AND DECLARATIONS (17—19) The specific trivial name ‘‘ laverani’’ as published in combination with the generic name ‘*‘ Haemamoeba’”’ by Labbé in 1894 for the Malignant Tertian Malaria Parasite and the trivial names ‘‘ quartana’’ and ‘“‘ tertiana ”’ published at the same time by that author for the Quartan Malaria Parasite and the Benign Tertian Malaria Parasite respectively, then treated as subspecies of the Malignant Tertian Malaria Parasite 158. In 1894 (Arch. Zool. exper. gén. (3) 2 : 55—258, pls. 1—10) Labbé published a long paper entitled ‘‘ Recherches zoologiques et biologiques sur les Parasites endoglobulaires du Sang des Vertebrés.”” In the fourth section of this paper, which was headed ‘“‘ Les Parasites endoglobularires du Sang de l Homme ”’ (: 160—170) the author reviewed the work of Laveran and his successors. After rejecting (: 168) the conclusion of Grassi & Feletti that the crescent-formed parasites should be separated — generically from the other human malaria parasites, Labbé posed a more radical question in the following words: “ Le parasite de la malaria constitue-t-il une espece unique, polymorphe ?” After observing that this important question was one which it was difficult to decide, Labbé proceeded to consider the position of the two varieties (“‘ vari€étés *?) separated by Golgi, namely the ‘“* Parasite de la quarte”’ (i.e. the Quartan Malaria Parasite) and the “‘ Parasite de la tierce” (i.e. the Benign Tertian Malaria Parasite). Of these he observed that they seemed to be fairly well established (“‘ assez bien établies ’’) by the characters which he proceeded to summarise. He added that they were accepted by practically every author, except Laveran who applied the expression ‘‘ corps sphériques” alike to the parasite of the ‘tierce’ and to the parasite of the “‘ quarte.” A little later (: 169) after discussing the conclusions reached by Mannaberg, Labbé gave as his opinion that these two parasites constituted varieties of a single parasite (“‘ Il semble donc a peu pres établi que les parasites tierce et quarte constituent des variétés d’un méme parasite, variétés qui ne sont peut-étre pas absolues et irréductibles.’’) ‘ OPINION 283 141 159. Having thus reached a taxonomic decision on the status of the Benign Tertian Malaria Parasite (the “* parasite tierce ’’) and the Quartan Malaria Parasite (the “‘ parasite quarte ’’), Labbé turned his attention to the crescent-formed parasite (“‘ les crois- sants’”’). As regards these latter, he considered that their claim to specific rank (“‘ spécificité’’) was poorly established (‘‘ mal établie ’’), for he agreed with Laveran that the crescent-formed parasites “‘se confondent avec les corps amoeboides ”’ and that small spherical elements gave birth to both these forms of parasite (‘de petits éléments sphériques donnent naissance a ces deux formes parasitaires”’). Labbé concluded, therefore, that the parasite in question, i.e. the Malignant Tertian Malaria Parasite, occurred in two forms, the one elongated and crescent-shaped, the other amoeboid. The passage in which Labbé gave this as his conclusion reads as follows :— . . . Nous pensons donc que, de méme que chez les Proteosoma et les Halteridium nous avons vu le parasite prendre soit une forme allongée, pseudo-grégarienne, soit une forme amoeboide; ces deux formes aboutissant uniformément a une forme ronde, puis a la sporulation; de méme, chez le parasite de malaria, que nous pouvons appeler Haemamoeba Laverani, nous pouvons voir deux formes parasitaires différentes: lune allongée, semi- lunaire, l’autre amoeboide. 160. Labbé summed up his views in regard to the taxonomy of the human malaria parasites in the following diagnosis (: 170):— HAEMAMOEBA Grassi Syn. Haematophyllum Metschnikoff, Oscillaria Laveran, Plas- modium Marchiafava et Celli, Haematomonas Osler, Haemamoeba +Laverania Grassi et Feletti. Parasites endoglobulaires du sang de l’homme caractérisées par un dimorphisme de structure: 1° Des formes amoeboides 4 mouvements assez vifs: 2° Des formes allongées, semi-lunaires, immobiles. Ces deux formes, 4 protoplasme hyalin, 4 structure d’amibes, douées d’un noyau vésiculaire a nucléole excentrique, aboutsissent a une forme ronde qui constitute une spore nue. Cette spore se divise en un petit nombre (quarte) ou en un grand nombre (tierce) de sporozoites nucléés groupés autour d’un reliquat et disposés en rosace ou en morula. 142 OPINIONS AND DECLARATIONS Il y a un stade de dégénérescence a flagella. Les parasites réduisent l’-héemoglobine en mélanine et ont une action pathogéne sur ’organisme. Le développement se fait en deux ou trois jours. Une seule espéce: Haemamoeba Laverani, comprenant des variétés H. L. var. quartana, H. L. var. tertiana. 161. The following conclusions may be drawn from the extracts from Labbé’s paper quoted above :— (1) Labbé considered that there was only one species of human malaria parasite. This species he placed in the genus Haemamoeba erroneously referred by him to Grassi instead of to Feletti & Grassi. He gave a new trivial name (/averani) to this species, clearly in honour of his compatriot Laveran, by whom the first of the human malaria parasites (the Malignant Tertian Malaria Para- site) had been originally described. Labbé did not indicate why he proposed to substitute this name for the name malariae Laveran, 1881 (published by Laveran in the combination Oscillaria malariae). Indeed at no ~ point in his paper did Labbé refer to the trivial name — malariae Laveran. (2) At the time when Labbé wrote his paper—and for some time thereafter—taxonomists commonly held the view that, if a racial or other variety was distinguished from a species previously regarded as made up of a single uniform population, it was appropriate to give a name to the newly distinguished racial or other variety but not necessary to distinguish by name the racial form to which the specific trivial name had originally been given. Accordingly, where an author stated that he recognised two varieties, his statement was equivalent, in the terms of the present Rég/es, to a statement that he recognised three subspecies, namely the nominotypical subspecies (which takes as its trivial name the name of the species itself) and two other subspecies. Labbé stated (: 169) that he recognised two varieties of the human malaria parasite, namely the “‘ parasite quarte ’’ (= the Quartan Malaria Parasite) and the “ parasite tierce” (=the Benign Tertian Malaria Parasite). In the light of the OPINION 283 143 explanation given above, this should be understood to mean that Labbé recognised a single collective species, consisting of three subspecies, namely :— (a) the nominotypical subspecies of Haemamoeba laverani Labbé, that is to say, the taxonomic unit originally described as Oscillaria malariae by Laveran in 1881, after whom this taxonomic unit was re-named by Labbé (=the parasite responsible for malignant tertian malaria fever); (b) the subspecies responsible for quartan malaria fever; (c) the subspecies responsible for benign tertian malaria fever. (3) Labbé added, on page 170, that what, in accordance with the then current terminology (explained in (2) above), he called the “‘ species ’’ (i.e. the nominotypical subspecies) itself occurred in two “forms” (the word “ forme ”’ being used in this connection in contrast to the word *“ varieté ’”’ employed by Labbé to denote the subspecies quartana Labbé and tertiana Labbé). The first of these forms was ‘“‘ amoeboide, 4 mouvements assez vifs,”’ the second “ allongée, semi-lunaire, immobile.” (4) The name Haemamoeba laverani var. quartana was given by Labbé to the “parasite quarte” (=the Quartan Malaria Parasite). (5) The name Haemamoeba laverani var. tertiana was given by Labbé to the “ parasite tierce’? (=the Benign Tertian _ Malaria Parasite.) 162. As will be seen from his diagnosis of the genus and its species (quoted in paragraph 160 above) Labbé referred to the Benign Tertian and Quartan Malaria Parasites (under their French names “ tierce’ and “‘ quarte ’’) and immediately thereafter gave the new trivial names tertiana and quartana to these parasites. Thus Labbé did give, for each of these parasites, an “‘ indication,” as required by proviso (a) to Article 25 of the Régles. The extreme brevity of the “ indication ”’ so given might have prompted 144 OPINIONS AND DECLARATIONS a doubt whether the data afforded were sufficient to enable a taxonomic judgment to be reached in regard to the identity of the taxonomic units to which the two foregoing names were applied, if it had not been for the fact that earlier in his paper (on page 168) Labbé gave a succinct definition both of the “ para- site tierce’’ and of the “ parasite quarte.”” The names fertiana Labbé and guartana Labbé may therefore both be accepted as having been duly published in the manner prescribed by the Régles. (It will be noted that, unlike the cheironyms tertianae and quartanae (discussed in paragraphs 134—136 above) commonly alleged to have been published as trivial names by Celli & Sanfelice in 1891, the trivial names tertiana Labbé, 1894, and quartana Labbé, 1894, were duly published in the nominative singular in agreement with the generic name and in the same gender as that name, as required by Articles 8 and 14 of the Régles.) 163. Having now established that the trivial names /Javerani, quartana and tertiana were duly published by Labbé in accordance with the provisions of the Rég/es and having identified the taxo- nomic units to which these names severally apply, we must turn to consider the status of each of these names in relation to other names given to the same taxonomic units. 164. The trivial name “ laverani”’ Labbé, 1894: The taxonomic unit to which Labbé applied the specific trivial name, /averani (but not the taxonomic units which he regarded as “‘variétés”’ of it and to which he applied the varietal (subspecific) names quartana and tertiana) was (as has been shown above) the Malignant Tertian Malaria Parasite. The trivial name /averani Labbé, 1894, is thus a junior subjective synonym of the malariae Laveran, 1881, as published in the combination Oscillaria malariae, which applies to the same parasite. Labbé quite clearly did not regard himself as describing a new species of parasite. What he was doing—and what he realised that he was doing—was to givea new specific trivial name toa species which had already been described and named. That species was undoubtedly the species which had first been named by Laveran, whom Labbé now associated with that species by using his name in the Latinised form “ Javerani”’ as its specific trivialname. Accordingly, the specific trivial name /averani Labbé 1894, as published in the combination Haemamoeba laverani, might reasonably be regarded as not only a subjective synonym, OPINION 141 145 but also as an objective synonym, of the specific trivial name malariae Laveran, 1881, originally bestowed upon the Malignant Tertian Malaria Parasite in the binominal combination Oscillaria malariae. 165. The trivial name “ quartana”’ Labbé, 1894: The trivial name quartana was (as already shown) applied by Labbé as a varietal (subspecific) trivial name to distinguish the taxonomic unit so named from the Malignant Tertian Malaria Parasite (=Oscillaria malariae Laveran, 1881) (renamed Haemamoeba laverani by Labbé), of which he considered that his guartana was a variety. The taxonomic unit to which Labbé applied the name quartana was the “ parasite quarte,” i.e. the Quartan Malaria Parasite. The only name which, prior to the publication of the name guartana Labbé, 1894, had been published for the Quartan Malaria Parasite is the name malariae Feletti & Grassi, 1889, as published in the combination Haemamoeba malariae. As already observed (paragraph 90) this latter name cannot be used for the Quartan Malaria by those malariologists—the mayjority—who regard this species as congeneric with the Malignant Tertian Malaria Parasite, for to these specialists both species are referable to the genus Plasmodium Marchiafava & Celli, 1885, in which genus the name malariae Feletti & Grassi, 1889, for the Quartan Malaria Parasite is a junior secondary homonym of the name malariae Laveran, 1881, the oldest available name for the Malig- nant Tertian Malaria Parasite. Nor, in my view, can the name malariae Feletti & Grassi be used for the Quartan Malaria Parasite even by those malariologists who do not regard the species so named as congeneric with the Malignant Tertian Malaria Parasite. For the fact that this name is a junior secondary homonym and must be rejected and replaced by authors who put the species so named in the same genus (Plasmodium) as the species malariae Laveran, 1881, means—under Articles 35 and 36—that this name is permanently rejected and cannot be used by anyone!’. 19 At the time when this passage was written, it was still a matter of argument whether a rejected secondary homonym (such as the name malariae Feletti & Grassi 1889, in the genus Plasmodium) was rejected permanently or whether it could be used by those authors who held a taxonomic view according to which the name in question was not a secondary homonym of some older name. As explained in Footnote 7, the Thirteenth International Congress of Zoology, Paris, 1948, decided that a name rejected and replaced for the foregoing reason can never be validly used again. 146 OPINIONS AND DECLARATIONS In these circumstances, under the Régles, the name gquartana Labbé, 1894, as published in the combination Haemamoeba laverani var. quartana, is the oldest available trivial name for the Quartan Malaria Parasite. 166. The trivial name “ tertiana’’ Labbé, 1894: The trivial name tertiana was (as already shown) applied by Labbé as a varietal (subspecific) trivial name to distinguish the taxonomic unit so named from the Malignant Tertian Malaria Parasite (= Oscillaria malariae Laveran, 1881) (renamed Haemamoeba laverani by Labbé), of which he considered his tertiana was a variety. The taxonomic unit to which Labbé applied the name tertiana was the ‘* parasite tierce,”’ 1.e. the Benign Tertian Malaria Parasite. The oldest available trivial name for that parasite is vivax Grassi & Feletti, 1890, as published in the combination Haemamoeba vivax, of which, therefore, the trivial name tertiana Labbé, 1894, is a junior subjective synonym. (20) The specific trivial name ‘‘ falciforme ’’ as published in combination with the generic name ‘‘ Haematozoon’”’ by Thayer and Hewetson in 1895 for the Malignant Tertian Malaria Parasite 167. As will be seen from paragraphs 107 above, Antolisei & Angelini in 1890 published a description of the Malignant Tertian Malaria Parasite, which they called ematozoo falciforme, which, for the reasons which I have explained, I consider is, on being corrected to Ematozoo falciforme, a name which must be regarded as an acceptable name under the Rég/es. In 1895 in the important work entitled “The Malarial Fevers of Baltimore” which occupies the whole of volume 5 of the Johns Hopkins Hospital Reports, Thayer & Hewetson, (: 27), without offering any explanation of what they were doing, introduced the name Haematozoon falciforme for the Malignant Tertain Malaria Parasite in a discussion of the paper by Antolisei and Angelini referred to above. In introducing this name, Thayer & Hewetson used a small letter instead of a capital for the first letter of the ee —_— J se el Se a ei, oe ime Ge) OPINION 283 147 generic name which appeared as haematozoon. It is likely, that Thayer & Hewetson, in taking this action, did not look upon themselves as publishing a new name, considering rather that they were correcting the defective name published five years earlier by Antolisei & Angelini. It is not possible, however, in a case of this kind to look behind the published words, in order to seek out the intention of the author concerned. For the reasons which I have explained fully when discussing the situation created by the publication of the generic name Ematozoo Antolisei & Angelini, 1890, with a small initial letter instead of with a capital letter (paragraphs 110 to 118 above), which is exactly parallel with the present case, I am of the opinion that the name /falciforme as published by Thayer & Hewetson is not invalidated by reason of the fact the generic name was written with a small initial letter (as haematozoon) instead of with a capital (as Haematozoon). It might however be argued that the name falciforme so published was no more than a usage of the same name as previously published by Antolisei & Angelini, but on the whole it seems better to treat it as a new name attributable to Thayer & Hewetson themselves.?° 168. The name falciforme Thayer & Hewetson, 1895, applies, as has been explained above, to the Malignant Tertian Malaria Parasite which ever since 1881, had possessed the available name malariae Laveran, (paragraph 45). Accordingly, the name falciforme Thayer & Hewetson, 1895, as published in the com- bination Haematozoon falciforme, is a junior subjective synonym of the name malariae Laveran, 1881, as published in the combina- tion Oscillaria malariae. Moreover, as the name /falciforme Thayer & Hewetson, 1895, applies to the same species as falci- forme Antolisei & Angelini, 1890, it is invalid by reason of being a junior secondary homonym of that name. 20 When at Copenhagen in 1953 the Fourteenth International Congress of Zoology revised Article 19 of the Régles (the Article governing the emendation of names), it included in the revised Article a provision that an invalid emendation (such as Thayer & Hewetson’s emendation to Haematozoon of the name previously published as Ematozoo by Antolisei & Angelini) is to be attributed to the authors by whom that emendation was made and is to bear priority as from the date on which it was published and not the earlier date on which the name in its unemended form was first published (1953, Copenhagen Decisions zool. Nomencl., : 45, Decision 73(3)). 148 OPINIONS AND DECLARATIONS (21—25) The terms “ tertianae,”’ ‘* quartanae,”’ “‘ undecimanae,” ‘““sedecimanae””’ and ‘“ vigesimo-tertianae ” published by Lewkowicz in 1897 in combination with the generic name ‘** Haemosporidium ” Lewkowicz, 1897, and _ erroneously alleged by later authors to have been published as specific trivial names for human malaria parasites 169. The issue of 6th February 1897 of the “‘ Erste Abteilung ”’ of the Centralblatt fiir Bakteriologie, Parasitenkunde u. Infektions- krankheiten (1897, Zbl. Bakt. (Erste Abt) 21(4) : 129—133) contains a short paper by Lewkowicz entitled ‘‘ Ueber den Entwickelungsgang und die Einteilung der Malariaparasiten,” in which he introduced a new classification of the human malaria parasites, which he presented (: 132—133) schematically as follows :— [132] Haemosporidia Entwickelungsdauer 2 und 3 Tage, I. Gruppe endoglobulare Entwickelung, erwachsene Formen sph§risch. a) Haemosporidium tertianae,Entwickelungsdauer 2 Tage, b) quartanae, SEOs [133] Entwickelungsdauer itiber 3 Tage, II. Gruppe extraglobulare Entwickelung, erwachsene Formen halbmondférmig. c) Haemosporidium z Entwickelungsdauer unbes- timmt (erwachsene Formen hdufig cigarrenf6rmig, s.o.) d) undecimanae, Entwickelungsdauer 10 Tage. e) : sedecimanae, : 15 f) : vigesimo-tertianae, : 22 170. The parasites, it will be noted, were divided into two main groups. The first comprised the ‘‘ Parasiten der Tertiana und Quartana ”’ and the second the “‘ Parasiten der sog. acyklischen Wechselfiebers ’ (: 130). The parasites of the first group were OPINION 283 149 characterised by Lewkowicz as having an endoglobular develop- ment, spherical shape in the adult forms, and a period of development of 2 to 3 days; those in the second group by extra- globular development, crescent shapes in the adult forms and a development period of more than 3 days. In the latter group Lewkowicz recognised four species. For the first of these species (a species in which the adult forms were stated to be frequently cigar-shaped) the period of development had not been determined but was considered to be probably somewhat shorter than that of the other species placed in the second group, this species thus forming a transition between the two groups (: 132). The periods of development of the three other species of the second group were given as 10, 15, and 22 days respectively. 171. The question now arises as to whether the terms “ ter- tianae”’ and “ quartanae’’ applied by Lewkowicz to the two species of his first group and the terms “ undecimanae,”’ *““ sedecimanae ” and “‘ vigesimo-tertianae ’’ applied to the second, third and fourth species of his second group should be regarded as new specific trivial names. 172. The parasites concerned are certainly distinguished by an indication, definition or description, as required by proviso (a) to Article 25 of the Régles and if the terms used were published as new names, they would undoubtedly possess rights under the Law of Priority as from the date of the publication of Lewkowicz’s paper. But, as explained in the case of the terms “ quartanae,” *“tertianae ’ and “ quotidianae ”’ as used by Celli & Sanfelice in 1891 (see paragraphs 134—136 above), an adjective, when used as a specific trivial name, must be placed in the nominative singular and in the same gender as the word constituting the generic name. In the present case the generic name (Haemo- sporidium) is a neuter noun and, in consequence, if Lewkowicz had intended that the Latin adjectives which he used in connection with these species should be regarded as specific or subspecific trivial names, he should have cited them as “ quartanum,” *“tertianum,” etc. In fact, as we have seen, he placed these adjectives in the genitive singular and cited them in the feminine gender. 150 OPINIONS AND DECLARATIONS 173. In these circumstances, it must be concluded that Lewkowicz used these Latin adjectives not as specific or sub- specific trivial names but as descriptive epithets. It must be noted however that, unlike the predecessors noted above, Lewkowicz used these adjectives in direct association with a generic name (Haemosporidium), without even the interposition of a noun acting as a specific trivial name. The use of these adjectives in the feminine singular in direct association with a generic name consisting of a neuter noun clearly shows that these adjectives were used by Lewkowicz as ordinary Latin adjectives applying to some noun, understood but not expressed, the noun in question being of the feminine gender and in the genitive case. The most probable explanation is that the noun in question was the word “malaria” and that the terms “ quartanae ”’ *“undecimanae ’’, etc., cited by Lewkowicz were regarded by him as being in grammatical agreement with the word “ malariae ”’ (i.e. the genitive singular of the word “ malaria’’). It is possible, however, that, as in the case of Celli & Sanfelice (paragraph 136), the word understood but not expressed was the word “ febris ” In this connection, it is important to note that, as shown in the table which he furnished, Lewkowicz was particularly concerned with the period of development (‘‘ Entwickelungsdauer ’’) of the various types of fever. Thus against the term “ quartanae,” Lewkowicz made the entry “‘ Entwickelungsdauer 3 Tage,’ thus indicating that in this case the period of development was three days and that the fever appeared on the fourth day. It is per- fectly clear that all five of the Latin adjectives used by Lewkowicz were employed to denote the period of development of the form of Haemosporidium concerned, and that these terms were not used by that author as specific or subspecific trivial names. If any confirmation of this conclusion were needed, it is provided by Lewkowicz’s treatment of the first form of Haemosporidium (lettered “‘a’’) in his “‘ II. Gruppe.” This is the only one of the Haemosporidia listed for which he did not know the period of development and it is also—no doubt, for this reason—the only one to which he did not apply a Latin adjective —— of an ordinal number. 174. If the five foregoing terms as published by Lewkowicz in 1897 had been acceptable as trivial names for human malaria OPINION 283 151 parasites, the first (“tertianae”) and second (“‘ quartanae ’’) would have been names for the Benign Tertian Malaria Parasite and the Quartan Malaria Parasite respectively, while the third (“‘ undecimanae ’’), fourth (‘‘ sedecimanae ”’) and fifth (“* vigesimo- tertianae ’’) would all have been names for the Malignant Tertian Malaria Parasite. For the reasons explained in the immediately preceding paragraph, it is clear, however, that these five terms were not published by Lewkowicz as scientific names and therefore that there are, in fact, no such names as the alleged names tertianae Lewkowicz, 1897, quartanae Lewkowicz, 1897, undeci- manae Lewkowicz, 1897, sedecimanae Lewkowicz, 1897, or vigesimo-tertianae Lewkowicz, 1897. (26) The specific trivial name ‘falciparum ” Welch as published in combination with the generic name ‘ Haematozoon” by Welch in 1897 for the Malignant Tertian Malaria Parasite 175. In 1897 (in Loomis’s Syst. Pract. Med. 1:36) Welch published the specific trivial name falciparum in the combination Haematozoon falciparum for the Malignant Tertian Malaria Parasite. (No definite evidence is available regarding the date in 1897 in which Welch’s book was published. It has accordingly here been treated as having been published after the paper by Lewkowicz discussed in the immediately preceding Section, as that paper is known to have been published as early as February 1897.) 176. In the passage referred to above Welch criticised the name falciforme Antolisei & Angelini, 1890, as published in the com- bination Ematozoo (=correction of ematozoo) falciforme (para- graph 102 above), on the ground that that name implied that the taxonomic unit so named was always falciform and that this carried with it the further implication that the name falciforme was applicable only to the crescentic forms. Welch accordingly proposed the name falciparum as being more appropriate, in that it indicated that “‘ the property of forming crescents is a distinctive character of the organism.” The present Rég/es, which were not 152 OPINIONS AND DECLARATIONS in existence at the time when Welch published the specific trivial name falciparum, provide (Article 32) that a scientific name, once published, cannot be rejected on the ground that it is inappro- priate. Accordingly, as the Régles have retroactive effect, Welch’s action in substituting the specific trivial name falciparum for the older name falciforme was incorrect. 177. For the reasons explained above, the trivial name falci- parum Welch, 1897, being only a nom. noy. pro the name falciforme Antolisei & Angelini, 1890, is a junior objective synonym of that name (and, incidentally, also of the name falciforme Thayer & Hewetson, 1895, as published in the com- bination Haematozoon falciforme) (paragraph 102 above) and is therefore an objectively invalid name. Even if this were not so, the name falciparum Welch, 1897, would not be required, for it is a junior subjective synonym of the trivial name malariae Laveran, 1881 (paragraph 45 above), the oldest available trivial name for the species to which both these names apply (namely the Malignant Tertian Malaria Parasite). Supplementary note on the cheironym ‘‘ tropica ’’ erroneously alleged to have been published in the combination ‘* Plasmodium tropica’’ by Koch in 1899 for the Malignant Tertian Malaria Parasite 178. In a letter dated 14th September 1944 Sir Rickard Christophers drew attention to the alleged existence of a specific trivial name tropica claimed to have been published by Koch in ~ 1899 in the combination Plasmodium tropica as a name for the Malignant Tertian Malaria Parasite. 179. The date on which this alleged name is claimed to have been published is two years subsequent to the close of the period (1881—1897) covered by the present paper and, if that were the date on which this name had actually been published, the problem created by that name would have no bearing upon the issues now OPINION 283 153 to be submitted to the International Commission on Zoological Nomenclature. For in that event the specific trivial name tropica would have been ante-dated by two years by the specific trivial name falciparum Welch 1897 (published as Haematozoonn falciparum), the specific trivial name now almost universally used for the Malignant Tertian Malaria Parasite and the name which for this reason it is desired should now be validated by the Inter- national Commission as the specific trivial name for that species. The name tropica has however a historical importance, for at one time it was widely used by German workers. For this reason and because, so long as the place (if any) in which this name was validly published remains undertermined, there is a risk that it may some day be found to have been published prior to the publication in 1897 of the specific trivial name falciparum Welch, I have felt it desirable to collect any information which might throw light on the question whether (and, if so, when), the name tropica was ever validly published as a specific trivial name. 180. Sir Rickard Christophers, writing on 14th September 1944, informed me that he had been unable to trace any paper in which the word tropica had been validly published by Koch as a specific trivial name, although later the word tropica attributed to Koch had been widely used in this sense for the Malignant Tertian Malaria Parasite by German authors. Koch had, however, used such expressions as “der tropischen Malariaparasiten”’ and * Parasiten der tropischen Malaria’ in his paper ‘‘ Ueber die Entwicklung der Malariaparasiten ”’ published in 1899 (Z. Hyg. InsectKr. 32 : 1—24); it was possible that it was through the use of these expressions in this paper that later authors had incor- rectly stated that in the same year (1899) Koch had published the specific name Plasmodium tropica (paragraph 178 above). Ina later letter (dated 16th October 1944), Sir Rickard reported that he had looked up all the papers by Koch that he could find but in none of these had Koch published the word tropica as a name in proper zoological form. In addition to the examples quoted above from his 1899 paper Koch had elsewhere used such expressions as “ Parasiten der Tropenfieber ” and “‘ Parasiten des febris tropica.” Schaudinn (1902, Arb. GesundhAmt. Berl. 19 : 169) and later authors had frequently used the expression * Tropicaparasit ’’ but had not published the word tropica as a 154 OPINIONS AND DECLARATIONS specific trivial name in a binominal combination as required in zoological nomenclature. Sir Rickard reached the conclusion that at most the name tropica had been applied to the Malignant Malaria Parasite as “‘ a semi-scientific name ”’ by workers engaged in clinical work. 181. Miss I. M. Bellis (4n the memorandum forwarded by Dr. C. M. Wenyon with his letter of 18th December 1944 to which reference had been made in paragraph 153 above in connection with the cheironym irregularis Sakharov, 1892) drew attention to the following statement by Ziemann (1906, in Mense’s Handbuch der Tropenkrankheiten 3: 356): “‘ Sacharoff nannte unser Perniciosa Febris meridiana. R. Koch nannte sie Tropica.” This statement, as Miss Bellis points out, referred to the fever and not to the parasite. Miss Bellis drew attention also to the fact that in the same work under the heading “ Einteilung der Malaria- parasiten ’ Ziemann mentions (: 279) “ Tropicaparasiten Koch’s”’ as a synonym of “ Perniciosaparasiten . . . der Italiener’’ and under ‘“ Die Perniciosaparasiten ’’ says (: 292): “‘ Robert Koch identifizierte dann die samtliche tropischen Perniciosaparasiten direkt mit den italienischen malignen Tertian-Parasitenformen . ..” but that in neither of these passages did Ziemann give the scientific name tropica for this species of parasite. In this con- nection aiso Miss Bellis cited the statement by Liihe (1906: 220) (to which reference has been made in paragraph 153 above in the discussion of the cheironym irregularis) that, in his view, there was so much confusion in the nomenclature of the human malaria parasites that it was much better to use vernacular names when referring to those species. Miss Bellis then pointed to the fact that in spite of this statement Liihe had given as full a list as he could of the scientific names so far bestowed upon the human malaria parasites, but that this list contained no reference to any specific trivial name consisting of the word tropica. 182. In the same memorandum Miss Bellis reported the result of a special examination which she had made of Koch’s malaria papers reprinted in Teil 1 of Band 2 of the ‘“‘ Gesammelte Werke von Robert Koch” published at Leipzig in 1912 and furnished the following extracts to illustrate the way in which Koch had OPINION 283 155 used the Latin adjective “‘ tropica”’ and the German adjective “tropischen ”’ in discussing Malignant Tertian Malaria:— (1) Berichte des Geh. Medizinalrats Prof. Dr. R. Koch tiber die (2) Ergebnisse seiner Forschungen in Deutsch Ost-Afrika, 1898 The following statements occur :— Die tropische Malaria unterscheidet sich von den anderen Arten der Malaria in vielfacher Beziehung, am deutlichsten aber durch den eigentiimlichen Krankheitsverlauf und durch die besonderen Blutparasiten, welche regelm4ssig bei derselben angetroffen werden... (: 308). Der Blutparasit des hiesigen Tropenfiebers entspricht im ubrigen volkommen der Beschreibung, welche von den Parasiten der angeblichen quotiadianen Malaria in anderen tropischen Landern von verschiedenen Forschern gegeben su (: 310). (A description with several figures follows, but no scientific name is given.) Uber die Entwicklung der Malariaparasiten, 1899 (Z. Hyg InsectKr. 32) The following passages occur :— Zu den echten Malariaparasiten, welche den angegebenen Merkmalen entsprechen, sind bisjetzt folgende Arten zu rechnen :— 1. Der Parasit des quartanen Fiebers 2. Der Parasit des tertianen Fiebers Beide sind durch die Untersuchungen von Golgi hin- reichend charackterisiert. 3. Der Parasit des Tropenfiebers .... Fur diesen Parasit ist durch Marchiafava und seine Schiller nachgewiesen, dass es eine besondere Art bildet und einen analogen endogenen Entwicklungsgang besitzt, wie die Parasiten der quartanen und tertianen Fiebers. (No scientific names were given by Koch in this passage.) 156 OPINIONS AND DECLARATIONS Von den drei Malariaparasiten des Menschen ist die endogene Entwicklung hinreichend bekannt und ich habe den Beschrei- bungen Golgis in bezug auf die Quartan- und Tertianparasiten und den Angaben Marchiafavas tiber die Tropenparasiten (identisch mit den sogenannten Astivo-Autumnalparasiten) eigentlich nichts hinzufiigen. (In this passage also no scientific names were given by Koch.) (3) Zusammenfassende Darstellung der Ergebnisse der Malaria- expedition, 1900 (Dtsch. med. Wschr., Nos. 49 u 50) Wir haben es somit nur noch mit den drei erwahnten Arten der Malaria zu tun. Aber wie sollen wir die dritte Art benennen. Ofters begegnet man die Bezeichnung “‘ Estivo- autumnalfieber,” in neuerer Zeit wird der Name “ maligne Tertiana”’ haufig gebraucht. Beide Namen scheinen mir nicht zweckmassig zu sein. Der Ausdruck “ Estivoautum- nalfieber’’ passt fiir die Tropen, wo doch diese Art von Malaria eigentlich zu Hause ist gar nicht; wollte man sie ‘“maligne Tertiana’’ nennen dann musste man die Tertiana des gemassigten Klimas zum Unterschied von jener als ‘““ benigne Tertiana’’ bezeichnen und zweien Taufen vorneh- men. Es scheint mir das richtigste zu sein, mit Riicksicht auf die eigentliche Heimat der Krankheit den von jeher gebrauchten Namen ‘“ Tropenfieber”’ beizubehalten und die spdteren lokalen Eindriicken entsprechende Bezeichnungen fallen zu lassen. (In this as in the previous passages anotg Koch did not give a scientific name.) 183. Finally, Dr. G. Robert Coatney (in a letter dated 13th September 1945) reported that he also had been unable to trace the use of the word tropica as part of a scientific name for a malaria parasite. 184. The conclusions which I draw from the evidence sum- marised above are:— (i) that Koch realised very clearly that the fever to which he applied the expression “ tropische Malaria” was the OPINION 283 (a same as the aestivo-autumnal fever found in temperate climates, i.e. the fever due to the Malignant Tertian Malaria Parasite; (ii) that Koch never published the word “tropica” as a specific trivial name in a binominal combination formed for the Malignant Tertian Malaria Parasite, for, if he had done so, he would (according to his own view, as summarised in (i) above) have only thereby given a new scientific name to a species already recognised (as Koch himself stated) by Marchiafava, i.e. he would only have created a junior synonym for the Malignant Tertian Malaria Parasite; (ili) that, when Koch did use the Latin word “ tropica’’ he used it not as a scientific name for the parasite but as a convenient technical designation for the fever to which the parasite gives rise. 185. The possibility that the name tropica was published by Koch in due zoological form may therefore be regarded as extremely remote. In the circumstances I think that the proper course is to treat the alleged name tropica Koch as a cheironym. Nevertheless, the possibility that this name was validly published in some paper which it has been impossible to trace cannot be entirelyruled out. This, as in the case of the cheironymirregularis, - is a risk which should, therefore, be taken into account by the International Commission on Zoological Nomenclature, when it comes to formulate its decision on the complex of problems raised in the present paper. PART 2 THE NAMES OF THE NOMINAL GENERA ESTABLISHED (OR ALLEGED TO HAVE BEEN ESTABLISHED) FOR THE HUMAN MALARIA PARASITES IN THE PERIOD 1881-1897 OPINION 283 159 186. In Part 1 of the present paper we have firmly established in the case of each specific trivial name bestowed upon a human malaria parasite in the period 1881-1897, the identity of the species to which the name was applied. We can now turn to the generic names given to these parasites during the same period, and, having ascertained, for each generic name, whether it is valid in the sense that it is not a junior homonym of some other name and having determined what is the nominal species which is the type species of the nominal genus to named, we shall be able to determine which generic names are available nomenclatorially and which either are invalid as junior objective synonyms of older names or, although available, are not required because of being junior subjective synonyms of older names. (1) The generic name “ Oscillaria ” Schrank, 1823, and the generic name “ Oscillaria ” erroneously alleged to have been published as a new name by Laveran in 1881 187. The first scientific name bestowed on any of the human malaria parasites was the name Oscillaria malariae given by Laveran in 1881 (Nature parasit. Accid. Impaludisme: 87) to the Malignant Tertian Malaria Parasite (paragraph 45 above). 188. Oscillaria was thus the first generic name used for a human malaria parasite, but a great deal of controversy has arisen as to its validity in view of the fact that at the time when Laveran first used this name for his malaria parasite (Oscillaria malariae, in 1881), there already existed a genus Oscillaria Schrank, 1823 (Nova Acta Acad. Caes. Leopold. Carol. 11 (No. 2): 533), which had been established for certain species which are now recognised to be algae. 160 OPINIONS AND DECLARATIONS 189. Two distinct questions arise in considering the validity of the generic name Oscillaria as used by Laveran in 1881 :— (1) Had the generic name Oscillaria Schrank, 1823, ever been applied to a genus in the Animal Kingdom ? (11) When Laveran used the generic name Oscillaria in first describing the human malaria parasite to which he applied the specific trivial name malariae, did he consider that he was introducing a new generic name for this species or did he consider that he was refering that species to the existing genus Oscillaria Schrank, 1823 ? 190. The status of the name Oscillaria Schrank in zoological nomenclature depends on whether the nominal genus to which it was applied has ever been regarded as a genus in the Animal Kingdom. To determine the answer to this question it is neces- sary to examine closely the manner in which in 1823 Schrank | introduced this generic name. It should be noted first that this name was published by Schrank in the serial publication cited in paragraph 188 above in a paper entitled “‘ Uber die Oscillatorien.” In this paper he included the following units in this group:— (i) Bacillaria. Schleichthieren. Under this head Schrank included a number of species previously described with the generic name Vibrio. (ii) Oscillaria. Oscillarie. Under this head Schrank included thirteen species previously described under the generic name Oscillatoria and four species previously described with the generic name Vibrio. (iii) Vibrio. Aelchen. Schrank placed eight species in this genus. OPINION 283 161 191. Schrank discussed the question of whether the species belonging to the group (Oscillatoria) dealt with in his paper were properly referable to the Animal Kingdom. On the one hand, he pointed, first, to their method of reproduction (‘‘ durch Theilung ’’) as suggestive of the Vegetable Kingdom, and second, to the fact that the organisms in question had been dealt with by Vaucher in a book on Freshwater Confervae (Hist. Conferv. : 172—178). On the other hand, he remarked with regard to the first point that reproduction “‘ durch Theilung”’ is not peculiar to the Vegetable Kingdom but has been observed also among small animals (“ Thierchen ’’) and with regard to the second point that in dealing with the species in question Vaucher “sah sie niemals fiir Pflanzen an, spricht von ihnen, wie man von Thieren spricht, und nennt sie sogar ausdriicklich Thierchen.”’ 192. At no point in his paper did Schrank express a definite view as to whether the organisms placed by him in the group Oscillatoria were to be referred to the Animal Kingdom or to the Vegetable Kingdom or possibly to some intermediate category. It seems to me, however, that the fact that Schrank himself in his original description of the genus consistently referred to the organisms concerned as “ Thierchen ”’ is sufficient to warrant the conclusion that the genus Oscillaria Schrank, 1823, was originally established as a genus of animals, for the foregoing expression would have been entirely inappropriate for species of a genus of plants. 193. We now come to the second of the two questions referred to in paragraph 189 above, namely that as to whether Laveran deliberately placed his malaria parasite in the genus Oscillaria Schrank, 1823, or whether by some accident he selected the word * Oscillaria”’ as the name for a new genus established by him, especially for the reception of his new species (the Malignant Tertian Malaria Parasite). We find in the first place that Laveran was certainly aware of Schrank’s paper, and this fact alone makes it unlikely that he would have chosen the name Oscillaria if he had intended to establish a new nominal genus. Nor would it seem intrinsically unnatural for Laveran to have regarded his malaria parasite as sufficiently closely related to the minute 162 OPINIONS AND DECLARATIONS motile organisms placed by Schrank in his genus Oscillaria to justify him in assigning his new species to the same genus. This view is strongly reinforced by Laveran’s own statement that the mobile filaments of what he called the “corps no. 2” often showed a great analogy with the “ oscillariées ” (see the penulti- mate paragraph of the passage quoted in paragraph 35 of the present paper). . In the second place, the fact that Laveran himself later abandoned the use of the generic Oscillaria is more consistent with the thesis that he realised that Schrank’s genus was unsuit- able for the reception. of his new malaria parasite (possibly because he had come to the view of other zoologists and botanists that it belonged to the Vegetable Kingdom and not to the Animal Kingdom) than with the alternative view that he deliberately abandoned a new name of which he was himself the author. On the whole, therefore, we can dismiss the possibility that, when in 1881 Laveran published the specific name Oscillaria malariae for the Malignant Tertian Malaria Parasite, he regarded himself as introducing a new generic name Oscillaria, for it may be con- cluded that he considered that he was describing his new species as a species of the genus Oscillaria Schrank, 1823. 194. In the light of the above considerations, we have now to consider the status under the Régles of the generic name Oscillaria as applied to the Malignant Tertian Malaria Parasite. The answer turns upon the interpretation to be placed upon Article 1 of the Régles when read in conjunction with Article 34. The point to be considered is the status in zoological nomenclature of a generic name applied to a genus originally regarded as belonging to the Animal Kingdom but subsequently transferred, on taxo- nomic grounds, to the Vegetable Kingdom. 195. Christophers & Sinton (1938: 1133), in considering the validity of the generic name Oscillaria as used by Laveran in 1881, concluded that “ there could be no question that Laveran’s name, being given to an animal, could not strictly be regarded as pre- © occupied by a genus of the Vegetable Kingdom.” This inter- pretation appears to have been based not upon the relevant portion of the substantive French text of Article 1 of the Régles OPINION 283 163 (or even upon the correct German translation) but upon the inaccurate English translation commonly in use. The substantive French texts of Article 1 (third paragraph) and Article 34 read as follows :— Article 1, paragraph (3) Si un étre est transporté du Régne animal dans le Regne végétal, ses noms zoologiques sont maintenus dans la nomenclature zoologique. Article 34 Tout nom générique est rejeté comme homonyme, s’il a été employé précédemment pour quelque autre genre d’animaux. 196. It is perfectly clear from the substantive French text of Article 1 that, if a taxonomic unit originally described as belonging to the Animal Kingdom is later regarded as belonging to the Vegetable Kingdom, the names given to that unit when it was believed to belong to the Animal Kingdom are maintained in zoological nomenclature. Applying this provision to the parti- cular case with which we are concerned, this means that, when the genus (supposedly of animals) named Oscillaria by Schrank in 1823, was later treated as belonging to the Vegetable Kingdom, the generic name Oscillaria Schrank, 1823, nevertheless retained © its position in zoological nomenclature. Accordingly, Article 34 still applies to that name, and in consequence any generic name consisting of the word Oscillaria published subsequent to 1823 as the name of a new genus of animals is, under that Article, to be rejected as a junior homonym. *1 The inaccurate English translation of Article 1, which appears to have been responsible for the misunderstanding, reads as follows :— Zoological nomenclature is independent of botanical nomenclature in the sense that the name of an animal is not to be rejected simply because it is identical with the name of a plant. If, however, an organism is transferred from the vegetable to the animal kingdom, its botanical names are to be accepted in zoological nomenclature with their original botanical status; and if an organism is transferred from the animal to the vegetable kingdom, its names retain their zoological status. The italicised passage is supposed to be a translation of the substantive French text of the paragraph (3) of this Article quoted in paragraph 195 of the present paper. 164 OPINIONS AND DECLARATIONS 197. The position as regards the validity of the generic name Oscillaria as applied by Laveran to the Malignant Tertian Malaria Parasite may thus be summarised as follows :— (i) If Laveran regarded himself as establishing in 1881 a new nominal genus named Oscillaria for the reception of the Malignant Tertian Malaria Parasite, the generic name Oscillaria Laveran, 1881, then published for the nominal genus so established, is invalid by reason of its being a junior homonym of the generic name Oscillaria Schrank, 1823, a generic name originally published for what was believed at that time to be a genus of animals but which was later recognised to be a genus of plants. (ii) If Laveran in 1881 deliberately placed the Malignant Tertian Malaria Parasite in the genus Oscillaria Schrank, 1823, he was simply using the latter name in a taxo- nomically incorrect sense. 198. The conclusion is, therefore, that the generic name. Oscillaria, as applied to the Malignant Tertian Malaria Parasite, is either nomenclatorially invalid or taxonomically incorrect. For the reasons explained in the preceding paragraphs, I believe that the second of these alternatives represents what really happened. In consequence, I am of the opinion that Laveran never looked upon the name Oscillaria as the name of a new nominal genus established by himself. However, this is one of the many matters on which it is desirable that a definite ruling should be given by the International Commission on Zoological Nomenclature when it comes to stabilise the names of the human malaria parasites. (2) The generic name ‘‘ Plasmodium ” Marchiafava and Celli, 1885 199. The generic name Plasmodium was first published by Marchiafava & Celli in 1885 (Fortschr. Med. 3 (24):791) as a generic name for what those authors regarded as the malaria parasite of Man. It is an available name in the sense that it is not a junior homonym of any previously published generic name. OPINION 283 165 200. Until the discovery described in paragraphs 55 to 56 of the present paper, it was always assumed (a) that Marchiafava & Celli placed no nominal species in their genus Plasmodium, and (b) that the single species which they did place in that genus was the species to which in 1881 Laveran had given the name Oscillaria malariae. As we shall see, both these assumptions were incorrect. 201. It has been shown earlier in the present paper (paragraph 55 above) that, when introducing the new generic name Plas- modium Marchiafava & Celli published also the new specific name Plasmodium malariae for the single species included by them in their new genus. This nominal species is therefore the type species of the genus Plasmodium Marchiafava & Celli by monotypy (Article 30, Rule (c)). 202. Up till the year 1946 it was universally assumed by malariologists that the species placed in the genus Plasmodium by Marchiafava & Celli was the Quartan Malaria Parasite, and that species has therefore been regarded by all authors as the type species of this genus. Sabrosky (1946), by going back to Marchiafava & Celli’s paper of 1885, was the first author to establish that the material treated by those authors as constituting the species placed by them in the genus Plasmodium included no individuals at all of the Quartan Malaria Parasite. The evidence provided by Dr. Martin Young included in Sabrosky’s paper showed that the greater part of this material consisted of the Malignant Tertian Malaria Parasite, though it included also some examples of the Benign Tertian Malaria Parasite. In order to give to the nominal species Plasmodium malariae Marchiafava & Celli, 1885, an entirely determinate character, I have in the present paper (paragraph 76) selected as a First Reviser (acting under Article 31) the examples of the Malignant Tertian Malaria Parasite to be the sole syntypes of the nominal species Plasmodium malariae Marchiafava & Celli. This action firmly attaches the foregoing name to the Malignant Tertian Malaria Parasite. 166 OPINIONS AND DECLARATIONS 203. The only generic name applied to any human malaria parasite prior to the publication of the name Plasmodium Mar- chiafava & Celli, 1885, was the name Oscillaria as used by Laveran in the combination Oscillaria malariae when in 1881 he first named the Malignant Tertian Malaria Parasite (paragraph 45). I have shown, however in paragraphs 197 to 198 above, that the name Oscillaria as used by Laveran was not a new name but represented only a taxonomically erroneous use of the earlier name Oscillaria Schrank, 1823. 204. We may therefore sum up the position as follows: The generic name Plasmodium Marchiafava & Celli, 1885, is the oldest available generic name for any species of human malaria parasite, and its type species by monotypy is the nominal species Plas- modium malariae Marchiafava & Celli, 1885, a nominal species which represents the Malignant Tertian Malaria Parasite. The fact that this species and not the Quartan Malaria Parasite is the type species of this genus would, if not remedied by the Inter- national Commission on Zoological Nomenclature under its Plenary Powers, cause serious confusion in the literature. That the Quartan Malaria Parasite had for so long been accepted as the type species of the genus Plasmodium Marchiafava & Celli, is undoubtedly attributable to the uncritical acceptance by all subsequent workers (up to the time of Sabrosky (1946)) of the authoritative but totally incorrect action of Schaudinn (1902) in designating the Quartan Malaria Parasite (misidentified by him, following Liuhe (1900), as Oscillaria malariae Laveran, 1881) as the type species of Plasmodium Marchiafava & Celli, 1885 (“‘ Plasmodium malariae (Der Quartanparasit) als typus der Gattung’’). A discussion of the probable origin of this erroneous determination will be found in paragraphs 46—51 above. (3) The generic name ‘‘ Haematomonas”’ Mitrophanow, 1883, and the generic name ‘‘ Haematomonas ” erroneously alleged to have been published as a new name by Osler in 1886 205. In 1886 in an “‘ Address on the Haematozoa of Malaria ” (1886, Philad. med. Times and 1887, Brit. med. J. 1887 (Vol. 1) : 556) Osler discussed the systematic position of “ the” human malaria parasite and expressed the view that, pending the acquisition of a OPINION 283: 167 better understanding of the affinities of this parasite, it should be referred to the-genus Haematomonas. Osler stated that this view had been suggested by Mitrophanow but he did not cite a reference to any paper by that author on this point. The following is an extract of the relevant portion of the account of Osler’s “* Address”’ published in the issue of the British Medical Journal referred to above :— There is sufficient evidence to show that the various forms are only phases in the life history of one of the flagellate protozoa belonging to the order Flagellata-Pantostomata. Mitrophanow suggests a new genus, Haematomonas, to include the monad haematozoa; but Crookshank, who has carefully worked out the affinities of the parasites of the rat, the fish and the surra disease, has referred them to the genus Trichomonas. The organism here described has not however, the characteristic marks of a Trichomonas; for it lacks the undulating fringe on one side and the caudal filament. Nor does it agree with the features of a Cercomonas; so that, meanwhile, until the true affinities are determined by an expert, its proper place seems to be the genus Haematomonas of Mitrophanow, which conveniently includes all monads parasitic in the blood. Thus: genus Haema- tomonas; species Haematomonas malariae. [no italics]. Defini- tion: Body plastic, ovoid or globose, no differentiation of proto- plasm, which contains pigment grains; flagella variable from one to four. Highly polymorphic, occurring in (1) amoeboid form, (2) crescents, encysted form; (3) sporocysts; (4) circular, free, pigmented bodies. The name designates the natural affinities of the parasite, its habitat and the conditions under which it occurs, on which grounds it seems preferable to that of Plasmodium malariae, suggested by Marchiafava & Celli. 206. Partly because Osler in his ‘“‘ Address’ did not cite any bibilographical reference for the name Haematomonas and partly, it may be supposed, because he adopted Haematomonas as the generic name for “‘ the’ malaria parasite (all the forms known to him being treated as forms of a single polytypic species), it has been commonly assumed that at the time when Osler’s ‘‘ Address ”’ was published, the name Haematomonas was no more than a manuscript name of Mitrophanow’s and, therefore, that Osler himself, by publishing this name, had become its author. In 1894, for example, Labbé (1894, Arch. Zool. expér. gén. (3) 2 : 170) cited the generic name “ Haematomonas Osler” among the 168 OPINIONS AND DECLARATIONS synonyms of Haemamoeba Grassi [recte Feletti & Grassi], the genus in which he placed the single species (Haemamoeba laverani) of human malaria parasite which he recognised. The alleged generic name Haematomonas Osler appeared again as recently as 1945 when it figured in a synonymy of the generic names of the human malaria parasites published by Christophers (1945, Rev. Inst. Salubridad y Enfermedades trop. 6 (4) : 216—219). 206. Actually, the name Haematomonas was published by Mitrophanow in a paper entitled “‘ Beitrage zur Kenntniss der Hamatozoon”’ published in 1883 (Biol. Zb/. 3:41), i.e. three years before the appearance of Osler’s “‘ Address.” In this paper Mitrophanow established the new genus Haematomonas for the reception of two new species of flagellate, the one Haematomonas cobitis found in the blood of a Mudfish (‘‘ Schlammpeizger ’’), the other Haematomonas carassii found in the blood of the carp (“ Karausch ”’). 207. With this information at our disposal, we see that what happened was that Osler was aware that Mitrophanow was of the opinion that “‘ the ”’ human malaria parasite was congeneric with the flagellates which he had described from the blood of the fishes referred to above and therefore considered that the human malaria parasite should be referred to the genus Haematomonas Mitrophanow, 1883. This view was provisionally accepted by Osler. 208. The position is, therefore, that Osler never published the name Haematomonas as a new generic name and that there is in consequence no such name as “‘ Haematomonas Osler.” This alleged name owes its existence entirely to a misunderstanding by later authors of Osler’s paper, due to their failure to read Mitrophanow’s paper of 1883. In these circumstances, the alleged name Haematomonas Osler, 1883, should be deleted from the synonymy of the names of the genera established for the human malaria parasites, though it must be included in any list of generic names erroneously supposed to have bestowed upon those parasites. OPINION 283 169 (4) The generic name “ Haematophyllum ” Metschnikoff, 1887 209. The generic name Hdmatophyllum was first published by Metschnikoff in a paper written in Russian and published in the Russian serial Russkaja Medicina (1887, Russk. Med. 12 : 207). It has not been possible to consult a copy of this paper, as there appears to be no copy of the above serial in Western Europe. Fortunately, however, Metschnikoff himself wrote an abstract of this paper which was published in the same year in the Centralblatt fiir Bakteriologie und Parasitenkunde (1887, Zbl. Bakt. 1 (21): 624—625). According to this abstract, which, though signed by Metschnikoff, is written in the third person, the title of his paper (when translated into German), was “ Zur Lehre von den Malaria Krankheiten.”’ In this paper Metschnikoff described some pathological-anatomical material obtained from the Bacterio- logical Station at Odessa. Of the three sets of material examined the two first consisted of amoeboid forms similar to those observed by previous workers (e.g. Marchiafava, Celli, Golgi); the third appeared to him to represent a stage of development not previously noticed. He went on to say that the name previously suggested for the malaria parasite, namely Plasmodium malariae (i.e. Plasmodium malariae Marchiafava & Celli, 1885) was unsuit- able and that he therefore proposed the new name Hdmatophyllum malariae. The words used by Metschinikoff were: “‘ Auf Grund seiner Befunde glaubt M. [=Metschnikoff], in dem Malaria- parasiten eine in die Nahe der Coccidien zu stellende Form erblicken zu k6nnen, welche er (wegen der Unbrauchbarkeit des_ vorgeschlagenen Namens: Plasmodium malariae) als Hdmato- phyllum malariae zu bezeichnen vorschlagt.” 210. It is clear from the passage quoted above that the generic name Hdmatophyllum was proposed by Metschnikoff as a nom. noy. pro the generic name Plasmodium Marchiafava & Celli, 1885, which he rejected solely on account of (as it seemed to him) the unsuitability of the word “ Plasmodium ”’ as the generic name for the malaria parasites of Man. According to Article 32 of the present Régles (which, however, it may be noted were not adopted until many years after Metschnikoff wrote his paper), a generic name, once published, may not be rejected on the ground of 170 OPINIONS AND DECLARATIONS inappropriateness, even by its own author (“‘ Un nom générique.., — une fois publié ne peut plus étre rejeté pour cause d’impropriété, | méme par son auteur’’). Accordingly, the name Hdmatophyllum | Metschnikoff, 1887, is no more than a substitute name for Plasmodium Marchiafava & Celli, 1885. ) 211. The orthography of this name has been expanded from © ‘““Hamatophyllum” to “ Haematophyllum” by those later — authors who have discussed Meschnikoff’s paper (e.g. Labbé, © 1894, Arch. Zool. expér. gén. (3) 2: 170; Christophers, 1945, Rey. | Inst. Salubridad y Enfermedades trop. 6 (4) : 216, 219). Strictly” speaking, however, this procedure is incorrect under the existing — Régles which require (Articles 19 and 20) that the exact original — orthography of a name is to be retained.”? 212. Since the name Hdmatophyllum Metschnikoff, 1887, is a © mere substitute name (nom. noy.) for the name Plasmodium — Marchiafava & Celli, 1885, it automatically takes, as its type species, whatever nominal species is the type species of the genus Plasmodium. That species, as shown in paragraph 201 above, is © Plasmodium malariae Marchiafava & Celli, 1885. The nominal species so named represents the Malignant Tertian Malaria Parasite (paragraph 45 above). 213. Although not invalid as a junior homonym of some older name, the name Hdmatophyllum Metschnikoff, 1887, is neverthe- — less an objectively invalid name, since it has, as its type species, © a nominal species (Plasmodium malariae Marchiafava & Celli, — 1885) which is also the type species of a nominal genus (Plas- — modium Marchiafava & Celli, 1885), the name of which is an available name and one which possesses priority over Hdmato-_ phyllum Metschnikoff, 1887. 22 Article 20, jointly with Article 19, was completely revised by the Fourteenth ~ International Congress of Zoology, Copenhagen, 1953. Undertherevised provisions diacritic marks are not to be used in zoological names and, where such a mark was ~ used on the first publication of a name, it is to be replaced by an appropriate com- | bination of letters (1953, Copenhagen Decisions on Zoological Nomemclature: | 57—58). Under this provision, the correct orthography for the name here under — consideration is Haematophyllum. OPINION 283 171 (5) The generic name ‘“ Haemamoeba ”’ Feletti and Grassi, 1889 214. The generic name Haemamoeba was first published by Feletti & Grassi in a paper entitled “‘ Sui parassiti della malaria.” This paper was (as explained in paragraph 78 above) first pub- lished in the form of an eleven-page pamphlet on 30th December 1889 and just over a fortnight later (on 15th January 1890) it appeared in the Riforma medica of Naples (Feletti & Grassi, 1890, Riforma med. 6 (11) : 61—64). No copy of the pamphlet pub- lished in advance of the appearance of this paper in the Riforma medica has been traced and in consequence it is not possible to quote the page reference for the publication of the generic name Haemamoeba. The following notes are based upon an examina- tion of the photostat copy of the issue of the Riforma medica containing this paper by Feletti & Grassi (which is reproduced in facsimile as Document No. 1, in Appendix 2 to the present paper). 215. In this paper Feletti & Grassi, after reviewing the work of their immediate predecessors, came to the conclusion that human malaria was due to two parasites, each referable to a distinct genus. For the species described by Laveran, Feletti & Grassi erected the new genus Laverania, while for the second species they erected the genus Haemamoeba. The passage in which these two generic names were thus published reads as. follows (: 63):— In ogni caso nessun zoologo, specialmente dopo le inoculazioni di Antolisei e Angelini, esitera ad ammettere che il corpo pig- mentato della terzana e della quartana é differente dalla semiluna. Noi proponiamo di riferire ’uno al nuovo gen. Haemamoeba e laltro al nuovo gen. Laverania. 216. On the following page (: 64) Feletti & Grassi (im the passage quoted in paragraph 81 above) referred to their new species (that responsible for the “terzana’’ and “ quartana ”’ fevers) under the new specific name Haemamoeba malariae, at the same time referring Laveran’s original species (Oscillaria malariae Laveran, 1881) to their new genus Laverania. 172 OPINIONS AND DECLARATIONS 217. Two preliminary points must be noted: first, the name ~ Haemamoeba was published by Feletti & Grassi with a brief ‘“‘ indication ’’ (as required by proviso (a) to Article 25 of the Régles) and is thus a validly published name possessing rights — under the Law of Priority; second, the name Haemamoeba Feletti & Grassi, 1889, is an available name in the sense that it is not a homonym of any previously published generic name (and is therefore not invalid under Article 34). 218. Since Feletti & Grassi included in this genus the single nominal species Haemamoeba malariae Feletti & Grassi, the genus Haemamoeba Feletti & Grassi was a monotypical genus from the standpoint of its authors. As shown in paragraphs 83—86 above, the species Haemamoeba malariae Feletti & Grassi, 1889, was, however, a composite species consisting partly of the Benign Tertian Malaria Parasite (“‘ terzana’’) and partly of the Quartan Malaria Parasite (“‘ quartana’’). In May of the following year in a postscript (quoted in paragraph 85 above) to a paper by the same authors (in which, however, their names were placed in the reverse order and therefore appeared as “‘ Grassi & Feletti’’) the Benign Tertian Malaria Parasite was recognised as a species distinct from the Quartan Malaria Parasite (and given the name Haemamoeba vivax), the earlier name Haemamoeba malariae Feletti & Grassi, 1889, being thus restricted to the Quartan Malaria Parasite. 219. The present Régles were not in existence when Grassi & Feletti thus restricted the specific name Haemamoeba malariae Feletti & Grassi, 1889, to the Quartan Malaria Parasite, but, in thus acting as the “first reviser’’ of the name, these authors proceeded in strict accordance with the provisions of Article 31 of the present Régles. 220. The nominal genus Haemamoeba Feletti & Grassi, 1889, is accordingly a monotypical genus with Haemamoeba malariae Feletti & Grassi, 1889 (=the Quartan Malaria Parasite) as its type species. This was the first occasion on which a nominal genus was established with the Quartan Malaria Parasite as type OPINION 283 173 species and, as the name Haemamoeba Feletti & Grassi, 1889, is a nomenclatorially available name, it is the valid generic name for the Quartan Malaria Parasite for all malariologists who consider that species as generically distinct from the Malignant Tertian Malaria Parasite. For those malariologists—the great majority— who consider that these two parasites should be referred to the same genus, the generic name Haemamoeba Feletti & Grassi, 1889, is a junior subjective synonym of Plasmodium Marchiafava & Celli, 1885, of which the Malignant Tertian Malaria Parasite (as Plasmodium malariae Marchiafava & Celli, 1885, a junior sub- jective synonym of Oscillaria malariae Laveran, 1881) is the type species (paragraph 204). (6) The generic name ‘ Laverania ” Feletti and Grassi, 1889 221. The generic name Laverania was first published by Feletti & Grassi in the same paper as the generic name Haemamoeba (discussed in paragraph 214 above), that is to say in the pamphlet * Sui parasiti della Malaria” published on 30th December 1889 as an advance issue of the same paper which had been sent for publication to the Riforma medica, in which serial it appeared about a fortnight later (on 15th January 1890) (Feletti & Grassi, Jan. 1890, Riforma medica 6 (11): 63). In view of the fact that it is not been possible to trace a copy of the pamphlet of 30th December 1889, it is not possible to quote a page reference for the original publication of the generic name Laverania. The following notes are based upon an examination of the photostat copy of the issue of the Riforma medica containing this paper by Feletti & Grassi, which is reproduced as Document No. | in Appendix 2 of the present paper. 222. Felletti & Grassi established their nominal genus Laverania jointly with the other nominal genus, Haemamoeba, which they then recognised. The passage containing their diagnoses for these two nominal genera has been quoted in paragraph 215 of the present paper, while the passage in which they cited by name the species which they referred to these genera has been quoted in paragraph 81. From the second of these passages it 174 OPINIONS AND DECLARATIONS will be seen that Feletti & Grassi included in their nominal genus Laverania a single species which they cited as Laverania malariae. It is clear from the passages quoted that Feletti & Grassi (while regarding the species Haemamoeba malariae as a new species) did not regard Laverania malariae as a new species and looked upon themselves as doing no more than transferring to the genus Laverania (established by themselves in honour of Laveran) the species of human malaria parasite responsible for the irregular fevers (“‘ delle febbri irregolari”?) which Laveran had been the first to discover and to which in 1881 he had given the name Oscillaria malariae. 223. The nominal genus Laverania Feletti & Grassi, 1889, is thus a monotypical genus with Oscillaria malariae Laveran, 1881 (=the Malignant Tertian Malaria Parasite) as its type species. The generic name Laverania Feletti & Grassi, 1889, is a nomen- clatorially available name in the sense (a) that it is not invalid under Article 34 of the Régles as a junior homynym of any previously published generic name and (b) that it has, as its type species, a nominal species which is not also the type species of a © previously established nominal genus that possesses an available name. 224. We have to note, however, that Oscillaria malariae Laveran, 1881, the type species of Laverania Feletti & Grassi, 1889, is subjectively identified with Plasmodium malariae Mar- chiafava & Celli, 1885, both these nominal species being regarded as representing the Malignant Tertian Malaria Parasite (para- graphs 45 and 77 above respectively). We have further to recall (paragraph 204) that the nominal species Plasmodium malariae Marchiafava & Celli, 1885, is the type species of the nominal genus Plasmodium Marchiafava & Celli, 1885. Thus, through the subjective identification of the species represented by the nominal species which are respectively the type species of these two genera, the nominal genera Laverania Feletti & Grassi and Plasmodium Marchiafava & Celli are subjectively identified with one another. Accordingly, though an available name, the name Laverania Feletti & Grassi, 1889, is a junior subjective synonym of the name Plasmodium Marchiafava & Celli, 1885. OPINION 283 175 (7) The generic name “‘ Ematozoo ” Antolisei and Angelini, 1890 225. As explained in paragraphs 104-108 above the generic name Ematozoo was published by Antolisei & Angelini in two papers each of which appeared in 1890. The exact date of publication (8th March 1890) is known for one of the papers in which the name Ematozoo appeared (i.e. the paper in the Riforma medica) but, as regards the other paper nothing is known regarding the date of its publication except that it was some time in 1890. In these circumstances, it is necessary to treat the name Ematozoo as having been first published in the paper which appeared in the Riforma medica. The reference to the original publication of the name Ematozoo Antolisei & Angelini, is, therefore, 8th March 1890, Riforma medica 6 (56) : 334. (This point is of theoretical interest only, for this generic name was published in exactly the same way in each of the two papers concerned.) 226. We have pointed out (paragraph 107 above) that the generic name Ematozoo was originally published with a small initial letter (as “‘ ematozoo’’) and in the same Section we have given our reasons (paragraphs 110—118) for considering that, although this method of publishing a new generic name was defective, in that it did not comply with the requirements of Articles 3 and 8 of the Rég/es, the infringement involved, though calling for automatic correction by later authors, is not such as to deprive either the generic name Ematozoo or the specific trivial name falciforme, published at the same time in the com- bination ematozoo falciforme, of rights under the Law of Priority (Article 25) as from the date in 1890, on which was published the paper by Antolisei & Angelini containing these names in the forms *“ematozoo”’ and “ ematozoo falciforme’’ respectively. 227. In each of the papers referred to above, the name Ematozoo Antolisei & Angelini, 1890, was published as the name of a monotypical genus with the new nominal species, Ematozoo falciforme Antolisei & Angelini, 1890, as type species. We have shown in paragraph 104 above that this name was applied in these papers to the parasite of semilunar form found in cases of the “‘ aestivo-autumnal Roman fever.”” In other words, this name applies to a form of the Malignant Tertian Malaria Parasite. 176 OPINIONS AND DECLARATIONS 228. We see, therefore, that Ematozoo falciforme Antolisei & Angelini, 1890, is a junior subjective synonym of Oscillaria malariae Laveran, 1881. At this point we have to recall (para- graph 204) that another junior subjective synonym of Laveran’s species is Plasmodium malariae Marchiafava & Celli, 1885, the type species of Plasmodium Marchiafava & Celli, 1885. Thus, the nominal species which are the respective type species of Ematozoo Antolisei & Angelini, 1890, and Plasmodium Marchia- fava & Celli, 1885, are both subjectively identified (on taxonomic grounds) with the same species. Accordingly the generic name Ematozoo Antolisei & Angelini, 1890, though an available name, is a junior subjective synonym of the name Plasmodium Marchiafava & Celli, 1885. (8—11) The generic names ‘ Cytamoeba,” ‘* Cytosporon,” ** Haemocytosporon ” and ‘* Cytozoon ” published by Danilewsky in 1891 229. In the paper published in 1891 (Ann. Inst. Pasteur 5 (12) : 758—782) discussed in paragraph 140 above in connection with the alleged specific and subspecific trivial name hominis, Danilewsky introduced no less than four new generic names for the human malaria parasites :— (1) Cytamoeba. This name was proposed on page 762 of Danilewsky’s paper as a substitute for the name Haemamoeba (“. . . jai proposé de remplacer la déno- mination du plasmodium malarique de lhomme, Haemamoeba, en celle de Cytamoeba.’’) (2) Cytosporon. Later in the same paragraph on page 762 Danilewsky introduced the generic name Cytosporon for the human malaria parasite (“‘ Aussi, et surtout a cause de la propriété fondamentale du microbe de donner des spores, je l’appellerai Cytosporon malariae.’). In the schematic table which appears on page 780 of Danilew- sky’s paper (and which is reproduced in paragraph 140 of the present paper), the generic name Cytosporon is given in the second column (jointly with the name OPINION 283 177 Cytozoon) as the name for a group which Danilewsky appears to have regarded as being of generic value, for, in addition to Cytosporon, he placed in it the well-known and thoroughly recognised genus Haemamoeba with Cytamoeba cited as a synonym. (3) Haemocytosporon. This name was introduced in a foot- note on page 762 of Danilewsky’s paper with a statement that it was the full form of the name published in the main text on the same page in the abridged form “* Cytosporon ”’ (“ On ne doit voir dans ce nom provisoire i.e. Cytospron (abrége de Haemocytosporon) aucune allusion...”’). (4) Cytozoon. This name appears in the first column of the table on page 780 of Danilewsky’s paper (reproduced in paragraph 140 above) as a generic name applicable to the parasites both of Man (“hominis”) and birds (“avium”), which embraces (in some unexplained manner) all the generic names cited in the second and third columns of the table. 230. The task of interpreting Danilewsky’s use of the foregoing names would present almost insuperable difficulties. For- tunately, however, it is not necessary to make this effort, for (as explained in paragraph 142 above) the International Commission on Zoological Nomenclature has ruled, in Opinion 101, that the Latin “‘ technical designations ” used by Danilewsky in this paper are not in harmony with the Rég/es and accordingly possess no rights under the Law of Priority (Article 25) as from the publica- tion of that paper. 231. The four so-called generic names (Cytamoeba, Cytosporon, Haemocytosporon Cytozoon) published in connection with the human malaria parasites by Danilewsky in 1891 have thus no status whatever in zoological nomenclature. 178 OPINIONS AND DECLARATIONS (12) The generic name ‘‘ Haemamoeba ” as used by Labbé in 1894 232. In the recent paper on the nomenclature of the human malaria parasites, to which reference has already been made, Christophers has included (1945, Rev. Inst. Sabubridas y Enfer- medads trop. 6 (4): 216, 220) on page 216 the generic name ‘* Haemamoeba Labbé, 1894” in a list of the generic names pub- lished for the human malaria parasites. On the same page he stated that the type species of this genus was “ O. malariae Laveran (1881) by designation.” In a note on page 220, he again referred to the genus “‘ Haemamoeba Labbé (1894).” On this occasion he added that this genus “ includes Haemamoeba and Laverania of Feletti & Grassi.’ In the same note he implicitly withdrew the earlier statement (made on page 216) that Oscillaria malariae Laveran, 1881, was the type species of this (alleged) genus, for he stated: “‘ He [i.e. Labbé] recognises only one species of malaria parasite, viz., H. Laverani, with varieties quartana and tertiana.” 233. These entries in the synonymy referred to above are - clearly due to some inadvertence, for (as shown in paragraph 160 above) in the paper in question (‘“ Recherches zoologiques et biologiques surles Parasites endoglobulaires du Sang des Vertebrés,” published in 1894, Arch. Zool. expér. gén. (3) 2 : 170) Labbé definitely attributed the generic name Haemamoeba to Grassi (actually he should have attributed it to Feletti & Grassi) and not to himself. Further, at no point in that paper did Labbé cite the name Oscillaria malariae Laveran, 1881, although at the beginning of the section of his paper dealing with human malaria parasites he referred in detail to Laveran’s discovery of “ the” parasite of human malaria and quoted at length (: 160—161) from the note of 24th December 1880 communicated by Laveran to the “‘ Société médicale des Hépitaux ” (reproduced also in the “Notes a l’Académie de Médicine~’) in which that authority made known the discovery which he had made (paragraphs 34 and 35 above). 234. The position is therefore that Labbé never published the name Haemamoeba as a new generic name of his own: he merely used the generic name Haemamoeba Feletti & Grassi, 1890 (which OPINION 283 179 he incorrectly attributed to Grassi). The alleged generic name ** Haemamoeba Labbé, 1894” is thus seen to be a non-existent name and therefore to have no place in the synonymy of the nominal genera established for the human malaria parasites. (13) The generic name “‘ Haematozoon” Thayer and Hewetson, 1895 235. We have seen (paragraphs 104—105 above) that in a paper entitled “‘The Malarial Fevers of Baltimore’? (1895, Johns Hopkins Hosp. Rep. 5:1—218, 2 pls.) Thayer & Hewetson stated (: 27) that in two papers (to which bibliographical references were given) Antolisei & Angelini had published a new name *““ haematozoon falciforme”’ for the parasite responsible for the *** aestivo-autumnal ’ Roman fever.’’ We have seen also (para- graphs 107—108 above) that in the two papers cited Antolisei & Angelini did publish a new name for this parasite but that the new name so published consisted of the words “ ematozoo falciforme.” It follows, therefore, that it was Thayer & Hewetson themselves who were the first to publish the generic name Haematozoon (printed in their paper “‘ haematozoon ”’ with a small initial letter), which appears to have been a deliberate emendation by them of the generic name Ematozoo Antolisei & Angelini, 1890 (which, as will be seen from the extract quoted in para- graph 107 above, was originally published by those authors as *ematozoo’’ with a small initial letter). 236. In our discussion of the generic name Ematozoo Antolisei & Angelini, 1890 (paragraphs 110—118 above) we have given our grounds for the view that the failure on the part of these authors to write the first letter of the word “ ematozoo’’ with a capital letter, though an infringement of Articles 3 and 8 of the Régles, which calls for automatic correction by later authors, does not constitute an infringement sufficiently serious to deprive the generic name Ematozoo Antolisei & Angelini, 1890, of rights under the Law of Priority. While noting that Thayer & Hewetson in their turn published the generic name Haematozoon without 180 OPINIONS AND DECLARATIONS a capital initial letter, we need not debate whether this omission on their part was sufficiently serious to invalidate this name, for the considerations in the opposite sense advanced in the case of the name Ematozoo Antolisei & Angelini, 1890, apply with equal vigour to the present case. 237. From the way in which Thayer & Hewetson introduced the name Haematozoon it is likely that they did not regard themselves as publishing a new name of their own but rather as correcting a name already published by Antolisei & Angelini, though (as explained in paragraph 235 above) it must be conceded also that, in introducing the spelling “‘ Haematozoon” in place of the spelling “‘ Ematozoo ”’ used by Antolisei & Angelini, Thayer & Hewetson did accept a personal responsibility for the revised spelling “‘ Haematozoon.” 238. As will be seen from paragraph 235 above, the generic name Haematozoon Thayer & Hewetson, 1895, was published with a single included species, Haematozoon falciforme. That species is accordingly the type species of the genus Haematozoon Thayer & Hewetson by monotypy under Rule (c) in Article 30. 239. We have already seen (paragraph 119 above) that the species Ematozoo falciforme Antolisei & Angelini, 1890, is the parasite responsible for the aestivo-autumnal Roman fever, that is, the Malignant Tertian Malaria Parasite, Oscillaria malariae Laveran, 1881. Another nominal species which has also been subjectively identified with that species is Plasmodium malariae Marchiafava & Celli, 1885. The last-named species is the type species of Plasmodium Marchiafava & Celli, 1885. Since Plas- modium Marchiafava & Celli is a nomenclatorially available name, not being invalid as a homonym under Article 34, and the type species of this genus is subjectively identified as conspecific with the type species of Haematozoon Thayer & Hewetson, 1895, we see that Haematozoon Thayer & Hewetson, 1895, is junior subjective synonym of Plasmodium Marchiafava & Celli, 1885. OPINION 283 181 240. Taxonomically, therefore, the name Haematozoon Thayer & Hewetson, 1895, is an unnecessary name which would not be required, even if it were an available name. In fact, however, the name Haematozoon Thayer & Hewetson, 1895, is not an available name, for it is invalid under Article 34 as a junior homonym of the older name Haematozoon Leisering, 1865 (Arch. path. Anat. 33(1): 125) published for a genus in the Class Nematoda. (14) The generic name ‘‘ Haemosporidium ” Lewkowicz, 1897 241. In a paper published in the issue of 6th February 1897 of the “Erste Abteilung” of the Centralblatt fir Bakteriologie, Parasitenkunde u. Infektionskrankheiten, Lewkowicz published a short paper entitled “‘ Ueber den Entwickelungsgang und die Einteilung der Malariaparasiten ’’ (1897, Zb/. Bakt. (Erste Abt.) 21(4) : 129—133), in which he introduced a new classification for the human malaria parasites, all the species of which he placed in a genus to which he applied the new name Haemosporidium (: 132—133). He did not cite, or otherwise refer to, any of the earlier generic names proposed for these parasites nor did he explain why he considered it necessary to reject those names in favour of the new name Haemosporidium. It may be inferred, however, that he made this change on the ground that his new name was more appropriate than any of those previously proposed, for, when discussing (: 130) the young spores (“‘ Die jungen Sporidien ’’) of the Quartan and Benign Tertian Malaria Parasites (the two species placed by him (: 132) in Group I (“ I. Gruppe ’’) of the genus Haemosporidium), Lewkowicz added the following footnote with reference to the word “‘ Sporidien ” :—‘‘ Dieser von Danilewsky vorgeschlagene Name scheint mir fiir die zoologische Bezeichnung der Malariaparasiten der entsprechendste.”’ 242. Article 32 of the Régles provides that a generic name, once published, cannot be rejected on the ground of inappro- priateness. Accordingly, if, in fact, this was the ground on which Lewkowicz rejected the earlier generic names published for the human malaria parasites and substituted the new name Haemo- sporidium in their place, his action was invalid. 182 OPINIONS AND DECLARATIONS 243. The name Haemosporidium Lewkowicz, 1897, is an available name in the sense that it is not a junior homonym of any previously published generic name in the Animal Kingdom. Its nomenclatorial status will, therefore, turn entirely upon the identity of the species selected as its type species. 244. Lewkowicz did not himself designate a type species for this genus, nor was its type species automatically determined under Rules (b), (c), (d) or (f) in Article 30 of the Régles. Accord- ingly, Rule (g) in that Article applies to this name and in consequence the type species of this genus is whatever nominal species is first selected as such by a subsequent author. So far as I have been able to ascertain, no such type selection has ever been made for this genus. 245. Lewkowicz, as has already been explained (paragraph 169), placed in his genus Haemosporidium all the species of human malaria parasite that he recognised, but did not cite those species under binominal names, contenting himself with applying to each a simple technical term (in the form of a Latin adjective), indi- cating the period of development of the fever arising from the parasite in question. Accordingly, the generic name Haemo- sporidium Lewkowicz, being the name of a nominal genus established without any included species cited under a binominal name, falls to have its type species determined under the provisions of the Commission’s Opinion 46 (1912, Smithson. Publ. 2060 : 104—107). The Ruling given in this Opinion is obscure and unsatisfactory, and it is much to be hoped that at its next Session the International Commission on Zoological Nomenclature will submit to, and secure the approval of, the International Congress of Zoology proposals for the clarification of the Régles in regard to the type species of nominal genera established without cited nominal species . It is fortunately not necessary to examine in detail the Ruling given in the foregoing Opinion, for the one provision in it which is clear happens by a lucky chance to be applicable in the present case. The portion of the Ruling given in the so-called ““ Summary ”’ with which we are concerned reads as follows: “.. . if several species are referred to but not men- tioned by name, one of these must be taken as the type.” Later OPINION 283 183 the Ruling turns to the case where a nominal genus is established with a diagnosis but with no indication as to the species which should be treated as being included species, and on this part of the subject the Ruling given was that “ the first species published in connection with the genus [i.e. the first validly established nominal species cited as belonging to the nominal genus con- cerned] (as Ac/astus rufipes Ashmead, 1902) becomes ipso facto the type.” In the present case, we must note (1) that we know what are the taxonomic species regarded by Lewkowicz as belonging to his genus Haemosporidium, and (2) that so far no author has formally cited under a binominal name any of the included species as being a species referable to this genus. It follows, therefore, that it is still open to any author formally to determine the type species of the genus MHaemosporidium Lewkowicz, 1897, under the procedure sear by the Ruling given in Opinion 46. 246. Accordingly, in order to secure a determinate content for the nominal genus Haemosporidium Lewkowicz and therefore a definite position for the generic name Haemosporidium Lewkowicz in the synonymy of the names of the nominal genera established for the human malaria parasites, I now propose to take the formal action required under Opinion 46 to provide the foregoing nominal genus with a type species. To this end, acting in accord- ance with the procedure prescribed in the foregoing Opinion, I hereby formally (1) “‘ recognise ’’ the Quartan Malaria Parasite as one of the species included by Lewkowicz in the nominal genus Haemosporidium established by him in 1897, (2) cite the name of the foregoing species (Haemamoeba malariae Feletti & Grassi, 1889), in combination with the generic name Haemosporidium Lewkowicz 1897, thus forming the combination Haemosporidium malariae (Feletti & Grassi 1889), and (3) designate (and select) the foregoing nominal species to be the type species of the nominal genus Haemosporidium Lewkowicz, 1897. 23 The Ruling given in Opinion 46 was clarified by the International Commission on Zoological Nomenclature at its Session held at Paris in 1948 (1950, Bull. zool. Nomencl. 4 : 160, 346) and the Ruling so clarified was incorporated in the Rég/es. Opinion 46 itself was thereupon repealed for interpretative purposes, that is, for every purpose except that of historical record (1950, ibid. 4 : 165—166). The action here taken in relation to the name Haemosporidium Lewkowicz is in harmony with the provisions adopted by the Paris Congress. 184 OPINIONS AND DECLARATIONS 247. The nominal species Haemamoeba malariae Feletti & Grassi, 1889, is also the type species of the nominal species Haemamoeba Feletti & Grassi, 1889 (see paragraph 220 above). Thus, the same species is the type species both of Haemamoeba Feletti & Grassi, 1889, and of Haemosporidium Lewkowicz, 1897. As the name Haemamoeba Feletti & Grassi is a nomenclatorially available name (and the valid generic name for Haemamoeba malariae Feletti & Grassi), the name Haemosporidium Lewkowicz, 1897, is invalid, being an junior objective synonym of Haema- moeba Feletti & Grassi, 1889. OPINION 283 185 PART 3 SYNONYMY OF THE SPECIFIC TRIVIAL NAMES BESTOWED (OR ALLEGED TO HAVE BEEN BESTOWED) UPON THE HUMAN MALARIA PARASITES IN THE PERIOD 1881- 1897 AND OF THE NAMES OF THE NOMINAL GENERA ESTABLISHED (OR ALLEGED TO HAVE BEEN ESTABLISHED) FOR THOSE SPECIES IN THE SAME PERIOD 248. In Part 1 of the present paper we have firmly established the identity of the taxonomic species represented by each of the nominal species established, or alleged to have been established, for species of human malaria parasite during the period 1881-1897. In addition, we have determined for each of the specific names concerned whether it possesses status under the Law of Priority and whether it is an objectively available name. Similarly, in Part 2, we have determined for each of the nominal genera established, or alleged to have been established, for species of human malaria parasite whether it is a name which possesses status under the Law of Priority. Further, by determining the nominal species which, under the Régles, is the type species of each of the validly established nominal genera, we have firmly established the content of those genera for nomericlatorial pur- poses. We are therefore now in a position to construct synony- mies of the names bestowed upon each of the species of human malaria parasite and thus to determine for each of those species what, under the Régles, is its oldest available specific trivial name. It is possible also now to arrange in order of priority the names bestowed upon nominal genera established for one or other or the species of human malaria parasite. 249. As will be seen, the synonymies constructed in the fore- going manner are necessarily very complicated, and it has been considered that it would facilitate their presentation and make them more readily understandable, if for the present purposes the bibliographical references were to be omitted. In order, however, that these may be readily available I give in Section (1) of the present Part an alphabetical list of all the names, both generic and specific, which appear in the present paper, together with 186 OPINIONS AND DECLARATIONS the bibliographical references for those names. Next are given the specific synonymies (Section (2)) and then a list of generic names with the type species of the nominal genera so named (Section (3)). In Section (4) I show, on the basis of the particulars given in Sections (2) and (3), what, under the Régles, is the valid name (binominal combination) for each of the human malaria parasites, (a) on the basis of the view that all three species are congeneric with one another (the view most commonly held by malariologists), and (b) on the basis of the view that two genera should be recognised. In Section (5), I compare the names applicable to the human malaria parasites under the Régles with the names universally applied to those species by malariologists. As will be seen, the two sets of names differ in almost every possible respect. (1) Original references for the names cited in the present paper 250. The following are the original references for the names bestowed or alleged to have been bestowed upon genera and species of human malaria parasites discussed in the present paper :— Cytamoeba Danilewsky, 1891, Ann. Inst. Pasteur 5(12) 762 Cytosporon Danilewsky, 1891, Ann. Inst. Pasteur 5(12) : 762 - Cytozoon Danilewsky, 1891, Ann. Inst. Pasteur 5(12) : 780 Ematozoo (correction of ematozoo) Antolisei & Angelini, 1890, Riforma medica 6(56) : 334 falciforme, Ematozoo (correction of ematozoo), Antolisei & Angelini, 1890, Riforma medica 6(56) : 334 falciforme, Haematozoon, Thayer & Hewetson, 1895, Johns Hopkins Hosp. Reps. 5 : 27 (usually cited by the title of the paper which constitutes this voulme, namely The Malarial Fevers of Baltimore) falciparum, Haematozoon, Welch, 1897, in Loomis’s Syst. pract. Med. 1 : 36 Haemamoeba Feletti & Grassi, Dec. 1889, Sui parasiti della malaria (a pre-print of a paper published in Januazy 1890 (Riforma medica 6(11) : 63)) OPINION 283 | 187 Haematomonas Mitrophanow, 1883, Biol. Zbl. 3: 41 Haematomonas Osler, 1886, Philad. med. Times and 1887, Brit. med. J. 1887 (Vol. 1) : 556 Hdmatophyllum Metschnikoff, 1887, Russk. Med. 12 : 207 Haematozoon (correction of haematozoon) Thayer & Hewetson, 1895, Johns Hopkins Hosp. Reps. 5 : 27 (usually cited by the title of the paper which constitutes this volume, namely The Malarial Fevers of Baltimore) Haemocytsporon Danilewsky, 1891, Ann. Inst. Pasteur 5(12) : 762 nota Haemosporidium Lewkowicz, 1897, Zbl. Bakt. (Erste Abt.) 21(4) : 132—133 hominis, Laverania, Danilewsky, 1891, Ann. Inst. Pasteur 5(12) :780 immaculata, Haemamoeba, Grassi, 1891, Zb/. Bakt. 10 : 517 irregularis, Plasmodium malariae, Kruse, 1892, Hyg. Rdsch. 2 : 467 irregularis, Haemamoeba febris, Sakharov, 1892 \a cherionym) laverani, Haemamoeba, Labbé, 1894, Arch. Zool. expér. gén. (3)2 : 170 Laverania Feletti & Grassi, Dec. 1889, Sui parasiti della malaria (a pre-print of a paper published in January 1890 (Riforma medica 6(11) : 63)) malariae, Haemamoeba, Feletti & Grassi, Dec. 1889, Sui parasiti della malaria (a pre-print of a paper published in January 1890 (Riforma medica 6(11) : 64)) malariae, Laverania, Feletti & Grassi, Dec. 1889, Sui parasiti della malaria (a pre-print of a paper published in January 1890 (Riforma medica 6(11) : 64)) malariae, Oscillaria, Laveran, 1881, Nature parasit. Accidents Impaludisme : 87 malariae, Plasmodium, Marchiafava & Celli, 1885, Fortschr. Med. 3(24) : 791 Oscillaria Laveran, 1881, Nature parasit. Accidents Impaludisme Bi Oscillaria Schrank, 1823, Nova Acta. Acad. Caes. Leopold. Carol. 11 (No. 2) : 533 Plasmodium Marchiafava & Celli, 1885, Fortschr. Med. 3(24) : 791 praecox, Haemamoeba, Grassi & Feletti, May 1890, Arch. ital. Biol. 13(2) : 300 188 OPINIONS AND DECLARATIONS gquartana, Haemamoeba laverani var., Labbé, 1894, Arch. Zool. exper. gén. (3)2: 170 guartanae, Haemosporidium, Lewkowicz, 1897, Zbl. Bakt. (Erste Abt.) 21(4) : 132 quartanae, Plasmodium malariae M. & C. Varieta A, Celli & Sanfelice, 1891, Ann. Ist. Igiene sper. Univ. Roma (n.s.) 1: 61 quartanae, Plasmodium malariae, Kruse, 1892, Hyg. Rdschr. 2 : 466 quotidianae, Plasmodium malariae M. & C., Varieta C., Celli & Sanfelice, 1891, Ann. Ist. Igiene sper. Univ. Roma (n.s.) 1: 61 sedecimanae, Haemosporidium, Lewkowicz, 1897, Zbl. Bakt. (Erste Abt.) 21(4) : 133 tertiana, Haemamoeba laverani var., Labbé, 1894, Arch. Zool. exper. gén. (3)2: 170 tertianae, Haemosporidium, Lewkowicz, 1897, Zbl. Bakt. (Erste Abt.) 21(4) : 132 tertianae, Plasmodium malariae M. & C., Varieta B., Celli & Sanfelice, 1891, Ann. Ist. Igiene sper. Univ. Roma (n.s.) 1: 61 tertianae, Plasmodium malariae, Kruse, 1892, Hyg. Rdschr. 2 : 466 tropica, Plasmodium, Koch, 1899 (a cheironym) undecimanae, Haemosporidium, Lewkowicz, 1897, Zbl. Bakt. (Erste Abt.) 21(4) : 133 vigesimo-tertianae, Haemosporidium, Lewkowicz, 1897, Zbl. Bakt. (Erste Abt.) 21(4) : 133 vivax, Haemamoeba, Grassi & Feletti, May 1890, Arch. ital. Biol. 13(2) : 300 (2) Synonymy of the specific trivial names bestowed (or alleged to have been bestowed) upon the human malaria parasites in the period 1881—1897 251. In the present Section I give a synonymy for each of the three species of human malaria parasite with which we are concerned. To the names shown in these synonymies must be added one trivial name, hominis Danilewsky, 1891, as published in the combination Laverania (paragraphs 140—143), which probably applies to the Malignant Tertian Malaria Parasite but is at present indeterminate and can safely be left so, in view of the fact that the International Commission on Zoological Nomen- clature has ruled (Opinion 101) that the paper in which it was published is not available for nomenclatorial purposes. OPINION 283 189 (a) The Malignant Tertian Malaria Parasite 252. The following is a synonymy of the names bestowed (or alleged to have been bestowed) upon the Malignant Tertian Malaria Parasite. The paragraph references cited are to the paragraphs in the present paper in which the names in question are discussed in detail :— (i) Validly published names for the Malignant Tertian Malaria Parasite Oscillaria malariae Laveran, 1881 (paragraphs 34—45) Plasmodium malariae Marchiafava & Celli, 1885 (paragraphs 53—77) Laverania malariae Feletti & Grassi, December 1889 (not a new name but a usage of the trivial name malariae Laveran, 1881) (paragraphs 91—93) Ematozoo (correction of ematozoo) falciforme Antolisei & Angelini, 1890 (paragraphs 102—119) Haemamoeba laverani Labbé, 1894 (paragraphs 158—166) Haematozoon (correction of haematozoon) falciforme Thayer & Hewetson, 1895 (paragraphs 167—168) Haematozoon falciparum Welch, 1897 (paragraphs 175—177) (ii) Validly published names which have been applied to the Malignant Tertian Malaria Parasite but erroneously because the nominal species so named were established for parasites of birds Haemamoeba praecox Grassi & Feletti, May 1890 (paragraphs 96—101) Haemamoeba immaculata Grassi, 1891 (paragraphs 120—127) 190 OPINIONS AND DECLARATIONS (iii) Technical terms published in connection with the Malignant Tertian Malaria Parasite and erroneously treated by later authors as names bestowed upon that species Plasmodium malariae M. & C., Varieta C., quotidianae Celli & Sanfelice, 1891 (paragraphs 128—139) Plasmodium malariae irregularis Kruse, 1892 (paragraphs 144—150) Haemosporidium sedecimanae Lewkowicz, 1897 (paragraphs 169—174) Haemosporidium undecimanae Lewkowicz, 1897 (paragraphs 169—174) Haemosporidium vigesimo-tertianae Lewkowicz, 1897 (paragraphs 169—174) (iv) Reputed but non-existent names Haemamoeba febris irregularis Sakharov, 1892 (paragraphs 151—157) Plasmodium tropica Koch, 1899 (paragraphs 178—185) (b) The Quartan Malaria Parasite 253. The following is a synonymy of the names bestowed (or alleged to have been bestowed) upon the Quartan Malaria Parasite :— (1) Validly published names for the Quartan Malaria Parasite Haemamoeba laverani var. quartana Labbé, 1894 (paragraphs 158—166) OPINION 283 191 Haemamoeba malariae Feletti & Grassi, 1889 (invalid because a rejected junior secondary homonym of the name malariae Laveran, 1881, consequent upon the two species so named having been united in the same genus, Plasmodium Marchiafava & Celli, 1885) (paragraphs 78—90) (ii) Technical terms published in connection with the Quartan _ Malaria Parasite and erroneously treated by later authors as : names bestowed upon that species Plasmodium malariae M. & C., Varieta A., quartanae Celli & _ Sanfelice, 1891 (paragraphs 128—139) Plasmodium malariae quartanae Kruse, 1892 (paragraphs 144—150) Haemosporidium tertianae Lewkowicz, 1897 (paragraphs 169—174) (c) The Benign Tertian Malaria Parasite _ 254. The following is a synonymy of the names bestowed (or alleged to have been bestowed)upon the Benign Tertian Malaria Parasite :— (i) Validly published names for the Benign Tertian Malaria Parasite Haemamoeba vivax Grassi & Feletti, 1890 (paragraphs 94—95) Haemamoeba laverani var. tertiana Labbé, 1894 (paragraphs 158—166) (ii) Technical terms published in connection with the Benign Tertian Malaria Parasite and erroneously treated by later authors as names bestowed upon that species Plasmodium malariae M. & C., Varieta B. tertianae Celli & Sanfelice, 1891 (paragraphs 128—139) Plasmodium malariae tertianae Kruse, 1892 (paragraphs 144—150) Haemosporidium tertianae Lewkowicz, 1897 (paragraphs 169—174) 192 OPINIONS AND DECLARATIONS (3) Names of the nominal genera established (or alleged to have been established) for the human malaria parasites in the period 1881—1897 255. The time has come now to bring together the names of all the nominal genera established (or alleged to have been established) for human malaria parasites in the period 1881—1897. It would have been possible at this point to arrange these names in synony- mies based upon current taxonomic judgments as to the number of genera which should be recognised, but this course would have presented certain inconveniences in view of the fact that there is not unanimity among malariologists on the question whether two genera or one genus only should be recognised on taxonomic grounds. In these circumstances it has been judged preferable at this stage to set out the names concerned in a single list arranged by reference to the dates on which the names were published. Altogether, there are sixteen names to be considered. Of these, seven are validly published names and each is applicable to a nominal genus having one or other of the human malaria parasites as its type species. Two are validly published names which have however been applied to the human malaria parasites in error, the - nominal genera so named being based upon species of other — groups. Next, there are four names (all published by Danilewsky in 1891), the content of which is indeterminate, no type species having ever been designated or selected, but this is of no practical consequence in view of the fact that the International Commission on Zoological Nomenclature has ruled (Opinion 101) that the paper in which these names were published is not available for nomenclatorial purposes. Finally there are three reputed but non-existent generic names to which it has been necessary to give consideration. Generic names given or applied to human malaria parasites (a) Validly published generic names published for nominal genera having species of human malaria parasite as type species Plasmodium Marchiafava & Celli, 1885 (type species, by mono- typy: Plasmodium malariae Marchiafava & Celli, 1885 (=The Malignant Tertian Malaria Parasite), a junior subjective synonym of Oscillaria malariae Laveran, 1881) (an available generic name) (paragraphs 199—204) OPINION 283 193 _ Hématophyllum Metschnikoff, 188724 (type species, by monotypy: Plasmodium malariae Marchiafava & Celli, 1885) (invalid, because a junior objective synonym of the name Plasmodium Marchiafava & Celli, 1885) (paragraphs 209—213) ie Feletti & Grassi, 1889 (type species, by monotypy: Haemamoeba malariae Feletti & Grassi, 1889 (=the Quartan Malaria Parasite)) (an available oe name) (paragraphs 214—220) Laverania Feletti & Grassi, 1889 (type species, by monotypy: Oscillaria malariae Laveran, 1881) (an available name, but a junior subjective synonym of the name Plasmodium Marchiafava & Celli, 1885) (Under a possible but much less probable interpretation, the type species of Laverania Feletti & Grassi, 1889, is, by monotypy, the nominal species named Laverania malariae Feletti & Grassi, 1889. If such a name had been published by these authors, it would be a junior subjective synonym of Oscillaria malariae Laveran, 1881, and in this case also the name Laverania Feletti & Grassi, 1889, would be a junior subjective synonym of Plasmodium Marchiafava & Celli, 1885.) (paragraphs 221—224) Ematozoo (correction of ematozoo) Antolisei & Angelini, 1890 (type species, by monotypy: Ematozoo (correction of ematozoo) falciforme Antolisei & Angelini, 1890, a junior subjective. synonym of Oscillaria malariae Laveran, 1881) (an available name but a junior subjective synonym of the name Plasmodium Marchiafava & Celli, 1885) (paragraphs 225—228) Haematozoon (correction of haematozoon) Thayer & Hewetson, 1895 (type species, by monotypy: Haematozoon falciforme Thayer & Hewetson, 1895, a junior subjective synonym of Oscillaria malariae Laveran, 1881) (invalid, because a junior homonym of Haematozoon Leisering, 1865 (Class Nematoda) ; in addition through the foregoing subjective identification of 24 This name was originally published as Hadmatophyllum, but under a decision taken by the Fourteenth International Congress of Zoology, Copenhagen, 1953, when revising Article 20, a name so published is in future to be written out as Haematophyllum. See Footnote 22. G 194 OPINIONS AND DECLARATIONS the type species of the above nominal genus, the name Haemato- zoon Thayer & Hewetson, 1895, is a junior subjective synonym of the name Plasmodium Marchiafava & Celli, 1885) (para- graphs 235—240) Haemosporidium Lewkowicz, 1897 (type species, by selection, under Opinion 46 by Hemming in the present paper: Haema- moeba malariae Feletti & Grassi, 1889) (invalid, because a junior objective synonym of the name Haemamoeba Feletti & Grassi, 1889) (paragraphs 241—247) (b) Validly published generic names which have been applied to the human malaria parasites but incorrectly because the nominal genera so named were established for species of other groups Oscillaria Schrank, 1823 (the name of a nominal genus of Algae) (paragraphs 187—198) | Haematomonas Mitrophanow, 1883 (the name of a nominal genus established for two flagellates found in the blood of species — of fish) (paragraphs 205—208) (c) Generic names of indeterminate content published in a paper which the International Commission on Zoological Nomen- clature has ruled is not available for nomenclatorial purposes Cytamoeba Danilewsky, 1891 (paragraphs 229—231) Cytosporon Danilewsky, 1891 (paragraphs 229—231) Cytozoon Danilewsky, 1891 (paragraphs 229—231) Haemocytozoon Danilewsky, 1891 (paragraphs 229—231) (d) Reputed but non-existant generic names Oscillaria Laveran, 1881 (paragraphs 187—198) Haematomonas Osler, 1886 (paragraphs 205—208) Haemamoeba Labbé, 1894 (paragraphs 232—234) OPINION 283 195 (4) The correct names for the Human Malaria Parasites under a strict application of the “ Régles ” _ 256. Having now compiled a synonymy, both generic and specific, for each of the human malaria parasites, we are at last in a position to establish what is the correct name under the Régles for each of these species. In the view of the majority of malariologists, all three of the species concerned are congeneric with one another, but in the view of some specialists two genera should be recognised, the Malignant Tertian Malaria Parasite being placed in one genus and the Quartan Malaria Parasite and the Benign Tertian Malaria Parasite in another genus. The oldest otherwise available trivial name (malariae Feletti & Grassi, 1889) for one of these species (the Quartan Malaria Parasite) is the same as the oldest available name (malariae Laveran, 1881) for another of these species (the Malignant Tertian Malaria Parasite), and, as these species have (by many workers) been united on taxonomic grounds in a single genus, the later-published of these names cannot be used even by those malariologists who regard these two species as being congeneric with one another; for at the time that the two nominal species so named were united in a single genus, the later-published name became a junior secondary homonym of the earlier-published name, and, under Articles 35 and 36, a name rejected and replaced as a junior secondary homonym can never be used again.?° Thus, the specific trivial names of the human malaria parasites are not affected by the differing taxonomic judgments of malariologists on the question whether the three species concerned should be placed in the same genus or whether one of them should be placed in a second genus, and it is only the generic names that vary according to which view is held on the question of generic relationship. 257. With the foregoing explanation, we may now present the following statement showing what under the Régles is the correct name (binominal combination of a generic name and a specific 25 The point here raised was still a matter of discussion when this passage was written, but by a decision by the Thirteenth International Congress of Zoology, Paris, 1948, the point at issue was settled in the sense indicated in this passage. See Footnote 7. 196 OPINIONS AND DECLARATIONS trivial name) for each of the three species of human malaria parasite with which we are concerned (a) for those specialists who hold on taxonomic grounds that all three species are congeneric with one another and (b) for those specialists who regard the Quartan Malaria Parasite and the Benign Tertian Malaria Parasite as congeneric with one another but consider that the Malignant Tertian Malaria Parasite is referable to a separate genus :— TABLE 1 The correct names for the human malaria parasites according to whether two genera or one genus only are recognised on taxonomic grounds Name of species Name of species Vernacular name specified in Col. (1) specified in Col. (1) of human malaria if only one genus if two genera are parasite is recognised on recognised on taxonomic grounds taxonomic grounds (1) (2) (3) Malignant Tertian | Plasmodium malariae Plasmodium malariae Malaria Parasite (Laveran, 1881) (Laveran, 1881) Quartan Malaria | Plasmodium quartanum | Haemamoeba quart- Parasite (Labbé, 1894) anum, Labbé, 1894 Benign Tertian Plasmodium vivax Haemamoeba vivax Malaria Parasite (Grassi & Feletti, Grassi & Feletti, 1890) 1890 (5) Comparison of the names applicable to the human malaria parasites under a strict application of the “‘ Réegles ” with the names universally used for those species 258. In the immediately preceding paragraph we have shown in tabular form what are the correct names for the three human malaria parasites, (a) if only one genus is recognised on taxonomic grounds and (b) if two genera are so recognised. In the following table (Table 2) the names so established are compared with the OPINION 283 197 names universally used for these species on the basis of the Ruling given (though invalidly) by the International Commission on Zoological Nomenclature when in its Opinion 104 it placed the generic names concerned on the Official List of Generic Names in Zoology. This comparison is made first to show the position for these specialists who take the taxonomic view that all three species are congeneric with one another and, second, for those specialists who consider that two genera are involved. TABLE 2 Comparison of the names applicable to the human malaria parasites under a strict application of the ‘“‘ Régles ” with the names now universally used for those species Section A. Position for those specialists who consider that all the human malaria parasites are congeneric with one another Scientific name Vernacular name Correct scientific universally in use of human malaria name under the (also the name parasite ** Régles ”’ recognised in ‘* Opinion ”’ 104) (1) Malignant Ter- | Plasmodium malariae Plasmodium tian Malaria (Laveran, 1881) falciparum (Welch, 1897) Parasite (2) Quartan Malaria | Plasmodium quartanum | Plasmodium malariae Parasite (Labbé, 1894) (Feletti & Grassi, 188976) (3) Benign Tertian | Plasmodium vivax Plasmodium vivax -Malaria Para- (Grassi & Feletti, (Grassi & Feletti, site 1890) 1890) _ °° This name is commonly but incorrectly treated as having first been published in 1890. Further, the names of the authors are normally cited incorrectly in the reverse order as “‘ Grassi & Feletti ’’ (see paragraph 47). 198 OPINIONS AND DECLARATIONS Section B Position for those specialists who consider that two genera are involved, namely, one for the Malignant Tertian Malaria Parasite and one for the Quartan Malaria Parasite and the Benign Tertian Vernacular name of human malaria parasite (1) Malignant Ter- tian Malaria | Parasite (2) Quartan Malaria Parasite 3) Benign Tertian Malaria Para- site 27 See Footnote 26. Malaria Parasite Scientific name universally in use (also the name recognised in “Opinion ”’ 104) Correct scientific name under the ** Régles ” Plasmodium malariae Laverania falciparum (Laveran, 1881) (Welch, 1897) Haemamoeba quartanum| Plasmodium malariae Labbé, 1894 (Feletti & Grassi, 188927) Haemamoeba vivax Grassi & Feletti, 1890 Plasmodium vivax (Grassi & Feletti, 1890) | | OPINION 283 199 PART 4 PROPOSAL FOR THE STABILISATION OF THE NAMES FOR THE HUMAN MALARIA PARASITES ON THE BASIS OF CURRENT USAGE 259. The nomenclature, both generic and specific, for the human malaria parasites adopted by the International Commission on Zoological Nomenclature in its Opinion 104 is the nomenclature now almost universally accepted by malariologists and has been in general use for nearly fifty years. As can however be seen from the table (Table 2) given in paragraph 258, in which the conclusions reached in the earlier part of the present paper have been summarised, the currently adopted nomenclature is incorrect in almost every particular. 260. It is obvious that any material change in the names for the human malaria parasites arising from purely nomenclatorial causes (that is, from the strict application of the Régles) would cause the greatest confusion in the literature of malariology and would serve no useful purpose whatever. There is therefore the strongest possible case for the use by the International Commission on Zoological Nomenclature of its Plenary Powers for the purpose of giving valid force to existing nomenclatorial practice and, incidentally, for validating the erroneous entries made by the Commission itself in its Opinion 104. We have therefore to consider carefully what is the action required in order to secure the desired end. As in the earlier part of the present paper, it will be convenient to consider this problem, first, in relation to the specific trivial names for the human malaria parasites and, second, in relation to the generic names bestowed upon those species. (1) The specific trivial name for the Quartan Malaria Parasite 261. The Quartan Malaria Parasite is universally known by the specific trivial name malariae Feletti & Grassi, 1889 (usually, though incorrectly cited as ‘“‘ Grassi & Feletti, 1890 ’’) as published in the combination Haemamoeba malariae. If the greatest confusion is to be avoided, it is essential that means should be found for retaining this name for this species. In order to 200 OPINIONS AND DECLARATIONS validate the use of this trivial name for the Quartan Malaria Parasite, it will, therefore, be necessary for the International Commission to use its Plenary Powers for the purpose of sup- pressing the name malariae Laveran, 1881, as published in the combination Oscillaria malariae, for the purposes both of the Law of Priority and of the Law of Homonymy. Further, it will be necessary for the Commission to use its Plenary Powers for the purpose of suppressing to a like extent the trivial name malariae Marchiafava & Celli, 1885, as published in the combina- tion Plasmodium malariae, a name bestowed independently by these authors upon the Malignant Tertian Malaria Parasite. Once these two names have been suppressed in this way, all usage of the trivial name malariae in the genus Plasmodium Marchiafava & Celli, 1885, prior to the publication of the name malariae Feletti & Grassi, 1889, for the Quartan Malaria Parasite will have been removed for nomenclatorial purposes. The latter name will, therefore, no longer be a junior secondary homonym of some earlier name malariae in the genus Plasmodium. At present, the name malariae Feletti & Grassi, 1889, as a rejected junior secondary homonym is, under Article 35 and 36, a permanently rejected name which cannot be brought into use again. This — difficulty can however be overcome by the use by the Commission of its Plenary Powers expressly to validate this name. By these measures—and by these measures alone—it will be possible for the International Commission to provide a valid basis for the continued use of the specific trivial name malariae Feletti & Grassi, 1889, as published in the combination Haemamoeba malariae, for the Quartan Malaria Parasite. (2) The specific trivial name for the Malignant Tertian Malaria Parasite 262. The trivial name falciparum Welch, 1897, as published in the combination Haematozoon falciparum, is the name now in general use for the Malignant Tertian Malaria Parasite. That name is, however, a long way down the list of names published for nominal species which are subjectively identified as represent- ing the foregoing parasite, and these names will all have to be suppressed by the International Commission on Zoological Nomenclature under its Plenary Powers before a valid status of OPINION 283 201 availability can be provided for the name falciparum Welch. In addition, there are two names which have priority over falciparum Welch which have been applied in the past to the Malignant Tertian Malaria Parasite but which, in my opinion, were bestowed by their respective authors not upon the foregoing species but upon parasites of birds. In order to provide a secure foundation for the name falciparum Welch, it will be necessary for the Inter- national Commission to take appropriate steps to prevent claims from being advanced on behalf of these names as against the name falciparum Welch. Finally, there is a name published before falciparum Welch which, in my opinion, is no more than a usage of a previously published name but which it has been claimed in the past as having been published as a new name for the Malignant Tertian Malaria Parasite. In this case also, action will need to be taken by the Commission to prevent any possible challenge from being advanced against the name falciparum Welch. 263. The five trivial names for the Malignant Tertian Malaria Parasite which have priority over the name falciparum Welch are the following :— malariae Laveran, 1881, as published in the combination Oscillaria malariae malariae Marchiafava & Celli, 1885, as published in the combina- tion Plasmodium malariae falciforme Antolisei & Angelini, 1890, as published in the com- bination Ematozoo (correction of ematozoo) falciforme laverani Labbé, 1894, as published in the combination Haema- moeba laverani falciforme Thayer & Hewetson, 1895, as published in the com- bination Haematozoon (correction of haematozoon) falciforme 264. Of the foregoing names, I have already recommended the suppression of malariae Laveran, 1881 (paragraph 261), and of malariae Marchiafava & Celli, 1885, as an essential part of the action needed for the purpose of validating the name malariae Feletti & Grassi, 1889, as published in the combination Haema- moeba malariae, for the Quartan Malaria Parasite. Here it is 202 OPINIONS AND DECLARATIONS necessary only to note that the suppression of these names forms an essential part also of any plan for validating the name falciparum Welch for the Malignant Tertian Malaria Parasite. A further part of any such plan will necessarily be the use by the International Commission of its Plenary Powers to suppress for the purposes of the Law of Priority but not for those of the Law of Homonymy the three other trivial names enumerated in paragraph 263 above, that is :(a) falciforme Antolisei & Angelini 1890, as published in the combination Ematozoo (correction of ematozoo) falciforme; (b) laverani Labbé, 1894, as published in the combination Haemamoeba laverani; (c) falciforme Thayer & Hewetson, 1895, as published in the combination Haematozoon (correction of haematozoon) falciforme. 265. Next, it will be necessary for the Commission to take steps to eliminate from consideration as possible names for the Malignant Tertian Malaria Parasite the following names which it has been claimed were published, in part, for that species : (a) praecox Grassi & Feletti, 1890, as published in the combination Haemamoeba praecox; (b) immaculata Grassi, 1891, as published in the combination Haemamoeba immaculata. The circumstances in which these names were published has been fully described in Part 1 of the present paper (praecox, paragraphs 96—101; immaculata, paragraphs 120—127). In the light of the evidence so brought forward it will be necessary for the International Commission to include in its decision on the names of the human malaria parasites a Ruling that these names apply exclusively to the avian parasites so named by the authors concerned. 266. Finally, it is desirable that, as part of any plan for validat- ing the name falciparum Welch, 1897, for the Malignant Tertian Malaria Parasite, the Commission should use its Plenary Powers to remove any possible doubt regarding the status of the name malariae as used in combination with the generic name Laverania by Feletti & Grassi in 1889 for the foregoing species, in so far as that name was on that occasion used by the above authors as a new name; for, if no such action were to be taken, this name, if published as a new name, would, on the suppression of the names malariae Laveran, 1881, as published in the combination Oscillaria OPINION 283 203 malariae, and malariae Marchiafava & Celli, 1885, as published in the combination Plasmodium malariae, become the oldest name for the Malignant Tertian Malaria Parasite, and claims might be advanced on its behalf as against the name falciparum Welch, 1897, even though it is at present an invalid name by reason of the fact that in the past it has been identified with, or rejected as a junior secondary homonym of, the name malariae Laveran, 1881. 267. By the combination of the three sets of measures set out above—and by no other means—it will be possible for the International Commission to provide that the name falciparum Welch, 1897, as published in the combination Haematozoon falciparum, shall be the oldest available specific trivial name for the Malignant Tertian Malaria Parasite. (3) The specific trivial name for the ee Tertian Malaria Parasite 268. The oldest available specific trivial name for the Benign Tertian Malaria Parasite is the name vivax Grassi & Feletti, 1890, as published in the combination Haemamoeba vivax. This is the name in universal use for this species. Accordingly, no action by the International Commission under its Plenary Powers is required in this case. (4) Certain terms consisting of Latin adjectives cited in the genitive case published as technical designations, but not as names, for human malaria parasites 269. Dealing—in Part 1 of the present paper—with the specific trivial names bestowed, or alleged to have been bestowed, upon one or other of the human malaria parasites, I had occasion at three points to draw attention to the publication of Latin adjec- tives in the genitive case in grammatical agreement not (as required by Article 14) with the generic name of the species concerned but with the specific trivial name of that species. For the reasons then explained I am of the opinion that the adjectives so published should be looked upon as constituting technical 204 OPINIONS AND DECLARATIONS designations for the species or “ varieties ’’ concerned but not as names possessing rights in zoological nomenclature. In each case I have rejected the claims which have been advanced in favour of the recognition of these terms as specific or subspecific trivial names. It is clearly necessary that, as part of any general cleaning-up of the nomenclature of the human malaria parasites, the status of these terms should be the subject of an authoritative decision. I therefore recommend that the Commission should rule that the terms in question contravene the provisions of Article 14 to such an extent as to put them outside the scope of the decision taken by the Paris Congress in 1948 that minor infringements of that and certain other Articles should be sub- ject to automatic correction (1950, Bull. zool. Nomencl. 4: 68) by later authors, that these terms should be regarded as technical designations and not as trivial names and that they are therefore to be rejected as possessing no status in zoological nomenclature. The terms which I ask should be made subject to the reeommenda- tion submitted above are: (1) the terms quartanae, tertianae and quotidianae published by Celli & Sanfelice in 1891 in connection with the specific name Plasmodium malariae Marchiafava & Celli, - 1885 (paragraphs 128—139); (2) the terms quartanae, tertianae and irregularis published by Kruse in 1892 in connection with the combination Plasmodium malariae (paragraphs 144—150); (3) the terms guartanae, tertianae, wundecimanae, sedecimanae and vigesimo-tertianae published by Lewkowicz in 1897 in connection with the generic name Haemosporidium Lewkowicz, 1897 (para- graphs 169—174). A recommendation that the International Commission should invite the International Congress of Zoology to insert words in Article 14 for the purpose of laying down the general principle involved in the foregoing cases is submitted at a later point in the present paper (paragraphs 279—280 below). __ (5) The generic name “ Oscillaria ”’ as used by Laveran in 1881 270. The first step needed to introduce order into the generic nomenclature of the human malaria parasites is to lay the ghost of the alleged generic name Oscillaria Laveran, 1881. In Part 2 of the present paper (paragraphs 187—198) I have given grounds for believing that Laveran never published this name as a new OPINION 283 205 generic name but merely applied it in 1881 to the species of malaria parasite which he then named malariae, because he considered it appropriate on taxonomic grounds to refer that species to the genus Oscillaria Schrank, 1823. In the same paragraphs I have shown that Schrank looked upon his genus Oscillaria as a genus belonging to the Animal Kingdom, and therefore that, under Article 1 of the Rég/es, the name Oscillaria Schrank, 1823, retains its status in zoological nomenclature, notwithstanding the fact that the species of Algae included by Schrank in this genus are not now regarded as belonging to the Animal Kingdom. In these circumstances, even if it could be clearly established that in his paper published in 1881 Laveran treated the name Oscillaria as a generic name then introduced by himself for the first time, the name Oscillaria Laveran, 1881, would be invalid, by reason of being a junior homonym of the name Oscillaria Schrank, 1823. 271. It is, therefore neither necessary nor desirable that the International Commission should use its Plenary Powers to suppress the generic name Oscillaria Laveran, 1881, for, as appears to me to be clear, no such name was ever published. It is however essential that the Commission should include in its. decision a Ruling that, in so far as Laveran may have published this as a new name and not as a usage of the name Oscillaria Schrank, 1823, the name Oscillaria Laveran, 1881, which would. in that event have been brought into existence, is to be rejected. as a junior homonym of Oscillaria Schrank, 1823. (6) The generic name ‘‘ Plasmodium ” Marchiafava and Celli,. 1885 272. As the result, mainly, of an uncritical following, by later authors, of pronouncements by Liithe and later by Schaudinn. (paragraphs 46—51), the nominal genus Plasmodium Marchiafava. & Celli, 1885, has been universally treated for many decades as having Oscillaria malariae Laveran, 1881, as its type species, and that nominal species has been consistently misidentified as representing the Quartan Malaria Parasite. In actual fact, as has been shown in Part 2 of the present paper, the type species 206 OPINIONS AND DECLARATIONS of this genus is the nominal species Plasmodium malariae Marchiafava & Celli, 1885, by monotypy (paragraphs 199—204) and this nominal species represents not the Quartan Malaria Parasite but the Malignant Tertian Malaria Parasite (paragraphs 53—77). 273. Very great confusion would arise if at this late date the type species of this genus were to be changed from the Quartan Malaria Parasite to the Malignant Tertian Malaria Parasite, in view of the fact that some malariologists regard these species as belonging to different genera, and a transfer of type species of the kind required by the strict application of the Régles would mean that the name Laverania Feletti & Grassi, 1889, universally used by specialists who on taxonomic grounds recognise two genera, would disappear as a junior subjective synonym of Plasmodium Marchiafava & Celli, 1885, while for such specialists the name Haemamoeba Feletti & Grassi, 1889, would replace the name Plasmodium Marchiafava & Celli, 1885, as the generic name for the Quartan Malaria Parasite. | 274. In these circumstances, there is, therefore, a very strong case for the use by the International Commission on Zoological Nomenclature of its Plenary Powers for the purpose of validating the current practice under which the Quartan Malaria Parasite is universally accepted as the type species of the genus Plasmodium Marchiafava & Celli, 1885, after setting aside the existing indica- tion, under Rule (c) of Article 30, of Plasmodium malariae Marchiafava & Celli, 1885 as the type species of this genus. It must be realised, however, that this will involve the use of the Plenary Powers to designate as the type species of the genus Plasmodium Marchiafava & Celli, 1885, a species not included by those authors in this genus when they published the name Plasmodium. Such use of the Plenary Powers in the interests of nomenclatorial stability will, however, constitute no innovation, for already in 1935 at Lisbon (Lisbon Session, 2nd Meeting, Conclusion 22) (1943, Bull. zool. Nomencl. 1 : 20—23) the Com- mission took precisely similar action for the purpose of validating the universally accepted usage of the name Satyrus Latreille, 1810 (Class Insecta, Order Lepidoptera), a decision which has since been formally promulgated as Opinion 142 (1943, Ops. Decls. int. Comm. zool. Nomencl. 2 : 67—80). OPINION 283 207 275. The action which is now recommended is that the Inter- national Commission, acting under its Plenary Powers, should set aside the indication, by monotypy, of Plasmodium malariae Marchiafava & Celli, 1885, as the type species of the nominal genus Plasmodium Marchiafava & Celli, 1885, and in its place designate as the type species of the foregoing genus the nominal species Haemamoeba malariae Feletti & Grassi, 1889 (the Quartan Malaria Parasite). (7) The generic name ‘“‘ Laverania ” Feletti and Grassi, 1889 276. The generic name Laverania Feletti & Grassi, 1889 (usually misdated as having been published in 1890, and incorrectly attributed to the above authors in the reverse order) is universally used as the generic name for the Malignant Tertian Malaria Parasite by those malariologists who regard that species as being generically distinct from the Quartan Malaria Parasite. This usage is, however, incorrect, for that species is also the type species of the nominal genus Plasmodium Marchiafava & Celli, 1885. The objections to the strict application of the Rég/es in this case have been summarised in paragraph 273 above and need not be repeated. 277. In order to secure that the name Laverania Feletti & Grassi, 1889, shall be available for use by those specialists who place the Malignant Tertian Malaria Parasite in a different genus from that to which they refer the Quartan Malaria Parasite, while placing the last-named species in the genus Plasmodium Marchiafava & Celli, 1885, it will be necessary, in the first instance, for the International Commission on Zoological Nomenclature to use its Plenary Powers to set aside the indication by monotypy of Plasmodium malariae Marchiafava & Celli, 1885, as the type species of the genus Plasmodium Marchiafava & Celli, 1885. For the reasons explained in paragraph 273 above, this action is in any case necessary, in order to secure the continued use of the generic name Plasmodium Marchiafava & Celli, 1885, in its accustomed sense and has already been recommended for that 208 OPINIONS AND DECLARATIONS reason alone. In addition, it will be necessary, before the name Laverania Feletti & Grassi can become the oldest available generic name for the Malignant Tertian Malaria Parasite for the Inter- national Commission on Zoological Nomenclature to take appro- priate action under its Plenary Powers to deprive the generic name Hdadmatophyllum Metschnikoff, 1887, of its prior rights. As has already been explained (paragraph 212), the name Hdmatophyllum Metschnikoff was published as a substitute name (nom. nov.) for the name Plasmodium Marchiafava & Celli, 1885, and accord- ingly takes, as its type species, the same species (Plasmodium malariae Marchiafava & Celli, 1885) as is the type species of that genus. Since that nominal species represents the Malignant Tertian Malaria Parasite, the effect, if no further action were to be taken, of changing the type species of the genus Plasmodium Marchiafava & Celli from Plasmodium malariae Marchiafava & Celli (the Malignant Tertian Malaria Parasite) to Haemamoeba malariae Feletti & Grassi, 1889 (the Quartan Malaria Parasite), would be to make the name Hdmatophyllum Metschnikoff, 1887, the oldest available generic name for the Malignant Tertian Malaria Parasite for those malariologists who regard that species as generically distinct from the Quartan Malaria Parasite. In order to make the name Laverania Feletti & Grassi, 1889, the oldest available generic name for the Malignant Tertian Malaria Parasite, it will, therefore, be necessary for the Commission to use its Plenary Powers in the case of the name Hdmatophyllum Metschnikoff, 1887, in the same way as that it is proposed that it should use those Powers in the case the name Plasmodium Marchiafava & Celli, 1885, that is, that it should set aside the indication by monotypy (through Rule (f) in Article 30) of the nominal species Plasmodium malariae Marchiafava & Celli, 1885, as the type species of Hdmatophyllum Metschnikoff, 1887, and should designate in its place the nominal species Haemamoeba malariae Feletti & Grassi, 1889, to be the type species of this genus. The effect of this action would be to make Hdmato- phyllum Metschnikoff, 1887, a junior objective synonym of Plasmodium Marchiafava & Celli, 1885 (in other words, to restore the relative position of these two nominal genera, a position which would have been upset if the type species of one of them (Plasmodium) had been varied under the Plenary Powers, while the type species of the other (Hdmatophyllum) remained un- changed.) By the foregoing means—and only by those means— OPINION 283 209 it will be possible for the International Commission to secure that the name Laverania Feletti & Grassi, 1889, shall be the oldest available generic name for the Malignant Tertian Malaria Parasite for those malariologists who regard that species as being generic- ally distinct from the Quartan Malaria Parasite. 278. The sole species placed in the genus Laverania Feletti & Grassi, 1889, at the time when they established that genus—and therefore the type species of that genus, by monotypy—was cited by those authors as “‘ Laverania malariae’ without the citation of an author’s name. In paragraph 222 of the present paper, I have expressed the view that there can be no reasonable doubt that the nominal species so cited was not a new nominal species then so named by themselves but was the nominal species Oscillaria malariae Laveran, 1881. In the past, however, the opposite view has been expressed by some specialists, and it is clearly necessary, therefore, that, in the decision now to be taken, the International Commission should leave no room for doubt on this question. This end could be secured by the adoption of either of two means: First, the Commission could rule under its Plenary Powers, that the name “‘ Laverania malariae” as cited by Feletti & Grassi in 1889 is to be interpreted as having the meaning which would have been conveyed if those authors had cited that species as “ Laverania malariae (Laveran, 1881) (=Oscillaria malariae Laveran, 1881).°> The second course would be for the International Commission to use its Plenary Powers to set aside the indication, by monotypy, of Laverania malariae Feletti & Grassi, 1889, or, as the case may be, Oscillaria malariae Laveran, 1881, as the type species of the nominal genus Laverania Feletti & Grassi, 1889, and in the place of the species so named, to designate the nominal species Haematozoon falciparum Welch, 1897, to be the type species of this genus. The latter course is, in my view, to be preferred, both because it is more consistent with the action proposed in the somewhat parallel case of the type species of the nominal genus Plasmodium Marchiafava & Celli, 1885, and partly because in any general clearing-up of the confused nomenclature of the human malaria parasites, it is desirable—because it is will promote clarity—that the nominal species which is to be designated as the type species of the genus concerned should be cited under the name which it is proposed 210 OPINIONS AND DECLARATIONS should in future be its valid name and not—as would be the case, if in the present instance if the type species of Laverania Feletti & Grassi, 1889, were to be designated by the name Oscillaria malariae Laveran, 188l—under an obsolete name which it is now proposed should be suppressed for all purposes. I accord- ingly recommend the adoption of the second of the two courses outlined above. i a j . oe ; OPINION 283 211 | PART 5 PROPOSED CLARIFICATION OF ARTICLE 14 OF THE “ REGLES ” ON THE QUESTION OF THE STATUS OF TERMS CONSISTING OF LATIN ADJECTIVES CITED IN THE GENITIVE CASE WHEN PUBLISHED AS THOUGH CON- STITUTING NAMES FOR SPECIES OR SUB- SPECIES _ 279. One of the serious difficulties encountered in determining the specific trivial names properly applicable to the human malaria parasites arises from the fact that no less than three authors (Celli & Sanfelice, 1891; Kruse, 1892; Lewkowicz, 1897) applied to these species terms consisting of Latin adjectives in the genitive case which, in the first two instances, had the appear- ance of being varietal (i.e. subspecific) trivial names and, in the third instance, of being specific trivial names. J have given grounds in Part 1 of the present Paper (paragraphs 134—138) for the view that, in fact, these adjectives were not intended by their respective authors to be trivial names in the sense prescribed in zoological nomenclature, but were looked upon by them rather as convenient technical designations for the species concerned. In formulating the recommendations submitted in Part 4 I have treated these terms in this light and have submitted a recommendation in regard for their disposal. While I am not aware of any other case where an author has published Latin adjectives in the foregoing manner, I think it highly probable that cases of this kind do exist, especially in the literature of groups of animals (notably Protozoa) which are of special interest in medicine and on which papers have been published by specialists, whose normal interests lie outside the field of systematic zoology. It seems to me, therefore, that it would be well to guard against the risk of misunderstand- ings arising in other fields similar to those which have caused so much confusion and inconvenience in the nomenclature of the human malaria parasites. 280. I accordingly recommend that, if, as I hope, the Inter- national Commission on Zoological Nomenclature approves the proposals which I have submitted for the stabilisation of the names of the human malaria parasites, it should at the same time recommend the International Congress of Zoology to clarify the 212 OPINIONS AND DECLARATIONS meaning of Article 14 in relation to the question discussed in the immediately preceding paragraph. The proposal which I now submit is that the International Commission should recommend the International Congress of Zoology:— to insert in the Régles provisions as follows:— (a) Where, in the case of a nominal species having, as its specific trivial name, a noun in the genitive case, a third term consisting of a Latin adjective is cited in gram- matical agreement not with the generic name of the species (as required by Article 14) but with the trivial name of the species, the Latin adjective so cited is not to be treated as having been published as a subspecific trivia] name, and a technical designation so published shall possess no rights in zoological nomenclature. Example: In the case of the term published as Plas- modium malariae quartanae by Kruse in 1892, the Latin adjective quartanae, being in grammatical agreement with the specific trivial name malariae and not with the generic name Plasmodium, is not to be treated as having been published as a subspecific trivial name. (b) Where a Latin adjective is cited in the genitive case in direct association with a generic name, the term so published is to be treated as being in grammatical agreement with a specific trivial name in the same case and number, understood grammatically though not ex- pressed, and a designation so published is to be rejected as a subspecific trivial name in like manner as in (a) above. Example : In the case of the term published as Haemosporidium quartanae by Lewkowicz in 1897, the Latin adjective quartanae is to be treated as being in grammatical agreement with the specific trivial name malariae, understood though not expressed, and is to be rejected as a subspecific trivial name. OPINION 283 213 PART 6 SUMMARY OF RECOMMENDATIONS (1) Recommendations for the stabilisation of the names for the human malaria parasites on the basis of current usage 281. Having now surveyed in detail the complicated and unfortunate history of the nomenclature of the human malaria parasites and having indicated the various measures which it would be necessary for the International Commission to take— mainly under its Plenary Powers—in order to provide a sound juridical basis for the names currently used for these species and thus to prevent the appalling confusion in the literature of malariology which would follow the strict-application of the Régles in the present case, I submit herewith in summary form the following recommendations for the consideration of the Inter- national Commission, namely that it should :— (1) cancel the incorrect particulars relating to the generic names Plasmodium and Laverania contained :— (a) in Opinion 104; (b) in consequence of (a) above, in the Official List of Generic Names in Zoology; | (2) use its Plenary Powers :— (a) to suppress for the purposes both of the Law of Priority and of the Law of Homonymy the under- mentioned trivial names for the Malignant Tertian Malaria Parasite :— (1) malariae Laveran, 1881, as published in the combination Oscillaria malariae; (ii) malariae Marchiafava & Celli, 1885, as published in the combination Plasmodium malariae ; 214 OPINIONS AND DECLARATIONS (iii) malariae Feletti & Grassi, 1889, as published — in the combination Laverania malariae, in — so far as this name was published as a new © name and not as the trivial name malariae Laveran, 1881; (b) to suppress for the purposes of the Law of Priority — but not for those of the Law of Homonymy the © under-mentioned trivial names published for the © Malignant Tertian Malaria Parasite:— (i) falciforme Antolisei & Angelini, 1890, as — published in the combination Ematozoo — (correction of ematozoo) falciforme; (ii) Javerani Labbé, 1894, as published in the combination Haemamoeba laverani; (111) falciforme Thayer & Hewetson, 1895, as P published in the combination Haematozoon — falciforme ; (c) to set aside the indication, by monotypy, of Plas- modium malariae Marchiafava & Celli, 1885 (the © Malignant Tertian Malaria Parasite) as the type species (i) of the genus Plasmodium Marchiafava & Celli, 1885, and (ii) of the genus Hédmato- — phyllum Metschnikoff, 1887 (the name of which ~ was published as a substitute name (nom. nov.) — for Plasmodium Marchiafava & Celli, 1885, in the — erroneous belief that the latter name could © properly be rejected on the ground of inappro- ~ priateness), and in place of the foregoing species ~ to designate Haemamoeba malariae Feletti & Grassi, 1889 (the Quartan Malaria Parasite) to be the type species both of the genus Plasmodium Marchiafava & Celli, 1885, and of the genus Hdamatophyllum Metschnikoff, 1887; — a ae re OPINION 283 215 (d) ‘to validate the under-mentioned trivial names :— (i) malariae Feletti & Grassi, 1889, as published in the combination Haemamoeba malariae, to be the name of the Quartan Malaria | Parasite, notwithstanding the fact that, prior to the suppression under the Plenary Powers of the trivial names consisting of the word “ malariae,” specified in (a) above, that name had been invalid as a junior secondary homonym in the genus Plasmodium; (ii) falciparum Welch, 1897, as published in the combination Haematozoon falciparum, to be the name of the Malignant Tertian Malaria Parasites; (e) to set aside the indication, by monotypy, of Laverania malariae Feletti & Grassi, 1889, or, as the case may be, of Oscillaria malariae Laveran, 1881 (being names for the Malignant Tertian Malaria Parasite suppressed under (a) above) as the type species of the genus Laverania Feletti & Grassi, 1889, and, in the place of the species so named, to designate Haematozoon falciparum Welch, 1897, to be the type species of that genus; (f) to validate the generic name Laverania Feletti & Grassi, 1889 (type species, by designation under the Plenary Powers, under (e) above: Haemato- zoon falciparum Welch, 1897, validated under the Plenary Powers, under (d) above), as the generic name of the Malignant Tertian Malaria Parasite ; (3) declare the under-mentioned generic names to be invalid or not required for the reasons severally stated below against the names in question :— 216 OPINIONS AND DECLARATIONS Generic name (1) (a) Oscillaria Laveran, 1881 (in so far as Laveran published this as anew name and not as Oscil- laria Schrank, 1823) (b) Hdmatophyllum Metschnikoff, 1887 (c) Haemamoeba Feletti & Grassi, 1889 (d) Ematozoo Antolisei & Angelini, 1890 Reason why generic name cited in Col. (1) is invalid or not required (2) Invalid because a junior hom- onym of Oscillaria Schrank, 1823, that name retaining under Article 1 its rights under Article 34, notwith- standing the fact that the genus so named has been transferred to the Vegetable Kingdom Invalid because a junior ob- jective synonym of Plas- modium Marchiafava & Celli, 1885, the two nominal genera having the same nominal species as type species Invalid because the type species of this genus (Haemamoeba malariae Feletti & Grassi, 1889) has, under (2)(c) above, been designated under the Plenary Powers to be the type species of the genus having the older name Plasmodium Marchiafava & Celli, 1885 Not required because its type species (Ematozoo falciforme Antolisei & Angelini, 1890) is a junior subjective syno- nym of Haematozoon falci- parum Welch, 1897, desig- OPINION 283 Generic name (1) (d) Ematozoo Antolisei & Angelini, 1890 ( continued) (e) Haematozoon Thayer & Hewetson, 1895 (f) Haemosporidium Lewkowicz, 1897 217 Reason why generic name cited in Col. (1) is invalid or not required (2) nated under the Plenary Powers, under (2) (e) above, to be the type species of the genus Laverania Feletti & Grassi, 1889 Invalid, because a junior homonym of Haematozoon Leisering, 1865; in addition, not required because its type species (Haematozoon falciforme Thayer & Hewet- son, 1895) is a junior sub- jective synonym of Haema- tozoon falciparum Welch, 1897, designated under the Plenary Powers, under (2) (ec) above, to be the type species of the genus Laver- ania Feletti & Grassi, 1889 Invalid because its type species Haemamoeba malariae Feletti & Grassi, 1889 (by selection under Article 30, Rule (g) and Opinion 46) is the same nominal species as that which, under the Plenary Powers, has, under (2) (c) above, been desig- nated as the type species of the genus having the older name Plasmodium Marchia- fava & Celli, 1885 218 OPINIONS AND DECLARATIONS (4) declare that the undermentioned trivial names, each of which was published as the name of a new avian parasite but in the description of each of which there appeared an incorrect statement that the parasite in question had been found in the blood of human malaria patients, are not available as trivial names for the Malignant Tertian Malaria Parasite (the parasite mis-identified with the avian parasite concerned), these trivial names adhering under the Régles to the avian parasites from which the original descriptions of these parasites were drawn up by their respective authors :— (a) praecox Grassi & Feletti, 1890, as published in the combination Haemamoeba praecox; (b) immaculata Grassi, 1891, as published in the com- bination Haemamoeba immaculata; (5) declare that the undermentioned terms consisting of Latin adjectives published in the genitive case, in agreement not with the generic name (as required by Article 14 (1)(a)) but with the specific trivial name, either expressed or understood, were published not as subspecific trivial names of human malaria parasites, but as technical designations for those species and that the Latin adjec- tives in question accordingly possess no status under the Régles as subspecific trivial names :— (a) quartanae Celli & Sanfelice, 1891, as published in connection with the combination Plasmodium malariae ; (b) tertianae Celli & Sanfelice, 1891, as published in connection with the combination Plasmodium malariae ; (c) quotidianae Celli & Sanfelice, 1891, as published in connection with the combination Plasmodium malariae ; (d) tertianae Kruse, 1892, as published in connection with the combination Plasmodium malariae; OPINION 283 219 (e) guartanae Kruse, 1892, as published in connection with the combination Plasmodium malariae ; (f) irregularis Kruse, 1892, as published in connection with the combination Plasmodium malariae; (g) tertianae Lewkowicz, 1897 as published in con- nection with the generic name Haemosporidium; (h) quartanae Lewkowicz, 1897, as published in con- nection with the generic name Haemosporidium; (i) undecimanae Lewkowicz, 1897, as published in con- nection with the generic name Haemosporidium; (j) sedecimanae Lewkowicz, 1897, as published in con- nection with the generic name Haemosporidium; (k) vigesimo-tertianae Lewkowicz, 1897, as published in connection with the generic name Haemo- sporidium ; (6) place on record:— (a) that there is no such generic name as Haemato- monas Osler, 1886, Osler in the passage in question not having published a new generic name but having referred to the previously published name Haematomonas Mitrophanow, 1883; (b) that the undermentioned generic and trivial names published for human malaria parasites by Dani- lewsky, 1891, possess no status under the Régles, the paper in which they were published having been declared by the Commission in Opinion 101 to be unavilable for nomenclatorial purposes :— (i) The generic names :— Cytamoeba Danilewsky, 1891 Cytosporon Danilewsky, 1891 Cytozoon Danilewsky, 1891 Haemocytosporon Danilewsky, 1891 220 OPINIONS AND DECLARATIONS (ii) the trivial name hominis Danilewsky, 1891, as published in the binominal combination Laverania hominis; (c) that the trivial name quartana Labbé, 1894, as published as a subspecific trivial name in the trinominal combination Haemamoeba_ laverani var. guartana, is not required for the Quartan Malaria Parasite, it being a junior subjective synonym of the trivial name malariae Feletti & Grassi, 1889, as published in the combination Haemamoeba malariae, validated under the Plen- ary Powers under (2)(d) above; (d) that the undermentioned alleged trivial names, not having been published, are cheironyms and accordingly possess no status under the Régles:— (i) irregularis Sakharov, erroneously alleged to have been published in. 1892 as a sub- specific trivial name in the combination Haemamoeba febris irregularis ; (11) tropica Koch, erroneously alleged to have have been published in 1899 in the com- bination Plasmodium tropica ; (7) declare that the trivial name vivax Grassi & Feletti, 1890, as published in the combination Haemamoeba vivax, 1s the oldest available trivial name for, and therefore the valid trivial name of, the Benign Tertian Malaria — Parasite; (8) declare that the trivial name tertiana Labbé, 1894, as published as a subspecific trivial name in the combina- tion Haemamoeba laverani var. tertiana, is not required for the Benign Tertian Malaria Parasite, being a junior subjective synonym of the earlier published trivial name vivax Grassi & Feletti, 1890; OPINION 283 221 (9) substitute the following particulars in regard to the generic names Plasmodium and Laverania in the Official List of Generic Names in Zoology in place of the particulars deleted therefrom in accordance with (1) above:— (a) Plasmodium Marchiafava & Celli, 1885 (type species by designation under the Plenary Powers: Haema- moeba malariae Feletti & Grassi, 1889) (the Quartan Malaria Parasite); (b) Laverania Feletti & Grassi, 1889 (type species by designation under the Plenary Powers: Haemato- zoon falciparum Welch, 1897) (the Malignant Tertian Malaria Parasite) (generic name to be used by authors who consider the Malignant Tertian (or Aestivo-Autumnal) Malaria Parasite to be generically distinct from the Quartan Malaria Parasite). (2) Recommendation for the clarification of Article 14 in relation to the status of terms consisting of Latin adjectives cited in the genitive case when published as though constituting names for species or subspecies 282. Arising out of the Recommendation numbered Recom- mendation (5) in the summary submitted in the immediately preceding paragraph, J recommend that the International Com- mission on Zoological Nomenclature should invite the Inter- national Congress of Zoology’ :— to insert words in the Rég/es providing as follows :— (a) Where, in the case of a nominal species having, as its specific trivial name, a noun in the genitive case, a third term consisting of a Latin adjective is cited in grammatical agreement not with the generic name of the species (as required by Article 14) but with the trivial name of the species, the Latin adjective so cited is not to be treated as having been published as a 28 This recommendation was approved by the International Commission on Zoological Nomenclature, on whose recommendation a provision on the lines suggested was inserted in the Régles by the Thirteen International Congress of Zoology, Paris, 1948 (1950, Bull. zool. Nomencl. 4 : 624—625). OPINIONS AND DECLARATIONS subspecific trivial name, and a technical designation so published shall possess no rights in zoological nomenclature. Example: In the case of the term published as Plasmodium malariae quartanae by Kruse in 1892, the Latin adjective quartanae, being in grammatical agreement with the specific trivial name malariae and not with the generic name Plasmodium, is not to be treated as having been published as a subspecific trivial name. (b) Where a Latin adjective is cited in the genitive case in © direct association with a generic name, the term so published is to be treated as being in grammatical agreement with a specific trivial name in the same case and number, understood grammatically though not expressed, and a designation so published is to be rejected as a subspecific trivial name in like manner as in (a) above. Example: In the case of the term published as Haemosporidium quartanae by Lewkowicz in 1897, the Latin adjective quartanae is to be treated as being in grammatical agreement with the specific trivial name malariae, understood though not ex- pressed, and is to be rejected as a subspecific trivial name. OPINION 283 223 . APPENDIX 1 The discovery by Golgi of the Quartan Malaria Parasite mey SIR RICKARD CHRISTOPHERS, C.LE., O.B.E., F.R.S. (University of Cambridge, Department of Zoology, Cambridge) (Enclosure to a letter dated 24th March 1944 addressed to Francis Hemming) Golgi’s complete works on malaria have been republished in a volume by Perroncito (Perroncito, A., 1929, Gli studi di Camillo Golgi sulla malaria, Rome). Five papers are given prior to 1890. I have been carefully over these and the only ones relevant to the present issue are:— Golgi, C. 1885. Sull’infezione malarica. Communicated R. Accad. di Med. di Torino at Meeting of 20th November 1885. Published Giornale della R. Accademia di Medicina di Torino in 1885, in Vol. 33, p. 734. Published later (in 1886) in extenso (di poi in esteso) in Archivio per le Scienze medische di Torino Vol. 10, p. 109. Republished (with the plate inscribed “ Arch. per le Sci. Med., Vol. X,’ so presumably same plate used as for the 1886 printing) Perroncito, loc. cit. 1929. Golgi, C. 1886. Ancora sulla infezione malarica. Communicated Soc. Med. Chir. di Pavia at Meeting of Sth June 1886. Published in 1886 in Gazetta degli Ospedali No. 53 (Wenyon gives Vol. 7, p. 419). 224 OPINIONS AND DECLARATIONS Golgi, C. 1889. Sul ciclo evolutivo dei parassiti malarici nella febbre terzana. Diagnosi differenziale tra i parassiti endoglobulari malarici della terzana e quelli della quartana. Published in 1889 in Arch. per le Sci. Med. Vol. 13, p. 173. In the first paper Golgi showed that in quartan malaria the pyrexial attacks correspond with segmentation of the parasite — and that there may be a double quartan or quotidian sequence of attacks, in which case they correspond to two or three generations of parasite respectively. He described the various stages the parasite goes through and gives a plate with unmistakable forms of quartan. Further he points out that the tertian fever is not associated with the quartan parasite “‘ Riguardo alla terzana si puo anzi dire, a priori, che il parassita rappresentante l’infezione © malarica, deve avere un ciclo di sviluppo diverso da quello della quartana e sue combinazioni.” In the second paper he returns to this latter point and describes the tertian parasite (named vivax by Grassi & Felletti). In the third paper he describes in detail the differences between quartan and tertian parasites, etc. There can be no doubt but that the first paper is that in which Golgi first recognised and described the quartan parasite, though he did not give it a zoological name. He not only gives beautiful reproductions in his plate of this parasite which include most of its characters but he also worked out its periodicity. He also clearly suspected another species to be present and practically said that this was not the quartan. —_ eee Oe eee ee a OPINION 283 225 APPENDIX 2 Reproduction in facsimile of the paper entitled Sui parasiti della malaria by Feletti (R.) and Grassi (B.) as published in the serial publication Riforma medica, Napoli, in its issue of 15th January 1890 and of the supplement to the foregoing paper entitled Sui parasiti della malaria : aggiunta alla nota preliminare published in the same serial publication in its issue of Ist March 1890 Documents 1 to 3: Feletti (R.) & Grassi (B.), 1890, Sui parasiti della malaria : facsimile reproductions of pages 62 to 64 of the issue of 15th January 1890 of the serial publication Riforma medica, Napoli, in which the foregoing paper was published Document 1: Facsimile reproduction of page 62 Document 2: Facsimile reproduction of page 63 Document 3: Facsimile reproduction of page 64 Documents 4 and 5 : Feletti (R.) & Grassi (B.), 1890, Sui parasiti della malaria : aggiunta alla nota preliminare : facsimile repro- ductions of pages 296 and 297 of the issue of Ist March 1890 of the serial publication Riforma medica, Napoli, in which the foregoing paper was published Document 4: Facsimile reproduction of page 296 Document 5: Facsimile reproduction of page 297 PAE See ae = I AE ig a i ‘ of ® oe Se! oe «gee a ee oo eae ee TS ome a er ee t <—~<- ee ne -- iy Sit Ter hens aloe wer ie | Fr £ SHIVA Ren io aes rs ve a haat D Gee ee At Wis tVe pipe alot, ii UE Lica lehoe ny} ribet 1 . , b : @ a _ f Pie et Ged sernseagad ie . ‘ . * . oY aAate iy tee, i MAAS Dig 4) 2 5th Show Whig 4 TAMVGSG ? Rabati ie 2 Ye: a rt pe) - a " * may Sh) oeae } Be ptt. ae ted oy j fair toa eg « 4 Wea aeR eB ewe eb .. iy ae + Z ne lina Sag * lias agi 6 ll Pelee sire ate itn is | | | | ee bilendo la anastomosi intestinale con l'e- - sclusione permanente della sede della oc- » clusione dal passaggio attivo delle fecce. + 18. La ripetizione dell’ accidente deve ' @ssere impedita con I’ accorciametto del Mesentere a mezzo di una ripiegatura pa- lela all’asse lungo dell’intestino, © con Ja sutura déll’apice della ripiegatura alla Fadice del mesentere. LAVORI E LEZION! ORIGINAL! RB. UNIVERSITA DI CATANIA (Prof. R. Feverti e B Grassi.—Sui parassiti della malaria. Lo stato presente della questione riguar- ante i parassiti della malaria ¢ questo. Da una parte Laveran, Marchiafava, Celli, Golgi, Guarneriecc. sostengono ‘che 6 causa della malaria un parassita da loro studiato, il quale presenta sicure ‘somiglianze, secondo Laveran con le 0- _ scillarie, secondo Marchiafava, Celli e _ @uarneri coi Micetozoi, con gli Sporozoi § coi Flagellati. Da un’ altra parte stanno Tommasi- €rudeli, Mosso, Maragliano, Hayem ece., i quali ammettono che il supposto essere vivo, cosi bene studiato dai sud- etti Autori, altro non sia che un prodot- dell’alterazione e del successivo disgre- nento dei globuli rossi. i fronte a due opinioni diametralmen- opposte, la maggior parte degli stu- si crede prematuro qualunque gindizio. ben vero che i corpi semilunari, i fla- ati, i plasmodi istintivamente (se ci si mette la parola) si giudicherebbero es- vivi. & vero chei cosi detti cielt evo- ini, messi in luce specialmente da quel acissimg osservatore che é il prof. Igi, a primo aspetto sembrano riferi- i gg ad esseri inferiori. Ma non é n vers She il globulo rosso, alterandosi, presentare forme, che hanno somi- anza con le semilune, con le marghe- coi corpi flagellati ece. Somiglianza olontana, ma sufficiente a far sorgere | dubbi. Com’ é vero del pari che, pre- Mandosi la febbre malarica comune- sente ad accessi, si pud supporre che le | erazioni dei globuli rossi corrisponda- ll’ andamento dei singoli accessi, e | telino percié dei cicli evolutivi. Tea gli studiosi che dubitavano, erava- foi pure. Ed uno di noi (Grassi) “ee@eri al collega Celli, che la piu ov- fia goluzione dell’ oscuro quesito doveva bwarsi nello studio delVintima struttura gupposto parassita, e sopratutto nella Rastragione che esso possiede un nu- bo, Tl Gelli e il Guarneri si misero ‘appanto sopra questa strada. A noi sem- “sen Herd che essi non siano riusciti a toc- la meta; e ci pare assai ragionevole ila prudente riserva che adoperano nell’in- terpretare i nuovi fatti trovati. E invero: “gn nucleo che si colora meno, 0 Come ] lasma; un endoplasma, talvolta ad lato dell’ éctoplasma, e inoltre chiaro, molto debolmente, (nelle fi- My . kaise DJ _DIeA - zed LA Rifu gure pero é incoloro, aentre l’ectoplas- ma sarebbe colorabile-molto intensamen- te; i granuli di pigmento non occupanti mai |’ endoplasma; dei flagelli con pig- mento; il nucleo delle semilune ritenuto simile a quello dei coccidi; un principio del differenziamento del nucleo ecc., ci par- vero espressioni oscure e fatti poco pro- vanti I’ asserto degli Autori. Bisogna con- fessare pero che I’ interessantissimo lavo- ro di questi Autori segno un vero pro- gresso, perché fece nascere la speranza di potere sciogliere definitivamente la que- stione. I dubbi, che abbiamo yoluto qui accen- nare, ci mossero ad imprendere nuove ri- cerche. Molti furono i tentativi infruttuo- si, durante i quali, ora propendevamo a credere che il supposto protozoo fosse uno pseudoparassita, perche non presen tava indubitabili caratteri di essere vi- vente; ora invece ritenevamo il contrario, perché impressionati dai corpi in segmen- tazione e dalle semilune, che ci parevano troppo belli per poter essere prodotti da alterazioni d’ un globulo rosso. Pero dopo molte e molte oscillazioni siamo arrivati alla conclusione desiderata. Il problema fu da noi posto in questi termini: Se il plasmodio e la semiluna sono Rizopodi, o Missomiceti, 0 Chitridi (che altro vero protisto possono essere’), debbono avere quasi sicuramente un nu- cleo, e questo deve nella riproduzione comportarsi come tutti i nuclei degh es- seri Vivi. : Ora noi, modificando alquanto i metodi di ricerca finora usati, siamo appunto riusciti a vedere costantemente e molto esattamente, e percio a potere con sicu- rezza interpretare, quanto avevano gia osservato Celli e Guarneri. Non solo, ma abbiamo potuto seguire il nucleo an- che durante il processo di divisione dei corpi pigmentati. Riserbando al lavoro completo di espor- re i metodi e di far rilevare punto per punto quanto spetta agli altri Autori, vo- gliamo qui semplicemente riassumere le nostre osservazioni e le conseguenze che ne deriyano. Il cosi detto plasmodio (1) consta di pla- sina (corpo o sostanza cellulare), in cui l’ ectoplasma e l’endoplasma non sono di- stinti (il che sta in rapporto specialmente con Ja sua molto imperfetta locomozione e col suo peculiare modo di nutrizione), e di un evidentissimo e grosso nucleo, ve- scicolare, come si trova in molti Rizopo- di. Il nucleo é per lo pit molto eccentri- co; possiede una delicatissima membrana, succo nucleare (nucleochilema), che pare semisolido, e reticolo. Questo reticolo nu- (1) Questi studi si riferiscono specialmente ai pla- smodi della quartana. Il termine plasmodio, nel senso usato dagli A. e da noi seguito per render facile la lettura della presente Nota, é Gel tutto improprio Sug- geriamo di sostitairvi il termine amibula, quando é@ 5 cethas ; Pavesi ancor piccolo: amtba, quando @ grande: amiba in rie cessive diyisione impiccoliscono. Tutti questi fone” pogo si denominera quindi il corpo pigmentato. cleare @ rappresentato da un nodo, forma di nucleolo pill 0 meno eccentrigg il qualé é di varia forma: rotondeggiante. o subtriangolare, 0 subquadrangolare : da esso partono parecchi fili (3-4) delicg tissimi (nei plasmodi | ancor piccoli noy sono quasi mai visibili), che vanno very la membrana. Questa assume talora, pg preparati artificialmente colorati, una tip. | ta come quella del corpo nucleoliforme,§ e pud mostrare degl’ ispessimenti egng mente colorati; se la colorazione appar tenga alla membrana stessa, od a delieg, tissimi fili del reticolo nucleare ad ggg addossati, non sappiamo decidere. Il ply sma ora presenta un aspetto reticolarg (spugnoso), ora mostra dei granuli finis.| simi (microsomi), che si colorano faci]. mente ad es. col bleu di metile. Nel pla sma poi possono trovarsi dei corpuscolj, che sono evidenti frammenti di globulo| rosso (cibo ancora indigerito), oppure de}, corpuscoli di melanina (residuo della die, gestione). Alle volte vi si notano anche: uno o parecchi vacuoli non contrattili. Rey: Come si vede, il corpo qui deseritto, tenendo conto che presenta movimenti ameboidi, puo fin dora eiudicarsi un! amis ba senza vacuoli contrattilé (in molti pro} tozoi parassiti maneano 1 vacuoli con trattil1). Man mano che va avvicinandosi alla maturazione, il plasmocdio ingrandisce, di- venta poi tondeggiante (corpo pigmentate degli Autori) e presenta spesse volte il suo -plasma distinto in duc strati: uno) esterno contenente i granuli di pigmen-! to, e uno interno, che ne e privo (questo si colora come il corpo jucleoltorme). Ingrandisce in proporzione forse pit i nucleo che il plasma. Vel nucleo poi it grossa relativamente molto di pin quel: corpiecciuolo nucleoliforme , che sopra # & accennato; esso cresce tanto da finite per occuparlo in gran parte. Abbiamo trovato molti esemplari in cui questo com picciuolo si era allungato a guisa di stoncello; molti altri in’cui tendeva adh vidersi o s’era gia diviso in due; molti} altri in cui ciascuno dei due tendeva pr) re a dividersi, 0 gia s era diviso in due altri (questi stadi corrispondono circa) 15, IS ore prima dell’ accesso); molti altt| esemplari, infine, in cui se ne contavano) 5, 6,8, 10 e piu. Ad un momento, che no) abbiamo ben determinato, attorno ad & gnuno dei corpiceinoli nucleoliformi, ¢ neoformati per divisione, disponesi suce nucleare ed una delicatissima membre) nella (1). (1) il fenomeno in discorso ci pare 4i verifichl : ogni guccessiva divisione, avendo trovato succo nucle, re attorno a ciascun corpicciuolo gia quando appena due, quattro, ecc, Ad aumentare la sostanss OF Corpo uncleoliforime, ossia la cromatina, concorre babilmente lo strato interny del plasma che 208 tiene pigmento. I corpicciuoli nucleoliformi acq il volume dello stesso corpicciuolo nell’ amibe finchd sono in numero di due o quattro; dopo (es consumato lo strato interno del plasma) ad P&rrebbero accennare ad ava riproduzione ft a { —————————— tb dllora un’amiba, ossia una ok , Abbiame nla plurinucleata (riproduzione endoge- ‘pa). Pid tardi, attorno ad ogni nucleo, di- onesi, pure in un momene e in un,;mo- ido che non abbiamo ancora precisato, un po’ di p plasma. Resta infruttuoso i] pigmente, e con es- so una sostanza, che probabijmente 6 une P tane periferica del plasma: il pigmen- to @ questa sostanza di solito si accumu- Jang in uD piccolo globo residuale. A poco a poco i nuclei, formatisi come testé si disse, col rispettivo plasma si allontanano jalquanto Y uno dall’ altro; e formano un muechietto , che (almeno nei casi da noi osservati) ha per lo pin forma di calotta pil 0 meno alta. Cosi si hanno le ben note figure di sporulazione, o di segmentazio- ne. In questo mucchietto distinguiamo : 1° una membranella, che © lo strato peri- feri¢o, 0, 5@ si ammette, la tanto contra- stata membrana del globulo rosso; 2." un globo residuale (nucleus de reliquat) (per lo piu collocato nel centro della faccia piana della calotta); 3.° i nuclei Vescico- lari (disposti per lo pit in uno strato che delimita-appunto la figura di calotta ) ‘ontenenti un evidente corpo nucleolifor- me pil o meno eccentrico, e¢ cireondati da plasma molto scarso, specialihente in corrispondenza a circa una meta del nu- cleo. Denomineremo questi corpi neoforma- tisi gimmospore , ce tors’ anche potremmo gia dirli amibule, Varia molto il numero delle eimnospo- te, che ogni amiba puo produrre, e forse anché varia alquanto il modo di divisione del nucleo (in alcuni esemplari troviamo ta gimnospora grande e parecehie pic- cole). Ti modo di riproduzione qui accennato trova sicnro riscontro nell lmoeha pigmen- tifera secoperta e deseritta da uno di noi (Grassi) e, riguardo al comportarsi del nucleo , nell cfwocha coli da uno di noi (Grassi) studiata. E.certo che il globulo rosso, il quale od 8 sfornito, od ha soltanto residui del m- dleo, alterandosi, non potrebbe riprodurre nuclei presentanti i caratteri suddescritti. TInuclei delle gimnospore sono perfet- tamente uguali a quello del plasmodio, da cui sono stati gvenerati (1). Recezionalmente le gimnospore si fon- dono in un vero plasmodio (con parecchi nuclei); alle volte lo fanno, com’ é gia no- to, i plasmodi endoglobulari coesistenti mun globulo rosso. E probabile che le SuMnospore con movimenti ameboidi in- Yadano il globulo rosso giovane, al mo- Ca ee mentezione (Arnold). Non osiamo pero darne una in- terpretazione definitiva: come pure non neghiamo che ei nvelei in discorso manchi Ia linina :plastina), La Straordinaria piccolezza del nostro oggetto ¢ la diffi- Soltd @' osservarlo spiegano la nostra riserva. (t) Supporre che il nucleo del plasmodio sia i} nu- 0 del globulo rosso giovane é assurdo, specialmente Perched: 1° if nucleo del plasmodio @ differente da uello del Slobulo rosso giovane; 2° perch? le semi- + Pill avanti) negli uccelli oceupano i proto- del Slobulo rosso, lasciando intatto i} nucleo. E ei BIE, eros 2 a ee cere LA RIFORMA eee eee mento in cui il’suo*nucleo si-é disfatto. | Con cid potrebbe star benissimo in rap- porto il fatto, anche da noi ripetutamente riscontrato, che ad un momento, i in cui il sangue é ricco di corpi in segmentazione, ne segue un altro in cui essi mancano o sono molto scarsi, e nei globuli rossi (del sangue ottenuto con la puntura del dito) non sono ancora presenti i plasmodi (1). La perfetta identita che abbiamo riscon- trata tra, il plasmodio appena entrato nel globulo rosso e la gimnospora dimostra che molto probabilmente tra l'uno e V’al- tra non v’ é stadio intermedio. I corpi semilunari derivano, com’é no- to, da cosi detti plasmodi, 0, giusta la no- stra: denominazione, da amibule. Queste hanno i caratteri sopracitati; ma quando diventano adulte, assumono appunto |’ a- spetto di semiluna. Le semilune differi- scono dal corpo pigmentato specialmente per Ja forma, per la disposizione de! pig- mento e per la quantita del plasma. Nella parte mediana della semiluna, quasi addossato alla faccia concava, e quindi sempre discosto dalla faccia con- vessa, sta 11 nucleo vescicolare, eccentri- co, percid, come nei corpi pigmentati. Ti nucleo spesso ha un corpo nucleolifor- me semplice, talvolta un corpo nucleoli- forme che si direbbe tencente a divider- si. I] plasma ha aspetto di quello del corpo pigmentato, pero 6 molto pit ab- bondante. Il pigmento sta per lo piu ad- dossato alla delicatissima membrana del nucleo. . Non é del tutto impossibile che la se- miluna, la cui membrana é uguale a quel- la del corpo in sporulazione, si riproduca per gemmazione, come credono Celli e Guarneri. Noi pure abbiamo verificato il fatto, ma non abbiamo potuto in aleun modo stabilire che nelle supposte gem- mule entri una parte del nucleo. Incli- niamo invece a credere che si tratti di un’ alterazione (diffluenza). In ogni caso . nessun zoologo , special- mente dopo le inoculazioni di Antoli- seie Angelini, esitera ad ammette- re che il corpo pigmentato della terzana e della quartana é differente dalla semi- Inna. Noi proponiamo di riferire l’uno al nuovo gen. Haemamoeba e Valtro al nuovo gen. Laverania. Nel periodo giovanile di amibula (pla- smodio secondo tutti gli A.) sono perfetta- mente identici |’ uno all altro. Pid tardi si differenziano grandemente anche per la forma, forse cid dipendendo dalla dif- ferente parte del globulo rosso, che ven- gono ad occupare. I generi in discorso, nello stato attuale delle nostre cognizioni, debbono inscri- (1) Se fosse vero che le gimnospore invyadessero sol- tanto i globuli rossi giovani, il valore della vera fa- gocitosi (che non osservamme ancora nel sangue fis- sato all’istante coi vapori di ac. osmico) si ridurrebbe assai. Ma di cid parleremo pid estesamente dopo nuove osservazioni. Sei en a padi Sa oI versi tra i Sarcodini (Rizopodi) nella tri- bu delle Amoebizormes. Le amibe in riposo (corpi pignicntatil & © ancora piuttosto giovani ( contenenti * gia pigmento), ma pitt frequentemente le: semilune (quest’ ultime spesso diventando! prima rotonde) possono emettere dei pro- lungamenti, che contengono 0 no pigmen- to, e che per lo pid non si possono dire né flagelli né pseudopodi , ma di solito rassomigliano pid a quelli che a questi, Tn quanto alle semilune, noteremo, che in tutti e cinque i mélarici con semailune da noi studiati nel novembre scorso qui a Catania , vedemmo i cosi detti corpl fla- gell&ti, sempre, tanto nell’ apiressia , che sottc la febbre? e cosi vicino come lon- tano da essa. Per trovarne sicuramente, | basta osservare il sangue ad una sufi- | ciente temperatura e impedirne lo schiac- ciamento, mettendo ad es. ai quattro an- goli del vetrino coproggetti un po’ di cera impastata coll’ olio. Neppure una volta trovammo forme fla- | gellate nel sangue, prima che fossero tra- ~ scorsi circa 20 minuti da che avevamo fatto il preparato. Non le trovammo nel sangue istantaneamente fissato coi vapori || di -acido osmico. Questi fatti collimano | con quanto ha osservato Danilewski, q nelle semilune degli uccelli. Consideran- © do che a produrre i flagelli in discorso ~~ concorre, secondo ogni verosimiglianza, soltanto il plasma,neghiamo che essi sia- : no capaci di riprodurre il corpo da qui | derivano. Sospettiamo invece che esprima- | no nient’ altro che fenomeni qgonici, for- | se paragonabili a quelli presentati da | certi flagellati , 1 quali nel periodo ago- nico si forniscono di uno pseudopodo, | che emettono e ritirano rapidissimamen- | te (Grassi, Blochmann). II sospet- to é avvalorato dalle svariate forme, che possono assumere i flagelli in discorso; dal fatto, che si distaccano via poco dopo che sono stati emessi , dal fatto che alle volte il corpo, invece di fornirsi di fla- gelli, presenta una sorta di orlo ondulan- ~ te, ecg. Infine il sospetto é giustificato dal: fatto da uno di noi (Feletti) mi- nutamente studiato: che dai globuli rossi possono uscire dei filamenti simili ai fla- gelli {n discorso, e come questi distac~' carsi, locomuoversi ecc. Giacché accennammo alle osservazioni di Danilewski, diremo che qnest’Au-' tore ha trovato negli uccelli delle forme, | che ritiene eguali a quelle della malaria. | Noi facciamo notare che veramente una | forma da lui riscontrata ha somiglianza | con Ja semiluna di Laveran,ma noné | della stessa specie, come risulta anche da un semplice corfronto delle figure (1). as r PR ca ree " In conclusione, sono certamente paras- w Finora non abbiamo trovato nel sangue dei mam- ~ miferi forme parassitarie paragonabili_a quelle della malaria; solamente uno di noi (Grassi) ba visto nel’ - di vari pipistrelli un Trypanosoma simile quello dei ratti. a saul # Ps ae } ‘ aeeeeee ernest rte liar Ont ne enn | Sovati descritte ¢ ‘meta dei ca, Safari Gelli : ‘Allinghan 5B 5‘ miglanza appartengo Ps, Vintestino n : rticolarmente agli Ajuindi si dovette si : sistono per lo meno duy °° ™Manovre ab ba malariae (delle feb *° s°™Plici e non verania malariae (dell¢#*: 208 Sono ari >, mentre nella ¢ _jeezioni, non vi ae lifficile e minuziosa, | naturalmente od E CLINICHE 2 pdistamente nel fond | SCIENTIFIC! i casi la ricerca a del mae ana ane ote we +) Facsimi Laveran,March | no ai Rizopodi | moebiformes, e ne esistono per lo meno due _ Specie: la Haemamocba malariae (delle feb- bri regolari) e la Laverania malayiae (delle febbri irregolari). o> ee RIVISTA DELLE CLINICHE E DEGL' ISTITUT! SCIENTIFIC COLLEGIO REALE DEI CHIRURGI DI LON- DRA. — (Prof. Bryant). Della colotomia ilia- Ca e lombare ; can particolare rignardo alla scelta dell’oporazione (I). * Colotomia lombare per cancro. Casi urgenti 100, di cui 55 segniti da buon Be SUCcesso: in 45 (casi troppo tardivi) morte entro un mese. g » 18 morte dopo 12 mesi. » 19 » tra il primo ed il secondo anno. » 12 » tra il secondo ed il terzo anno. » 6 Sono tuttora in vita, eiod 3 tra due anni © mezzo e cingue dopo l'operazione, ed 1 dopo sei anni. Casi non urgenti, 60: Nessuna morte nel mese successivo: 18 (cio 15 °/,) morirono nei primi 12 mesi. 24 (cioe 34 °/,) vissero da uno a due anni. : 16 (cioé 22 ©’) vissero da due a tre anni. 12 (cioé 17 °7,) sono ancora viventi, di cui 8 dai due ai sei anni dopo l’eperaziune. 38 °/, di tntto il numero sopravvissero allope- razione dai 2 ai sei anni. Aggiungo a questa statistica che. nei tre quarti dei*pazienti sopravvissuti alla operazione, questa ebbe un pievo risultato,in quanto che tutti i ma- -teriali fecali venivano emessi dall'ano artificiale. con gran vantaggio della lesione locale per eni Yoperazione erasi esecuita , essendo affatto ces- gata su di essa Virritamento prodotto dal passaz- gio delle fecce. Nell'altro quarto questo risulzato tanto desiderato non erasj ottenuto completamen- te, ele fecce, specialmente se liquids, continna- ¥ano in parte a passare per Ja via normale, az- ; >. giungondo irritaziune alla lesione ceciudente, n= eet che questi infermi intanto . ritraevano Yantaggi E ~ dalle lavande fatte dal reito uv dallano artificia- |. le, e dallapplicazione di suppositori calmanti, Ed ora credo opportuno discutere i] valore re- _,. Jativo dell’ operazione lombare od estrayerito- _ neale o della iliaca od intraperitoneale, quest’ ul- _tima preconizzata superiore alla prima special- ~ mente da Reeves, Hebert Allingham, Har- vison Cripps, e Chavasse. Gli argomenii principali su cui questi chirurgi si fondano per proclamare la superiorita della operazione iliaca sono: 1° Che essa in sé stessa = é pia facile della lombare ; 2° che Ja incisione | — dddominale permette di chiarire la diagnosi nei ‘ _easi oscari, prima di aprire l'intestino; 3° che per questa via non & possibile scambiare il grosso intestine con il tenne, coi duodeno, con lo sto- ~ maco, e che le possibili anomalie del colon non no rendere frustranea l’operazione, potendo essere l’addome esplorato in tutto il suo ambito; 4° che V'intestino pud prestamente e senza fativa essere portato fuori dell’ incisione, e per conse~ fissato stretiamente alla cute, senza pro- ‘durre indebita tensione sui punti di sutura; 5° che eolotomia lombare i! prolasso frequente e iderevole dell’ intestino induce seri disturbi CTR = Se x ‘ x Be * & siti della malaria le forme descritte da lafava Celli ecc.; secondo ogni verosimiglianza appartengo- © particolarmente agli 4- . neers a a ee ee pr e 6°, finalmente, che la Pasizione inguinale della ferita 8 molto pit conveniente all'infermo per la pulizia , e per l’applicazione aj apparecchi con- tentivi delle fecce e dei 88s intestinali. Esamineremo tutti questi progi in base alla espericnza clinica dei differenti casi, ammettendo che. la diagnosi di occlusione intestinale per can- cro © per altra causa sia certa, Tutti questi casi posson» andar divisi in tre grandi categorie: Nella prima vanno i casi Urgenti, associati a sintomi acuti generali ed a forte distensione dell’addome; nella seconda’ quclli dj occlusione intestinale cronica (prevalentemente dell'ultima porzione del retto senza sintomi generali acuti, ma con di- stensione dell’ addome ; nella terga i casi di oc- clusione rettale non urgenti » in cui i sintomi, quantunque bene sviluppati, non si associano a distensione dell’ addome. Ora | Operazione della colotomia ha le sue indicazioni Speciali per «cia- scuna di queste categorie di casi, Loperazione iliaca & essa pit facile della lom- bare ? Eseguita nelle due prime categorie , ciot quando vi é distensione addominale, l’operazione iliaca , cheeché ne dicano i snoi fautori, & diti- cile ed associata a pericoli indiscutibili. In questi casi lintestino tenue con assai faci- lita protrude e si avanza fuori dell'addome, e, quau- do vi sia necessita di ricereare lintestino grosso, di esplorare l'addome a scopo diagnostico, i pericoli consecutivi sono sempre tali da non essere di- sprezzati. Nella terza categoria di casi , essendo Yaddome non disteso, flaccido, la difficolté di rin- venire Vintestino é di molto ridotta. D'altra parte la distensione dell'addome costituisce una condi- ' zione favorevole per la colotomia lombare e la ricerca dell'intestino non é cireondata, per lo pin, da nessuna dificolta. Solo quando il paziente é pingue, pnd esservi qnalche difficolta nella ricerca dell’ intestino, ma in generale si riesce a dimi- nuire € superare questa ditiicolta, se il chirurgo dirige le sue ricerche in avanti » verso la ripie- gatura del peritoneo, e non in dietro, verso Vadipe lombare ; ¢ se nel tempo stesso fa girare il pa- ziente dalla posizione semiprona, che si era pre- scelta per cominciare loperazione , alla semisu- pina, in modo che Vintestino meccanicamente cada verso la incisione lombare: e listessa pra- tica riesee Lene quando I’ intestino @ totalmente 6 parzialmente vuoto, Neila colotomia iliaca, intanto fanche ammet- tendy che la difiicolts di rinvenire lintestino sia Minore che nejia lombare , i pericoli necessaria- mente inerenti alla incisione del peritonea pa- rietale debbono essere tenuti in considerazione. auche da quei chirurgi, che ‘credono potersi il peritoneo sano aprire senza alcun timore. Nel- Yoperazione lombare il peritoneo viene inciso solo in casi eccezionali, ed in questi casi non vi é nessuna ragione di differenza di gravezza tra Ja ferita del peritoneo nella regione iliaca e quella nella regione lombare. I pericoli della peritonite, qualunque essi sieno, saranno gli stessi per ambo le regioni. Nelle 170 colotomie lombari da ™e eseguite , solo due yolte, scientemente, ho Aperta la cavith peritoneale, senza che percid fosse Conseguito al- cun pericolo, Laonde sono obbligato a concludere che Voperazione iliaca non é pit facile della lom- bare, quando l’addome & disteso, quantunque ri- conosca che cid possa essere, quando }'addome & flaccido, Ne! cadavere la ricerca del colon dalla incisione iliaca pud sembrare, ed difficile, ma nell’ infermo l'incisione e la manovra del ricereare col dit cosa non peritoneale zioni iliache eseguite dall’Alling ham} terzi di queile del Cripps, l'intesting : tossi alla incisione, © quindi si dovette nae nel cavo peritoneale con manovre ab} complicate , le quali, se semplici non pel chirurgo sperimentato, non gono senza risultati per l'ammalato, mentre nella en} lombare, sete re eccezioni , non yj Sark. gno di questa ricerca difficile e minugi Tintestino , se disteso naturalmente od art mente, 8i presenta immadiatamente nel fond: incisione, Ed in tutti i casi la ricerca del « dalla incisione lombare non importa tgnte ap coltt, e noa implica tanti organi, quanto nell iliaca. Conclado quindi che la ricerca del} inte. stino riesce pit facile e meno Pericologa E regione lombare anziché dalla iliaca, Quanto al secondo argomento avanzato a fa del metodo iliaco, essere con questo, ciod, bile di verificare la diagnosi dei casi oscuri di aprire Vintestino , io osservo solo che 8 ama ‘difficile che possa csser7i una oscurita di diagnoy) nelle occlusioni rettali, le quali possono in gray parte essere esplorate dal dito, e nelle coliche, j cui sintomi non possono presentare alcun dubbig di interpretaziene. Ed anche se j sintomi elinig non indichino abbastanza chiaramente J, dell’occlusione, vi & Vanatomia patologica che insegna come, nei 4/; dei casi di occlusione, quests sia sita in sotto dell’ angolo splenico dal colon, e quindi in sotto di un taglio fatto nella = gione iliaca sinistra, e come nel rimanente quints dei casi loperazione lombare destra con qi completa certezza sia adeguata agli scopi che a prepone. Dimando quindi- Pud una regola di pre tica chirurgica , che ha sempre corrisposto a cura delle. ocelusioni eroniche del Tetto, essere modificata, per poter rispondere nei casi eecerion nali e non gid nei pil che frequenti, @ si dere q wodificare Ja pratica generalo chirurgica per tarla ai soli casi di eccezione / 4 La terza ragione a favore della colotomia iliag | @ che per questa via non é possibile I’ errore di capitare sul tenue intestino, sul duodeno o sulle: stomaco, e che le anomalie del colon non possono rappresentare una ragione di esito sfavyorevole | dell’operazione, poicht dalla incisione addominale | Si puod esplorare l'addome in tutte le direzioni. ~ Quanto al primo fatto, io credo che gli ert non sinno possibili,e, quando avvengono, sian da attribuirsi a mancanza di acenratezza nell’ & peratore, anziché all’ operazione in sé stessa. In ogni specie di attiviti, ¢ quindi anche in qué propria del chirurgo, sono questi non devono costituire fondamento di leggi speciali. Quanto poi alle anomalie del colon, di euti fautori della colotomia lombare gsi fanno arge mento validissimo , ed intorno alle quali 6 stat? tanto detto e scritto, id ho poco da dire. Am metto che le anomalie in parola possano oceor rere, ma tanto col metodo iliaco quanto col lom bare le probabilita di imbattersi in queste anomalig sono le stesse, ele difficoltaé Operative esisteran per ambo le Operazioni,y quantunque possano sere di forma differente. Ma dal punto di visit chirurgico l'anomalia di sede del colon 6 un fi / pit sfavorevole al metodo lombare che all’ ilie co ¢ Per mia esperienza devo rispondere di poiché in 170 casi, finora da me osservati, mi sone mai imbattuto in condizioni tali de durre serio ostacolo, e@ .:olto meno da Fé Yoperazione non favorovole nel risultato. Tn sol caso dovetti prolungare Vincisione in 6¥™ ed andare ad afferrare il colon al periore della pelvi , aprendg suturai i ini possibili errori, @ ee ee ee —— i quali come ultima ratio sono indispen- sabili. E, per adoperarli razionalmente, oc- corre che, insieme con le cause predispo- nenti, si ricerchino in ogni singolo caso le cause immediatamente occasionali dell’ a- gripnia ed a questa si conformi la terapia ipnotica individuale. La causa dell’agripnia é l’eretismo ce- _. Yebrale, in quanto che I’ organo psichico __ Soggiace all’azione consecutiva degli sti- ' moli del lavoro del giorno. Per calmare ' questo eretismo, oltre ad una giusta die- __ tetiea, al riposo del. cervello, alla calma campesire, é da aspettare buono effetto dal bromo, dalla piscidia, dal solfonale. dalla valeriana, dai bagni tiepidi usati la sera. 1 semicupi,; e in generale i procedimenti idropatici con bassi gradi di calore, rie- scono piuttosto eccitanti che calmanti e “non sono da raccomandare che come cura eonsecutiva. Se sono in giuoco processi cmotivi nella eoscienza, non si pud aspettar sussidio che da una terapia psichica x