J. PROTOZOOL. 9 ( 3 ) , 307-324 (1962). 307

Taxonomic-Nomenclatural Practices in Protozoology and the New International Code of Zoological Nomenclature*

JOHN 0. CORLISSt

Department o f Zoology, Eniversity of Illinois, Urbana, Illinois

SYXOPSIS. All zoologists are affected by provisions in the very recently published International Code of Zoological Nomen- clature, the first revised edition of these important rules to ap- pear in over 50 years. Common nomenclatural practices, often malpractices, of protozoologists and parasitologists who work primarily in taxonomic fields are revealed and discussed in light of recommendations and mandatory regulations to be found in the new Code. Some errors have been due solely to careless- ness ; others have involved misinterpretations of various direc- tives; still others have involved cases not adequately covered by the old R6gles. Certain mistakes of the past cannot be changed; but others are to be rectified upon discovery, accord- ing to mandates in articles of the new Code. Practical applica- tions of the rules of nomenclature are stressed, and examples are taken from actual situations found to exist throughout all major taxa of the phylum Protozoa.

Because of the value of such discussion in both new and re-

ROTOZOOLOGISTS working in widely differing P areas of research-biochemical, ultrastructural, parasitological, morphological, physiological, ecologi- cal, genetic, taxonomic-are all affected by provisions of the “International Code of Zoological Nomenclature Adopted by the XV International Congress of Zoology, London, July 1958”(16), the new set of rules which has very recently [November, 19611 come into force. Nevertheless, it is obviously the systematist whose work is most affected; and investigators in other fields will continue to rely heavily on his nomenclatural con- clusions. It seems a most appropriate time to consider some of the current or usual practices of protozoolo- gists and parasitologists in light of the appearance of this new, comprehensive set of regulations, essentially the first revision of the Code in over 50 years. But in point of fact a number of the provisions of the old Rkgles-changed or unchanged in the new Code-have been widely ignored, neglected, or misinterpreted in many publications of the 20th century.

The objectives of the present paper do not include a critical review or a complete summary of the Code;l rather. I should like to treat briefly but carefully some seven or eight major topics primarily concerned with common nomenclatural malpractices in fields of proto- zoology, with mention in paraphrased style of appro- priate provisions of the new rules. Such discussion of

* The support of National Science Foundation grant No. 10666 is gratefully acknowledged.

t This paper was prepared while the author was Visiting Professor of Zoology at the University of Exeter, Devonshire, England, during the academic year 1961-62.

visory work in protozoan systematics, the following major topics are given special consideration: matters of orthography. the original spelling of names and their justified or unjustified emendation ; authorships and dates of names, who is responsible and when, and how such data are properly cited; mandatory dates in the new Code, and their effect on both already estab- lished names and names not yet proposed; the principles of priority and conservation or continuity, and how the rules at- tempt to satisfy proponents of both of these diametrically op- posed “laws” ; the concepts of synonymy and homonymy, and proper methods of treating names which have become involved in such situations; family-group names, and the several special nomenclatural problems they present to protozoan taxonomists ; the major problem of types, and the peculiar position of pro- tozoologists with regard to the type concept, especially type- specimens for categories in the species-group ; miscellaneous con- siderations, several unrelated but significant topics not appro- priate for inclusion in preceding sections of the paper.

these neglected but important topics, available no- where in the literature, is germane to both new and revisory taxonomic work and thus warrants wide at- tention.

Errors or bad habits of a nomenclatural nature. often shown by esteemed workers who otherwise are systematists of recognized renown, may stem from four basic causes: ignorance of the rules, carelessness in their application, lack of clarity in the rules themselves, or total lack of a pertinent directive anywhere in the rules. The first two are quite often inexcusable. With regard to the second two, fortunately the new Code (16), in my opinion, has amply corrected the exasper- ating shortcomings of the old Rhgles, copies of which are easily available in several works( 14,18.22,30). Understandably, this has invoIved an increase in num- ber of articles (from 36 to 87) and thus in amount of reading matter. The clarity and precision of the Code. however, prevent its being considered a formidable. over-long, specialized treatise. Its Editorial Committee (N. R. Stoll, chairman) deserves the lasting gratitude of zoologists everywhere for finally producinq such a useful and usable document, worth its wait in gold.

‘The new Code is available, for one English pound or three American dollars, from the Internatlonal Trust for Zoological Nomenclature, 19 Belgrave Square, London, S.W. 1, England. This compact, hard-bound 176-page (+ i-xvii) publication con- tains important introductory material, appendices, glossary, and index, as well as the Code proper (Articles 1 through 8 7 ) . The only official language of the former Rhgles was French; the present rules are given simultaneously (printed side by side in facing pages) in French and English, and the texts in both languages are to be considered “equivalent in force, meaning, and authority.” The effective date of the Code is 6 Sovember 1961.

305 SOJIEKCLATCRAT, PRACTICES ASD THE YEW CODE

MATTERS OF ORTHOGRAPHY One of the most controversial areas in nomenclatural

systematic protozoology has been that concerned with the original spelling of names and ‘or their subsequent emendation. In the neky Code, Article 32 states very clearly that the oriqinal spelling is to be retained as correct except under special circumstances. given in a short list. However, if a name is discovered to be in- correct under thrsc circumstanccs, then it “is to be corrected wherever it is found.”

What are the circumstances under which correction is not only allowed hut is mandatory? These are con- sidered briefly in the paragraphs immediately fol- lowing.

lf the original spelling contravenes a mandatory pro- vision of Articles 26-31. These are simple. straight- forward rules, all of which need not be treated here. But several are particularly apposite. since they offer solutions to situations which have occurred quite fre- quently in naming protozoa.

If a specific name was based on a compound name published as two separate words by the original de- scriber these words are to be united without a hyphen in subsequent publications. Curiously enough. in- stances of total disregard for the ancient and basic principle of binominal nomenclature still occur in the protozoological literature. even within the past decade or two.

Diacritical or other similar marks are no longer al- lowed in a zoological name. In general. these are just to be dropped in correction. leaving the vowel. or con- aonant, in a purely latinized form. For example Hop- litophrya gkorgkvitchi becomes Hoplitophrya george- vitchi; and Juxtaradiophyra nazsensis, Juxtaradiophyra naisensis. But the German “ u ” with an umlaut-u- must be replaced by “ue”. This affects a number of name? in the protozoological literature; for instance, all those a t familial, generic. or specific levels com- posed in honor of Butschli. Lieberkuhn, and Muller. For example. Butschliidae automatically becomes. from the date of the new Code onward. Buetschliidae: Spirobiitschliella, even though often written (incor- rectly) “Spirobzitschliella,” becomes Spirobuetschli- ella; Iodamoeba butschlii, Iodamoeba buetschlii; Stera- tor miilleri, Stentor muelleri.

Family-group names must be formed by addition to the “stem” (not, strictly speaking, the actual gram- matical stem but the genztive without i ts case-ending) of the name of the type-genus the endings -idae for family. -inae for subfamily. It is considered a “justi- fied emendation” to repair a familial name not properly so formed. Thus a number of protozoological families are affected. For example. Astylozoonidae becomes Astylozoidae: Blepharocoridae, Blepharocorythidae; Condylostomidae. Condylostomatidae; Epiplocylidae,

Epiplocylididae ; Epistylidae. Epistylididae: Pycno- thricidae, Pycnotrichidae; Trypanosomidae, Trypano- somatidae. However: it is important to note that if the generic name involved is a Greek word latinized with a change in termination then the stem is that appropriate for the latinized form. Thus, although Trypanosomidae should be Trypanosomatidae (be- cause the type-genus is Trypanosoma), Spirostomidae was recently incorrectly emended ( 6 ) to Spirostomati- dae (because in this case the type-genus is Spirosto- muin not Spirostoma). This whole subject is consid- ered in more detail in a subsequent section of the present paper (see p. 317).

Species-group names must, of course. agree in gender with the generic name if they are in the form of adjec- tives in the nominative singular. Thus when the myxo- sporidian species Cystodiscus immersus was transferred to the genus Myxid ium, for example, its new name rightly became Myxid ium immersuna. Information is provided in the Code to aid in determination of the gender of genus-group names. If species-group names are formed from modern personal names there are prescribed endings: the genitive termination of -i if the personal name is that of a man, -ae if of a woman, and -orum or -arum for the appropriate plurals.

lf a clearly inadvertent error or printer’s mistake occurred in the original publication. At last this man- date of the rules has been clarified. A lapsus calami is rightly considered an “inadvertent error”; and it is permissable for a later worker to correct it. But not included in such a category are incorrect trans- literation, improper latinization, or use of an inappro- priate connecting vowel. Names involving the latter errors are to be left in the original spelling.

Thus Dobell’s( 10) wrathful attack on ‘ (Teranympha and other monstrous Latin parasites,” valuable though his paper was and still may be as a guide to persons who are going to be forming new names, carries no authoritative nomenclatural weight as regards names already published. Indeed, Cleveland( Z ) , whose quite innocent remark of doubt as to Koidzumi’s legalistic right to change Teranympha to Teratonympha occa- sior.ed Dobell’s vitriolic outburst, is vindicated: by the rules ‘(Teratonympha,” Dobell’s insistent emendation, is not valid. Furthermore, the familial name Terato- nymphidae should revert to Teranymphidae.

Kirby( 19)’ writing in a much milder vein than his contemporary Dobell, was concerned about the fact that strict application of priority in orthography, good or bad, might oblige protozoologists to change what had then become and still is customary spelling in the case of a number of important genera. He pointed out that Amoeba was originally spelled Amiba; Chlamy- domonas, Chlamidomonas; Condylostoma, Kondylio- stoma; Strombidium, Strombidion; Trichomonas, Tri- conzonas. Other examples could be added to this list.

NOMENCLATURAL PRACTICES AND THE N E W CODE 309

However. I believe that the new 50-year “statute of limitation” regulation (Art. 23; and see pp. 311, 313 of the present paper), very broadly interpreted, might be employed to take care of nearly all such cases: for the emended spellings, though unjustified, have the status of junior objective synonyms (Art. 33) and are thus available as valid names should the unused senior synonyms (the names in their original spelling) be considered as nomina oblita (forgotten names).

An example of a truly inadvertent error would be the following: if one stated his desire to name a species after a certain Dr. Willson and yet first published the name as X-us wilsoni. This must be emended to X-US willsoni. And no one will quarrel with the desirability of correcting errors obviously of a typographical na- ture: e.g., if Actinopkrys had first been printed, in- advertently, as Actinophyrs.

If a family-group name has an incorrect sufix or has been improperly formed from the name of its type- genus. There are three possible instances under this heading.

The suffix naturally has to be changed if the rank of the taxon is changed from family- to subfamily-level or vice versa. E.g., if Biitschli’s hypotrich subfamily Urostylinae should be elevated to familial level the spelling of the name obviously would have to be emended to Urostylidae.

If the name has had an incorrect ending in the older literature it is also to be automatically repaired. To give several examples from the many that exist in the protozoological literature: Ehrenberg’s families CoI- podea, Enchelia, Vorticellina, and Oxytrichina are properly written today as Colpodidae, Enchelyidae, Vorticellidae, and Oxytrichidae; Dujardin’s Bursari- ens, Parameciens, and Urceolariens become Bursari- idae, Parameciidae, and Urceolariidae ; Stein’s Ophry- oscolecina becomes Ophroscolecidae. Note that these repaired names are still to be cited with their original authorship and date.

Finally, a family-group name must be emended if the stem of the name of the type-genus has been in- correctly formed. This important matter, covered by Article 29 of the Code, has already been mentioned briefly above (p. 308) and is to be considered again later (see p. 317).

Recommendations concerned with spelling of names. In addition to mandatory provisions relevant to ortho- graphical matters there are a number of recommenda- tions, both in the Code proper and in its Appendices, which deal specifically with proper proceedings in for- mation of names. But it should be emphasized that these Recommendations do not have the weight of the mandates which comprise the definitive Articles them- selves. Names already formed, even though etymo- logically incorrect, must be allowed to stand (with the few exceptions already briefly discussed above).

For consideration by workers who are going to be forming new names in the future, it may be worthwhile to mention one or two of the recommendations (from Appendix D of the Code) regarding a category of names commonly used by protozoologists in describing new genera or species. In forming a species-group name from the name of a modern man that is not Latin, latinized, or of Greek origin, it is recommended that a single -i should be added to the entire name rather than the double -ii (that is, an ‘5” onto a theo- retical genitive “stem” which is considered to termi- nate in an “i” itself) so frequently employed in the older literature. Thus smithi rather than smithii (from Smith), steini rather than steinii (from Stein). But buetschlii (not Biitschlii: see p. 308) would still be cor- rect (from Biitschli). As I understand it, however, names once formed with -ii should not be changed; thus Colpoda steinii should remain so, but a new spe- cies named for Stein in another genus preferably would be X-us steini.

I t is recommended that in forming generic names from compound personal names only one of the com- ponents, the better known, should be used. Also pre- fixes should either be omitted, as in the case of a nobiliary particle or one indicating Christian sainthood or miscellaneous others, or combined with the name proper. Some examples involving names of protozoolo- gists: cunhai, for da Cunha; faurei, for FaurC-Fremiet ; geleii, for von Gelei; puytoraci, for de Puytorac; but macdonaldi, for McDonald; and demorgani, for De Morgan.

Status of emendations. To return to the subject of proper emendation of names originally misspelled, it should be noted that the date and authorship of the original name are retained. This is as true for family- group names as for generic and specific names. Accord- ing to Article 19, an emendation, whether justified or unjustified, is an available name; but an incorrect spelling, whether it appears in the original or subse- quent works, has no standing in nomenclature, is not an available name, and thus may not later be consid- ered as a synonym. Such misspelled names, however, may be listed in synonymies for bibliographical and historical reasons, as was done with generic names in my recent book on classification of ciliates(6).

A justified emendation is credited to the original proposer of a name, as mentioned above, not to the reviser. But an unjustified emendation falls as a junior objective synonym of the original name, and it is to be considered the responsibility-as shown by the au- thorship and date-of the person who made it.

AUTHORSHIPS AND DATES OF NAMES

Articles 22 and 51, and parts of others, are concerned with the proper manner of citing the authorship and date of a zoological name. Although such citations are

310 SOMENCLATURAL PRACTICES A N D THE KEW CODE

optional, since they do not form a part of the official name of a taxon. it is recommended (Appendix E of the Code) that every zoological paper involving names of animals (generic or below) should include citation of the authors and dates “a t least once” in the publi- cation. Because there seems to be a considerable amount of confusion in the general protozoological and parasitological literature, the matter merits further discussion here.

The recommended form of citation for a f ami l y or genus may be seen in these examples: Trypanosomati- dae Doflein. 1901 : .-lrcella Ehrenberg, 1830. S o t e that no taxonomic conclusions are to be drawn from inspec- tion of these names. That is to say. the composition of such qroups may have been considerably altered by later workers: in fact, even the names may have been emencied (as in Trypanosomatidae) . But the original authors. with their dates, are nomenclaturally credited with--i.P., are to be held responsible for-the names of these particular taxa.‘

Two common erroneous modifications of the method of citation just illustrated are these: ( 1 ) the author’s name and the date are regularly thrown into parenthe- ses: e.g., Trypanosomatidae (Doflein. 1301) : ;1rcella (Ehrenberg, 1830). ( 2 ) The names of the taxa are unreasonably credited to the author of whatever text- book or similar source in which the writer has appar- ently last seen them: c g . , Trypanosomatidae IYenyon: ilrcella Kudo!

The recommended form of citation for a species whose nomenclatural history is ar, uncomplicated one is identical to the examples given above. E.g., .-lrcella rwlgaris Ehrenberg. 1830: Selenidiunz potamillae JIac- kinnon & Ray. 1933. Incorrect ways of making these citations abound in the literature. IVhen a species has been transferred to another genus, there are several proper ways to give the reader information concerning the authors and dates involved. There are even more improper ways! Let us consider the widely studied species Tetrahymena pyriformis as an example, assum- ing the following historical data to be factual (see

* The Code should not be-and is not-concerned with mat- ters of taxonomic judgment. I t is perhaps unfortunate, how- ever, that no uniform way is suggested for citing the name of persons who have carried out major taxonomic revisions of a given group: drastically restricting, expanding or revising the characterization of the taxon or making a major curtailment or increase in the number of lower taxa (e.g., genera in a family) included in it. -4 style commonly employed-and it seems a sensible one-is to append the word “emendation” (customarily abbreviated to “emend.” or ‘‘em.”) followed by the name and date o i the ivorker responsible for the revision. Thus Arcella Ehrenberg, 1S30. might justifiably be written in further detail 2s: i l r r ~ l l a Ehrbg.. 1830. emend. Deflandre, 1928. In cases o i familial names perhaps protozoologists and parasitologists mould not be so hesitant to use the name of the original author if they felt more free to mention also the name of the most im- portant or most recent subsequent reviser of the concept of the taxonomic group when writing out the full name of that group.

Corliss & Dougherty(9) for details in this complicated case).

Ehrenberg first described the form in question, in 1830. as Leucophrys pyriformis. Lwoff transferred it to the genus Tetrahymena, in 1947. [Actually Maupas had transferred it to the genus Glaucoma, in 1883; but. any time after 1947, Maupas cannot be cited as an author: ix., his name cannot follow the binomen Tetrahymena pyriformis, since he was neither the origi- nal author nor the most recent reviser.] The manner of citing the name depends on the amount of informa- tion a writer wishes to give his readers. Any of the fol- lowing ways are correct:

Tetrnhymena pyriformis T. pyriformis [if generic name has already

Tetrahymena pyriformis (Ehrenberg) Tctrahymena pyriformis (Ehrenberg, 1830) Tetrahymena pyriformis (Ehrenberg) Lwoff Tctrahymena pyriformis (Ehrenberg, 1830)

been given in full]

Lwoff, 1947

once

Among the many incorrect ways the following are given because they have their frequent parallels‘in ac- tual protozoological literature (the names of the au- thors may or may not be accompanied by their respec- tive dates-examples will not be shown both ways in this particular list) :

pyri f ormis Tctrahymena pyriformis Ehrenberg Tetrahymena pyriformis Lwoff Tetrahymena pyriformis (Lwoff) Tetrahymena pyriformis (Lwoff ) Ehrenberg Tetrahymena pyriformis Lwoff (Ehrenberg) Tetrahymena pyriformis (Ehrenberg, Lwoff) Tetrahymena (Leucophrys) pyriformis (Ehren-

The problem posed by the last-cited example will be treated again in a subsequent section of the present paper (see p. 314) : the placement of a synonymous generic name in the location reserved by law (Art. 6 ) for the name of a subgenus is, unfortunately, a wide- spread habit among parasitologists and protozoologists. Leucophrys is not a subgenus of Tetrahymena. The above list of incorrect citations of authors would be greatly extended by addition of variants involving Maupas’ placement of this species in the genus Glau- coma; as already mentioned above, Maupas’ name cannot appear after the specific name in its most re- cent combination.

If the scientific name of a taxon is the responsibility of someone who is not the author of the publication first containing the name, then one is required to be- stow the credit where i t properly belongs (assuming the identity of the namer is known). This holds for any name a t the family-level or below (and, by logical ex- tension. should be used even a t higher taxonomic levels,

berg) Lwoff

NOMENCLATURAL PRACTICES AND THE NEW CODE 311

in my opinion). As an example, the names of five fami- lies of ciliates erected by KahI appeared first, accom- panied by short diagnoses, in the 5th edition of Doflein & Reichenow’s great protozoological treatise. These names should thus be credited as follows, using one of them in illustration : Conchophthiridae KahI in Doflein & Reichenow, 1929. Curiously enough, Kahl also pub- lished these names a t a later date, still as “new,” in publications of his own (see 6) ; this observation, how- ever, is not pertinent here (but see p. 322 of the pres- ent paper).

If an author’s name is very well known it may be abbreviated, even reducing it to initials, when it is used after a taxonomic name. All zoologists recognize “L.” as standing for Linnaeus, by tradition. In proto- zoology, “Ehr.,” “Ehrb.,” or “Ehrbg.,” for Ehrenberg, and “Duj.,” for Dujardin, are among a number of com- monly recognized abbreviations. “O.F.M.,” for 0. F. Miiller (not to be confused with J. Miiller of a later century or with a number of still more recent Mullers), has become a classic set of initials requiring no ex- planation.3

MANDATORY DATES IN THE CODE A number of provisions in the new Code refer to

exact dates or specified periods of time of nomencla- tural importance. Some of these, also present in the old Rkgles, have been rather routinely ignored, not alone by protozoologists? The dates concerned may be mentioned briefly, more or less in their chronological order.

(1) Year 1758. This is the “year zero” for the names of animals (Art. 3 ) : the starting point of zoo- logical nomenclature. Any names published prior to 1 January 1758 cannot be recognized as from those dates, although a number of protozoologists persist- ently cite Paramecium, for example, as “Paramecium Hill, 1752.” ( I t should be: Paramecium O.F.M.. 1 773.4)

(2) Statute of limitation. If a senior synonym has gone practically unnoticed for more than 50 years (Art. 23) it may be relegated to the position of a nomen oblitum, with conservation of the junior syno- nym which has been in general usage. Discussion of this important “Law of Continuity” (or “Law of Con- servation”) is reserved for a subsequent section of the present paper (see p. 313).

(3) Range of dates. There are a number of provi- sions in the Code (Art. 2 1 ) referring to the determina-

Otto Friderich is apt to be remembered by students as “Old Fellow” Miiller, an epithet not be taken at all disrespectfully. .4n up-to-date assessment of his contributions to protozoology, incidentally, is badly needed. ‘ Miiller actually spelled this generic name “Paramaecium,”

hut there has been quite general and widespread agreement for many years among taxonomic protozoologists, from Ehrenberg onward, that the “ae” be reduced to “e.”

tion or interpretation of the exact date of a taxonomic name, an important consideration in cases of alleged priority, of course. One in particular concerns works which are given a range of dates as their time of pub- lication ( e g . , “1880-1882,” “1927-1929”). If separate parts of such a work were not actually published at different known times then the date to be used is the latest one in the range given.

(4) Date of replacement names involving the type- genus of a family. If the generic name on which a family-group name is based is found to be a junior homonym (Art. 39), the replacement names for the genus, and for the family as well, are given the dates of the preoccupied invalid names. Although the gen- eral question of homonymy is to be considered in a subsequent section of the present paper (see p. 315), this specific matter can be clarified here with an actual example in protozoology( 5,6) ; the prescribed form of writing the new names with authors and dates can also be shown:

Epalxis ROUX, 1899, a junior homonym of Epalxis Cossmann, 1899, has been replaced by Epalxella Cor- liss, 1960. The familial name Epalxidae [always erro- neously spelled Epalcidae] Wetzel, 1928, automatically became Epalxellidae Corliss, 1960. But in order to preserve the family concept historically-i.e., to pre- vent take-over (only hypothetically possible in this particular case) by any subfamilial groups bearing dates earlier than 1960-the new Code (Arts. 39. 40) requires that the replacement names bear both dates. Thus the new names should henceforth be written as: Epalxella Corliss, 1960 (1899) ; and Epalxellidae Cor- k, 1960 (1928).

( 5 ) Pre-1900 familial names. If family-group names published before 1900 in vernacular form otherwise meet the requirements of the rules, have been fully latinized by subsequent authors, and have been gener- ally accepted by specialists of the group, they are to be considered available with original dates and author- ships (Art. 11). For example, family Bursariens of Dujardin should be written Bursariidae Dujardin, 1841 ; Family Ophryoscolecina of Stein, Ophryosco- lecidae Stein, 1859.

( 6 ) Generic names published before 1931. Such names are acceptable even if published without asso- ciated nominal species (Art. l l ) , if they meet certain other general requirements and were accompanied by a description, definition, or indication (Arts. 12, 16).

( 7 ) Generic names published after 1930. Genus- group names published after 1930 must conform more stringently to the several provisions of Articles 11-13. I t is especially to be noted that such a name must be accompanied by the definite designation of one of the included species as the type-species. There has been considerable laxity in publications of the past 30 years with regard to this mandatory provision: strictly

312 SOJIESCLATL-RAL PKACTICES 4x11 THE SEW ConE

speaking, genera so improperly erected are to be re- jected. although to date this drastic step has not been taken in the protozoological literature. to my knowl- edge.

( 8 ) Scw nainec puhlished anonymously after 1950. According to provisions of Articles 9. 14. and 17, taso- nomic names published anonymously after 1950 are not available: nor do anonymous papers constitute official publication, nithin the meaning of the Code. after that date. A name is or remains available, even though anonymous. if published before 195 1 .

(9 ) Importance of thr dates 1960-61. These very recent dates have mandatory significance in several different situations, each of uhich needs to be men- tioned separately.

( i ) Conditional naiizr5. After 1960. a new name pro- posed conditionally is not available (Art. 15) ; but a name SO iiroposed before 1961 cannot be rejected on such a basis.

( i i ) “I’aric’ties” and “lormap.” -\liter 1960. names propoced explicitly as xrarieties or forms. generally preceded bv “var.” or “forma” when used in the pro- tozoological literature. are not available (*Art. 15). That is. wch a name is to be ofticially regarded as of infrasubspccific rank. therefore outside the jurisdiction of the Code (Arts. 1, 4 5 ) . Thus it should be empha- sized here that ii a name first established with infra- subspecific rank is subsequently elevated to the level of a subspecies. or species. it takes as its authorship and date those of its elcration (.\rt. 10) . Varieties and forms of the past literature (i.e., before 1961). how- ever. are to be construed as of subspecific rank (Art. 45) . I t would be well if protozoologists adopted such a consistent policy with regard to these names: many “var’s” exist amonq the phytoflagellates (understand- ably, since phycologists recognize the rank”) : and “formae” are to be found scattered throughout various groups, c . g . . among the entodiniomorphid ciliates.

(iii ) Family-group names based on junior synonynzs. After 1960, a familial name is not to be changed if the name of the type-genus on which it is based falls as a junior synonym (A4rt. 40) . Xote that this does not in- clude cases involving junior homonyms (see p. 318). But if, before 1961. a family-group name was changed because of such synonymy-that is, a new name was formed to agree with the replacement name (valid senior synonym) of the type-genus-it is to be main- tained in the interests of stability if it has “won gen- eral acceptance.” I n such cases, the date of the newer familial name is to be considered that of the rejected familial name. In any possible subsequent arguments

,’ In fact. the International Code of Botanical Nomenclature ( 1 5 ) recognizes, in descending order of rank: subspecies, varie- ties and subvarieties, forms and subforms. But trinominal names (ternary combinations) can be used only if the rank of the third epithet, the infraspecific taxon, is stated directi? i n the name

of priority the conserved name would be considered the senior synonym, The recommended manner of indi- cating the dates is the same as that suggested for the somewhat parallel situation discussed under section 4, above. Two pre-1961 examples may be cited in illus- tration of application of Article 40:

The familial name Trimyemidae was proposed by Kahl. in 1933, as replacement for Sciadostom(at)idae Kahl. 1926, upon his discovery that Sciadostoma Kahl, 1926. was obliged to fall as a junior synonym of Trirnyema Lackey, 1925. Had all this occurred after 1960, Sciadostomatidae would have to be considered the only valid name for the family concerned. But since the change was made before 1961, and has won general acceptance, the replacement name Trimyemi- dae is to be considered the valid one. It should be written with authorship and dates as follows: Tri- myemidae Kahl, 1933 (1926).

The second example involves retention of a familial name based on a type-generic name which has fallen as a junior synonym. This case is also pre-1961, bu t the familial name based on the senior synonym has never won general acceptance. In 1913, Poche erected the Caenomorphidae to replace Gyrocorythidae [spelled Gyrocoridae by Poche] Stein, 1867, because Gyrocorys Stein. 1860. was recognized as a junior synonym of Cacnoinorpha Perty, 1852. Recognition of Gyrocory- thidae is supported by the Code; and i t is to be found in the literature more commonly than Caenomorphi- dae. As a familial name it already stood as a senior synonym, so no second date is needed after the name of the author. A complication involving the name Metopidae Kahl, 1927, with the family under con- sideration is treated briefly in a later section of the present paper (see footnote 9, p. 318).

(iv) Egec-tive date of the new Code. By the princi- pal provision of Article 84, the “International Code of Zoological Yomenclature adopted by the XV Interna- tional Congress of Zoology, London, July 1958” comes into force on the day of its publication, with all previ- ous editions of the Rules thereby being superseded. Cnfortunately, this all-important date was not able to be supplied anywhere in print in the Code itself; but it is now known to have been 6 November 1961 (26 ) .

PRIORITY AND CONSERVATION The new Code makes every effort to reconcile the

two seemingly diametrically opposed concepts: the “law” of priority, on the one hand, and the “law” of conservation or continuity, on the other. There is no doubt that the primary goal of the Code, to “promote stability and universality” (Preamble, 16), is generally better realized by recognizing the validity of the oldest available name for a given taxon than by endorsing wholesale conservation of more recently proposed names for such taxa. There are two major complica-

NOMENCLATURAL PRACTICES AND THE KEW CODE 3 13

tions, of course: how exactly to apply the Law of Prior- ity (Arts. 23-34); and when to invoke means of con- servation, either by direct use of certain provisions in the Code (see parts of Arts. 23, 40) or by calling on the International Commission on Zoological Nomen- clature to suspend otherwise mandatory provisions of the Code, in certain specific cases, through use of its plenary powers (Art. 79). A further potential compli- cation is that the rules of nomenclature are not to re- strict the “freedom of taxonomic thought or action” in any way (Preamble, 16).

As I stated in a work(6) published before the ap- pearance of the new Code, there are three general goals which may act as guides in nomenclatural aspects of taxonomy: stability (and thus universality), priority, and consistency. The second and third should be sub- servient to-and, when necessary, sacrificed for-the first of these. Discussion of a few protozoological ex- amples will illustrate application of pertinent provi- sions of the new Code and may be of aid not only to taxonomists who are involved in forming new names for new groups but also to investigators who are en- gaged in reviewing and revising taxonomic treatises of by-gone decades, work often enmeshed in unbelievably complicated nomenclatural tangles.

Article 23 is the Law of Priority in the new Code. By helpful coincidence (or was it so planned?) its num- ber is very close to the celebrated “Article 25” of the old rules( 14,18,30) on the same topic. Its exceptions, minor in nature save one, involve a number of subse- quent articles and are related to concepts of validity, availability, justified and unjustified orthographic emendations, and the like. Parts of Article 40 (related to preservation of certain family-group names, a sub- ject already discussed on p. 312 of the present paper) and Article 79 (on invoking the plenary powers of the Commission) are concerned in some measure with the matter of conserving names. But the section of Article 23 entitled “Limitation” is the major exception to the Law of Priority proper. Since it represents the heart of the “Law of Continuity,” it alone merits further particular discussion here.

The ‘150-yea~s” rule. A name that has remained in disuse as a senior synonym in the “primary zoological literature” for a period exceeding 50 years is to be con- sidered a nomen oblitum (forgotten name). Such an invalid name is not, therefore, to be used unless the International Commission on Zoological Nomenclature so directs. On the other hand, a worker discovering such a name is, after 1960, to refer it to the Commis- sion for placement either in the Official Index of Re- jected and Invalid Names or (if desirable) in the ap- propriate Official List of (acceptable) Names. Two ex- amples, of slightly differing nature, may be mentioned from the protozoological literature. These involve fa- milial names, a category of names also covered in a

subsequent section of Article 23: but, as I interpret the “statute of limitations,” such names are not offi- cially excluded from treatment under provisions of the “50-years” rule as well.

The well-known suctorian family Discophryidae Col- lin, 1912, is nominally a junior synonym of Rhyncheti- dae Kent, 1880. But the latter name has rarely been used since the turn of the century and would thus seem to have been well forgotten(6). There remains only the task of referring it to the Commission to establish it legally as a rejected name.

The gymnostome ciliate family Amphileptidae Biit- schli. 1889, should, by strict application of the prin- ciple of priority, fall as a junior synonym of Litonoti- dae Kent, 1880. With only one or two exceptions in relatively obscure publications, however, the familial name Litonotidae has not been resurrected; so the case is similar in this respect to that just cited for Rhyn- chetidae. In fact both of the senior synonyms in these two instances date from the same 19th century mono- graphic work, that of Kent.

The generic name involved in the second case- Litonotus-has, interestingly enough, also been in dis- use for a long time, essentially since Biitschli altered its spelling to Lionotus in 1889. Lionotus has been, and still is, in such universal usage that it might seem to warrant similar appeal to the statute of limitation. However, “Lionotus” was Butschli’s unjustified emen- dation of the original spelling of Wrzesniowski’s ge- neric name Litonotus; furthermore, Biitschli’s Liono- tus has to be rejected, anyway, because it is a junior homonym of Lionotus Agassiz, 1846, coleopteran, and Lionotus Thomson, 1870, hymenopteran (23). Thus we must become accustomed to Litonotus as the legiti- mate name in place of the currently more familiar, but incorrect, Lionotus; or we must petition the Commis- sion either to conserve Butschli’s name, in spite of the counts against it, or to reject both names-in the lat- ter case someone would have to propose an altogether new name for the group. Adoption of Litonotus has seemed the simplest and most sensible solution to me (5,6) ; and its close similarity to the word “Lionotus” should preclude its causing much inconvenience to the taxonomic specialists on this and related genera of gymnostome ciliates.

The examples of family-group names just cited per- haps better fall under a different section of Article 23, one which permits petition to the Commission for con- servation of a junior synonym as name of a family formed by union of two or more families, provided that “general usage” would be upset by strict applica- tion of the law of priority. I have followed the spirit of this provision of the Code, before it was published, in a dozen cases in my work on the systematics of the ciliated protozoa (6) . All that remains to be done in these cases is preparation of the proper request to the

314 x0MEKCLATURAL PRACTICES .4KD THE KEW CODE

Commission for acceptance of the names in the Official List of Family-Group Names in Zoology.

Preservation of relatively recent names. ,4n impor- tant subsection of the ‘‘statue of limitation” provision of Article 23 stresses that i t does not preclude conser- vation of names, important in applied zoology, of which the period of widely acceptable usage has been less than 50 years. ’4pparently the Commission may be petitioned in cases of both junior synonymy and jun- ior homonymy.

An established example in protozoology of preserva- tion of a junior homongm is the now familiar one of the spirotrich ciliate Stentor (see 13). This generic name, when used for a protozoan tason, had been pre- occupied by the little-used (available but invalid) name of a genus of monkeys. .A major example of a junior synonyin may be found in the complicated case of the hymenostome holotrich ciliate Tetrahymena. With the very recent but long anticipated appearance of the new Code. Dr. Dougherty and I have further delayed final submission of our necessarily lengthy petition(9) to the Commission until we can redraft it in light of certain provisions in the new rules. Over a dozen generic names are involved in this particular case; but Tetrahymena Furpason. 1940. is obviously the name to be preserved. since i t has now become so thoroughly entrenched in the protozoological and bio- chemical literature of the past 21 years(6. and perti- nent references therein).

There are other well-known names, some ancient, some quite recent. whose existence is threatened today from some quarters. One hopes that their cause will be taken u p in time. To mention just one example. Amoeba: see the latest discussion by Kudo(21), in a paper favoring retention of this time-honored name. Here again i t should be remembered that strictly no- menclatural matters can be-and generally should be -divorced from the broader taxonomic considerations of interrelationships. etc.. judcments generally based on morphological data. The valid name to be applied can either be determined by strict application of perti- nent provisions of the Code or be established contrary to these same provisions by successful petition to the Commission for their suspension, under the Comniis- sion’s plenary powers, in a given particular case.

ST?;OSI’JI’S’ AND HO?tIOSTMT Synonymous names. The existence of synonyms,

two or more different names for the same recognized zoological taxon. is common in the protozoological literature. Such cases are often inevitable and many have arisen legitimately ; although also innumerable instances could have been avoided entirely had investi- gators been more conscientious in acquainting them- selves with data available in the older literature.

-4 number of scattered provisions in the new Code

treat problems of synonymous names; particularly, of course, Article 23 (Law of Priority) and subsequent articles in that section of the Code. There are gen- erally no complications in application of such direc- tives. Recall that incorrect subsequent spellings of names are not to be treated as synonyms. -4 small but very important problem arises in con-

nection with the manner of citing a synonymous ge- neric name when giving the full name of a species. As recently mentioned briefly elsewhere( 5 - 7 ) , many pro- tozoologists and parasitologists have the unfortunate habit of writing names in the style illustrated in the following examples when they wish to indicate the, or a. former generic vehicle of the species in question:

Colpidium (Paramecium) colpoda Diophrys (Euplotes) appendiculata Einzeria (Coccidiurn) schubergi Haemamoeba (Plasmodium) gallinacea Hepatocystis (Plasmodium) kochi Histomonas (Amoeba) meleagridis Laaerania (Plasmodium) f alcipara Phacus (Euglena) pleuronectes Tetrakymena (Glaucoma) pyriformis Triclaomonas (Cercomonas) horninis Trypanosoma (Herpetomonas j lewisi

This is done indiscriminately, whether all the species of the former genus now belong in the genus of the new combination (i.e., the one generic name has fallen as a junior synonym of the other) or only one or more ( i . e . , the synonymy is only pro parte at the generic level). It is a distressing and misleading habit because the names, as written above, indicate that the species in question all have as subgenera the names enclosed in parentheses (Art. 6) ! Yet these “subgenera” may be, and often in actuality are, perfectly good, inde- pcndcnt, full genera in their own right, containing their own quite separate species, and possibly belonging to entirely different families or even different orders (e.g., Histomonas and . 4 m o ~ b a ) ,

The Code. unfortunately, provides no official way of writing the full name of a transferred species which in- cludes indication of the former generic name, although a recommendation in Article 44 condemns placing i t in any form between the generic and specific elements of a binomen. 1-et, admittedly, it is often most helpful to be given such information concisely, especially when most of the past literature may have dealt with the or- ganism in question under the name which has now fallen as a junior synonym. As far as the Code is con- cerned. the manner of indicating the former generic name is left to the ingenuity of the investigator, as lonq as he does not usurp the place of the subgeneric name-as was deliberately done in the “bad” examples given in the list above. Whether or not any subgenus happens to exist for any particular genus under con- sideration is beside the point.

NOMENCLATURAL PRACTICES AND THE NEW CODE 315

Here are some potentially acceptable possibilities for citation of the species given as “Laverania (Plas- modium) falcipara” in the preceding list:

Laverania (= Plasmodium) falcipara Lavernnia [Plasmodium] falcipara Laverania (“Plasmodium”) falcipara Laverania (syn. Plasmodium) falcipara Laverania (formerly Plasmodium) f alcipara Laverania falcipara (= Plasmodium falciparum) Laverania falcipara (syn. Plasmodium falciparum)

The shortest style shown, with the square brackets in place of parentheses, might be considered too danger- ously close to the required method for indicating a subgeneric name. Certainly neither i t nor any other among the first five examples adheres to the spirit of the recommendation in Article 44, mentioned above. I understand that a common form in entomology would have it read “Laverania falcipara [Plasmodium]”, a possibility meriting consideration by protozoologists. The longest way, used in the name given last in the list, is the safest; but, admittedly, it is not very con- cise. However, as Keen & Muller( 18) stress, in treat- ing this general subject, “clarity should not . . . be sacrificed for the sake of brevity.”

Treatment of homonyms which happen also to be synonyms is reserved for the following section.

Homonymous names. An entire section of the new Code, embracing Articles 52-60, is devoted to homony- my, the occurrence of the same ( ie . , orthographically identical) names for different zoological taxa (familial level and below). The Law of Homonymy (Art. 53) clearly states that any name which is a junior homo- nym of an available name (this includes even those cases in which the senior homonym has itself become a junior synonym elsewhere in the animal kingdom) must be rejected and replaced. Most of the subsequent articles in the section are concerned with minor points and refinements, clarifying situations in which there might be confusion over the existence of an homony- mous condition or not. A few examples follow.

Generic names are not to be considered homonyms if the difference is even as slight as only one letter (Art. 56). The now classical example of this in the parasitological-protozoological literature is the cele- brated Endamoeba-Entamoeba case, in which the “in- famous” Opinion 99, published in 1928, was finally superseded by the Commission’s publication, in 1954, of Opinion 312, validating both names for use in ref- erence to separate genera of parasitic amoebae. An- other case is that of Kahl’s( 17 ) unnecessary replace- ment of Spathidiodes by Spathidiella because he con- sidered the former name to fall as a junior homonym of a separate earlier generic name Spathidioides.

Variable spelling at the species-level (Art. 58) , how- ever, does not prevent homonymy if it involves certain letters or pairs of letters. For example, if one were to

describe a new species of Stentor as Stentor ceruleus, that name would fall straightaway as a junior homo- nym of S. coeruleus. Two species of any genus which might bear the specific names of litoralis and littoralis would similarly be subject to the Law of Homonym; and smithi, smithii, even if different gentlemen were being honored by such patronymics.

Although Article 53 insists on the rejection of a jun- ior homonym, it is not until Article 60 that one is in- structed as to how to replace it, other than to petition the Commission to use its plenary powers to conserve the name (such as was done for the ciliate genus Sten- tor: see p. 314 above). There are two sources of re- placements:

(1) Use of an existing available name. If the jun- ior homonym has any available synonyms, junior or senior0, the oldest must be adopted as the replacement name. In effect, then, no attention a t all need be given to junior homonyms already being treated as junior synonyms. If the junior homonym is a senior syno- nym first choice for its replacement name must be the next oldest synonym, used with its own authorship and date. An exception is made for names of families and their type-genera when the latter have been found to be junior homonyms (Art. 39) : the new family-group name, to be formed only from the (new) valid name of the type-genus (at least, after 1960), and the new generic name take the dates of the replaced names.

Examples of the use of either kind of synonym as replacement names for nominally preoccupied proto- zoan genera are abundant; a few have been selected from the Ciliophora( 5) for illustration here. Junior homonyms falling as junior synonyms of valid names include such exampIes as these: Drepanostoma, of Loxodes; Metopides, of Metopus; Oospira, of Gymno- dinioides; Triloba, of Teutophrys; Turbinella, of Uro- centrum. Junior homonyms replaced by junior syno- nyms include: Arachnidium, by Strobilidium; ‘4 ulax, by Cristigera; Campylopus, by Uronychia; Ervilia, by Dysteria; Hallezia, by Paracineta; Rhabdodon, by Orthodonella.

Replacement of a homonym is not the easy task it might first seem to be. A thorough searching of the older literature may be required in order to make abso- lutely certain whether or not available synonyms are already in existence.

( 2 ) Use of a new name. Only if there is no other solution to the problem should a new name (nomen novum) be proposed. I t is credited to the person first publishing it and bears the date of that publication. From that time on it must compete in priority with any synonyms possibly recognized by later workers. It should be noted that all the provisions in the Code which are concerned with “regular” name-making for

‘This is not “spelled out” in so many wcrds in Article 60 of the Code, bu t I assume it to be the case.

316 S O b f E N C L A T U R A L PRACTICES AND THE XEW- CODE

any zoological taxon in the family-group or below also apply to the publication of replacement names. I t might well be further noted that the ‘Code of Ethics” (Appendix A) , which has the same status as a recom- mendation in the Code proper, ought to be carefully adhered to: no replacement name should be published without notifying the preoccupied name’s author (if living) about the situation, giving him ample oppor- tunity to publish a substitute name first himself.

Examples in the protozoological literature of un- necessarily proposed new names include these( 5 ) : Charonella for Charon (when Charonina was already available) ; Craspedonotana for Craspedonotus ( H o - inalozoon available) : Trichoderum for Trichopelma (Leptopharynx available). Such unnecessary names cannot be disregarded; but they fall as junior syno- nyms. Examples of new names properly proposed for well-known ciliate groups include these( 5) : dctinobo- lina for .Ictinoboluz; Chilodonella for Chilodon; Co- closomides for Coelosoma; Cohnilenzbus for Lembus; Epalrella for Epalxis; Histriculus for Histrio ; Pol?- diniella for Polgdinium ; Pyxididla for Pyridium.

Today there is less excuse than ever before for pro- posal of names already in use in the zoological litera- ture. Both the writers themselves and reviewers on journal editorial boards should be willing-indeed, should feel it their duty-to check new generic names (the most common kind of potential homonym) while they are still in an unpublished state. This can be done with the aid of such very convenient publications as Xeave(23) and Schulze et a1.(27).

FAlIILT-GROL-I’ S.Al lES Although references to family-group names occur

scattered throughout the provisions of the new Code and although I have already considered a number of them-sometimes just in passing-in precedin, sec- tions of the present paper. I wish to treat the subject separately here for two principal reasons. Firstly. it is with familial names that most tasonomic protozoolo- gists appear to esperience greatest nomenclatural diffi- culties. And. secondly, the new Code gives much more attention to this important category of names than did the old Regles. By use of cross-references to preced- ing parts of the present paper whenever possible I shall hold actual repetition to a minimum.

In whole or in part, Articles 4, 11. 23, 29. 32, 35-41, 55. 60, 62-65, in particular, contain positive directives concerning family-group tava and their names. Only a few of these, however, need to be discussed here in any detail.

Composition of the “jamily-group.” The categories of the family-group include superfamilies. families, subfamilies, tribes. and “any supplementary categories required” (Art. 35). Each of these taxa is defined by reference to its type-genus. To the “stem” (genitive

without its case-ending) of the name of the type-genus the endings -idae for family and -inae for subfamily are to be added to form the proper names for these groups. I t is recommended, but not mandatory, that the endings -oidea for superfamilies and -ini for tribes be adopted for the other categories in the group (Art. 29).

Many family-group names in protozoology have been formed slightly improperly because of failure to recognize the stem of the name of the type-genus or, in the earlier literature, because of use of improper suffixes (see discussion, and some examples noted earlier). The Code allows-indeed, requires-subse- quent emendation of all such names, fortunately without loss of their original date and authorship ( . i r k 11, 29, 32). This topic also has been discussed in the preceding pages just cited.

I t is curious that protozoologists have consistently used the category of “tribe,” a t least during the past 75 years, to represent a suprafamilial instead of an infrafamilial taxon. For example, among the dino- flagellate Mastigophora, one finds tribes “Gymno- dinioidae” and “Peridinioidae” ; among the gregarine Sporozoa. tribes “Cephalina” and “Acephalina”; among the Ciliophora: gymnostome tribes “Prostoma- ta % ” “Pleurostomata,” and “Hypostomata” ; thigmo- trich tribes “Stomodea” and “Rhynchodea”; and peri- trich tribes “Loricata” and “Aloricata.” To conform with the new rules i t would seem that these categories can no longer be called tribes. In most of the above cited esamples the groups have already been trans- formed by some workers into subordinal taxa, a rank beyond the jurisdiction of the Code. The great ma- jority of the many tribes erected by Poche(25) have simply been dropped by writers of modern proto- zoological textbooks.

If subfamilies are erected i t should be obvious that there must be at least two of them. What should be equally obvious, but which has been disregarded more than once by protozoologists, is that one of the sub- families rnust be nominate, i.e., bear the same name as that of the family except for alteration in the suffix from -idae to -inae. This situation is paralleled by that in the case of names of genera and of species; there, however, the nominate subgenus or subspecies bears exactly the same name as that of the genus or species. Subfamilies, subgenera, and subspecies as well, inter- estingly enough, are relatively rare in taxonomic works on protozoa-perhaps sensibly so, until we are equipped with more precise methods of recognizing re- fined differences among such relatively undifferentiated organisms.

Relationships o j families and their type-genera. I n addition to their names being formed from the so- called stem of their type-genera, family-group taxa have other close relationships with these genera which

NOMENCLATURAL PRACTICES AND THE NEW CODE 317

are sometimes overlooked. All of these are important enough to warrant separate discussion, brief though it must be.

(1) Formation of familial from generic names. Since names of taxa in the family-group are derived from the names of included type-genera, and must be emended if originally improperly formed (Arts. 29, 32, 3 3 ) , the matter of their proper formation needs review. What is meant by the stem of the name of the type-genus is actually that name in the genitive case without its case-ending. In protozoology, as in zoology generally, it is usually very easy to recognize the stem of a ge- neric name: for the great majority of names ending in the nominative case in -us, -a, -um, -es, -is, -on, or even -os, one merely drops these endings. Then -idae is added to produce the proper familial name. Because of the frequency of errors nevertheless found in the literature, i t is worthwhile pointing out that if these common terminations are preceded by 4, or -y, the -i or -JJ is not dropped when the -idae is added ; converse- ly, such a letter is not to be added, either, if it is not already present. Thus Bursariidae, from Bursaria, is not to be spelled Bursaridae; and Podophryidae. from Podophyra [and a -phrys ending is treated similarly], not Podophridae. Nor is Conchophthiridae, from Con- chophthirus, to be written Conchophthiriidae. In spite of their superficial similarity to some of the endings to be considered in the following paragraph, the stem of -styla is just -styl; of -pa&, just -palx; of -stomurn, just -stom. The stem of -phylax, however, is -phylac; I wish to take this opportunity to emend the familial name “Thyrophylaxidae” which was inadvertently so misspelled in my book on ciliate systematics(6): cor- rected, i t should read, with author and date, Thyro- phylacidae Berger in Corliss, 1961.

I t would be far beyond the scope of the present paper to attempt to list here the stems of all possible Greek and Latin words or suffixes which might be in- volved in formation of familial names in zoology. But two dozen7 of Greek derivation, some of which are very common in the protozoological literature, should be noted. Most of these are already in use (or mis- use) in established family-group names in the phylum Protozoa; furthermore, they represent quite popular terminations which may be expected to turn up in ge- neric names proposed as types of families in future years. In the following list the termination of the nominative case of the generic name is given first, separated by a comma from the termination of the word in its genitive case without its case-ending. The examples are arranged alphabetically and are set up

‘Some of these samples occur in Table 2 of Appendix D of the Code; others I have added myself. I am indebted to Pro- fessor F. w. Clayton, Department of Classics, University of Exeter, England, for his careful checking of the entire list pre- sented here.

in parallel columns for ease in reading. In all instances the proper familial name is formed by adding -idae to the stem-termination given on the right-hand side of each double-column.

-anax, -anact -monas, -monad -aulax, -aulac -nema, -nemat -ceras, -cerat -pelma, -pelmat -coyax, -corac -pharynx, -pharyng -cork*, -coryth -sema, -semat -co?,ys, -coryth -soma, -somat -c y lis, -cylid -stigma, -stigma t -desma, -desmat -stoma, -stomat -odon, -odont -stylis, -stylid -lepis, -1epid -thigma, -thigmat -mastax, -mastac -thorax, -thorac -mastix, -mastig -thrix, -trich [*This is to be considered here as a corruption of ‘‘-coryb”l

( 2 ) Wominate categories within the family-group. All nominate taxa within the family-group retain the same type-genus and the same (single) original date and authorship (Art. 36). That is, if a family X-idae Ehrenberg, 1838 (for example), based on the included nominal genus X-us, is either elevated to superfamily- or lowered to subfamily, or if super- and subfamilies are both erected by subsequent workers, they all re- tain X-us as their type-genus and the names are all credited to Ehrenberg: X-oidea Ehrb., 1838; X-idae Ehrb., 1838; X-inae Ehrb., 1838. The subsequent workers would, of course, get credit for (be held re- sponsible for) the other super- and subfamilies estab- lished in the same publications in which the new nomi- nate taxa were obliged to be erected under Ehrenberg’s name. An interesting example complicated by the occurrence of homonymy is given below (p. 3 18).

(3) Type-generic name discovered to be a junior synonym. If the name of the type-genus becomes a junior synonym, the family-group name based on it is not to be changed, after 1960. Before 1961, if a change was made to a new familial name based on the dis- covered senior synonym, and it has won “general ac- ceptance,” i t is to be maintained for the sake of stabil- ity (Art. 40). This important relationship of generic and familial names has been treated in the preceding section on mandatory dates in the Code (see p. 312). It is advisable that i t be well noted by protozoologists, since such cases arise quite frequently in our literature. I n my own work on ciliate families I used stability as my guiding principle and was thus obliged to act with some inconsistency in cases involving junior synonyms of nominal type-genera(6). Fortunately most of my actions now find support in provisions of Article 40 in the new Code. The date of a conserved name ( e g . , a more recent substituted familial name based on the senior synonym of the nominal type-genus) , however, is to be that of the older, rejected name. Thus, for

318 NOMENCLATURAL PRACTICES A N D THE KENr CODE

example, Trimyemidae Kahl, 1933, for Sciadostomati- dae Kahl, 1926 [type-genus Trinzyema Lackey, 1925 (syn. Sciadostoma Kahl, 1926) 1 , should be written in the future as Trimyemidae Kahl, 1933 (1926).

(4) Familial name long unused. A familial name which has remained unused as a senior synonym for more than 50 years could. as I interpret the provisions of Article 23, be considered as a nomen oblitum. Ap- parently this rule is to be applied even if the name of the type-genus inaolved i s a well-known one. Thus I consider that the Code (in one provision or another of Art. 23) supports my decision to drop the unknown name Rhynchetidae Kent, 1880, for the suctorian fam- ily Discophryidae Collin, 191 2 , although the genus Rhyncheta Zenker is retained in the family, nomencla- turally quite unrelated to Discophyra Lachniann (6) . I draw a similar conclusion for the hymenostome fam- ily Pleuronematidae Kent. 1880. containing Pleurone- ma, Cyclidiiim, and other genera. in that I favor Kent’s well-established name over Cyclidiidae Ehrenberg. 1838 (6) .

(5) Type-gcneric name discozlercd to be a junior homonym. If the name of the type-genus falls as a junior hmionym then the names of all taxa in the family-qoup based on that name become invalid and must be replaced. The familial name must be changed to one based on the valid name of the genus involved.8 but both the new family-group name and the new name of the type-genus are to take the dates (not author- ships. thouqh! ) of the replaced names. in order to pre- vent a change to a subjectively different family and type-genus (Art. 39) .

For example. the entodiniomorphid ciliate genus Polydiniunt Kofoid. 1935, type of a subfamily Poly- diniinae Kofoid. 1935. is preoccupied by the dino- flagellate genus Polydiniunz Kofoid Kr Swezy. 192 1. The generic name which is the junior homonym has been replaced by Polydinivlla Corliss. 1960: and the subfamily, by Polydiniellinae Corliss, 1960: these nominal taxa should now be written with the date “1935” appended a t the end. enclosed in parentheses. Later during the same year in which my publication appeared( 5 ) . Noirot-TimothPe( 24) raised Kofoid’s subfamily to family, incorrectly labelling it as a ”fam. nov.” In a footnote she corrects the name proper to Polydiniellidae from I’olycliniidae, having noted my nomenclatural paper while reading proofs of her own work. But rather than considering it as a “new fam- ily.” dating from her publication. actually the full name of Soirot-Timothee’s family. with authorship and dates, must be given in the future as Polydiniel- lidae Corliss, 1960 ( 193 5 ) .

( 6 ) Selection of type-genus. By the provision of

.4 widely accepted familid name established before 1961 “under a different procedure,” home\ cr, is not to be upset

Article 64, a protozoologist erecting a new taxon in the family-group is free to select as type-genus any of the included genera. The genus need not be the oldest one in the group, although from time to time one finds this argument put forth (unjustifiably) as a reason for rejection of a familial name which happened not to be based on the oldest included nominal genus. It should be noted, although this is not, unfortunately, pointed out directly in Article 64, that a zoologist is not free to make such a selection if his “new” taxon includes certain genera which themselves are already types of former families. This important point is treated in the paragraphs immediately following this one.

Proper name in the case of an ainalgamated family. All too commonly when protozoologists or parasitolo- gists have formed a “new” family-group taxon by the union of two or more existing taxa of equivalent rank, they have failed to follow any consistent nomenclatural procedure. Type-genera of the previously erected fami- lies included in the amalgamation are often totally ig- nored! and a new name is proposed without regard for even the ethics of the situation: or else a name is chosen from among those already available, but seem- ingly purely at random, with no attention to priority. A large share of the blame must fall on the old Rkgles which were so very deficient in their coverage of this matter of such importance to workers engaged in large- scale taxonomic revisions. The new Code, fortunately, clarifies the situation greatly.

4ccording to a section of Article 23 (Law of Pri- ority), a family-group taxon formed by the union of two or more taxa of that group must take as its name the oldest valid family-group name made available by the amalgamation. There follows an “escape clause” in case relief from priority is in the best interests of stability and universality: the person joining the for- merly separate familial group may request the Com- mission to decide which name is to be accepted if the oldest name seems definitely a poor choice. Note even here, however, that the zoologist is expected to know thoroughly the taxonomic and nomenclatural history of all the groups involved-his own ignorance. alone, forms no legitimate excuse for the automatic over- throwing of priority.

In a t least two major cases“ Kahl, in his indispensa-

’ Leptopharyngidae Kahl, 1926, unnecessarily changed to Trichopelmatidae Kahl, 1931 (when Leptopharynx fell as a junior synonym of T r i c k o p e h a ) , was chosen for the name of a family o i trichostomes over the already available Micro- thoracidae M-rzesniowski, 1870. The name Metopidae Kahl: 1927, was established for a heterotrich group in which both Gyrocorythidae Stein, 1867, and Caenomorphidae Poche, 1913, were available(6). Two other instances might be mentioned in passing: Kahl’s( 1 7 ) favoring, seemingly without firm basis, Holophryidae Perty, 1852, over Enchelyidae Ehrenberg, 1838, as name of a major family of gymnostomes; and Ancistrumidae Kahl, 1931 (unnecessary replacement, anyway, for Ancistridae

NOMENCLATURAL PRACTICES AND THE NEW CODE 319

ble monographs on fresh- and salt-water ciliates pub- lished during the period 1926 to 1935 (see 17), chose to give new names to families of revised composition, families which, however, contained genera associated with available and valid older familial names. In spite of the justifiably authoritative weight of Kahl’s syste- matic work in general it seems inadvisable to petition for suppression of these older names which were widely used until Kahl unreasonably cast them aside in his publications of only 30 years ago (6) .

A second example may be found in the excellent taxonomic revision of families comprising the large ciliate order Hypotrichida published very recently by FaurC-Fremiet (1 1) . Among other proposals a family “Keronidae n. nom.” is established to contain Kerona, Epiclintes, Eschaneustyla, and Paraeuplotes. But the last-named genus brings along with itself the familial name Paraeuplotidae Wichterman, 1942, which natu- rally has priority over any name suggested for the first time in 1961. However, in this particular instance, Dujardin’s family “Keroniens,” with its suffix already emended (6) in accordance with provisions in Article 11, saves the name of FaurC-Fremiet’s choice: Keroni- dae Dujardin, 1841, may be considered the proper name for the amalgamated group.

Proper n a m e ( s ) in the case of a fragmented family. This section really forms a corollary to the preceding one, and the same nomenclatural directives may be applied. In proposing revisions a t the familial level, a protozoologist may well become involved in both sorts of situations simultaneously.

In splitting up a large family, two nomenclatural rules must be kept in mind: the new, smaller groups are to be given the oldest valid familial names avail- able for them; and the fragment containing the type- genus of the original single family must retain the original familial name. This second rule needs to be stressed in particular, since so frequently it is not fol- lowed. I suspect that the reasoning behind the drop- ping of the originaI aII-inclusive famiIiaI name is that the reviser probably wishes to get rid of the connota- tion (over-large, vague, polyphyletic group, etc.) asso- ciated with that name: he hopes that its complete re- jection, along with his revision of the characterization and composition of the group, will yield this happy result.

Thus in the recent important revisory work by FaurC-Fremiet ( 1 1 ) , also cited above, the commend- able taxonomic treatment of the grossly over-extended hypotrich family Oxytrichidae Ehrenberg, 1838, has resulted in proposal of a number of smaller families,

Issel, 1903) over Hemispeiridae Konig, 1894, for name of an important family of thigmotrichs. Perhaps unwittingly, Kahl, has, nonetheless, caused considerable confusion in such cases in literature published subsequently to his own work, for his im- pressive taxonomic monographs have also generally been ac- cepted as authoritative in all nomenclatural matters as well.

including resurrection of Pleurotrichidae Butschli, 1889. But in FaurC-Fremiet’s Pleurotrichidae one finds the genus Oxytricha; therefore “Oxytrichidae.” obvi- ously predating Butschli’s name, cannot be so easily disposed of. In fact, I can think of no way for legiti- mately getting rid of this widely used, well-known name: elevating it to the category of superfamily would be of no help, for it would still have to be recognized as the nominate family of its group (a fact often overlooked in the literature, incidentally, espe- cially with regard to the parallel situation of a family and its several subfamilies). As suggested in footnote 2, addition of the major reviser’s name follow- ing the name of the original author of the family may present an agreeable solution to the problem. In the case under discussion the family which FaurC- Fremiet has labelled Pleurotrichidae could be called Oxytrichidae Ehrbg., 1838, emend. FaurC-Fremiet, 1961.

T H E PROBLEM OF TYPES Some 24 pages of the new Code proper (Arts. 61-75)

are devoted to the “type-concept” in zoology; one- third of these deal specifically with the problem of type-specimens. Protozoologists, as a whole, have never shown much concern over type-genera of families or even type-species of genera, in spite of the rules of nomenclature, as examples in preceding sections of the present paper make all too clear. But even less attention has been paid to the concept of type-speci- mens for our species. News of this latter neglect is always received with disbelief, sometimes even horror, by vertebrate systematists. Indeed, it was my genuine worry when attending the meetings of the Colloquium on Zoological Nomenclature in London in July, 1958, vastly outnumbered as I was as a protozoologist, that some provision of the new Code would render the name of practically all protozoa invalid by making the desig- nation of type-specimens mandatory for recognition of any species! But the closest the new rules seem to come to this is in Article 45, in which the statement may be found, “Each taxon of the species-group is objectively defined only by reference to its type- specimen” [italics mine]. Or in the first recommenda- tion attached to Article 73, which reads, “A zoologist when describing a new species should clearly designate a single specimen as its holotype.”

I have used the rather harsh word “neglect” in the preceding paragraph. Actually it is not altogether in- excusable that protozoologists have designated so few type-specimens. As I have briefly pointed out else- where( 7), it has been only natural that myriads of quite obviously new species have been described solely from examination under the microscope of fresh, living ma- terial suspended in temporary mounts. Not only are there often few anatomical features of taxonomic value

3 2 C XOMENCLATURAL PRACTICES AND THE NEW CODE

which are improved by fixing or staining-this was particularly true in times of earlier light microscopy- but also many so-called permanent preparations may not retain recognizable specimens for more than a few years. Furthermore, in certain groups some of the im- portant diagnostic characteristics, including all non- morphological ones. of course, are simply not pre- servable in a fixed condition on a slide or in a vial. I n general i t is far easier to preserve in recognizable shape a coleopteran or lepidopteran insect. a bird. or a part of a fossil mammal, for esample. than a lep- tomonad or euglenoid flagellate. an astramoeba. or a podophryan suctoridan.

Types at the familial l e w l . Generally there is little trouble in recognizing the type-genus of a family-group taxon. even if the worker erecting the family has not mentioned the type in so many words. because the familial name is (supposed to be) formed from the name of that genus. However. there are several impor- tant situations in which it may be difficult to know the type of the family: if the familial name bears no relationship to an available name of any of its included genera; i f the names of the type-genus and of the fam- ily have become separated in the shuftle of a taxonomic revision (joining erstwhile separate families or break- inq up a large one): or if the familial name is based on a junior synonym of the valid name of the type- genus. The first and second esamples are. of course. improper situations which should be corrected immedi- ately upon discovery (see discussion above. p. 318). In the third example. which represents a perfectly legitimate case-in fact. it may involve a mandatory situation after 1960 (see -4rt. 40. and p. 317 of the present paper)-it would be strongly advisable for the author to include the name of the junior synonym in his list of genera. in parentheses following the valid name of the type-genus. For esample(6): Family Gyrocorythidae. genera Brpometopus, Caenomorpha (syn. Gyroc-orys) . Copemetopus, Ludio, Metopus, Pal- mariuna, and Tropidoatractus.

Types at the generic- l e w l . The type of a genus or subgenus is a species. For any genus-group name pub- lished after 1930. the author of the original publication must definitely designate a type-species (Arts. 13. 68). If he does not, his new generic name has no status and, strictly speaking, should not be recognized a t all by subsequent workers.

Type-species (which, incidentally, should never be called ”genotypes.” as they once widely were, because of the established usage of that word-with an entirely different meaning-in genetics) are given different terminology depending on their mode of origin: type by original designation, type by indication, type by subsequent designation. But discussion of such details, which can become quite complex and are primarily of interest to nomenclatural specialists only. are generally

beyond the scope of the present paper. Protozoologists and parasitologists involved in work demanding treat- ment of type-species should study carefully the provi- sions of Articles 67-70 of the new Code. The recom- mendations given for guidance to workers who must make subsequent designation of types are particularly clear and helpful (Art. 69).

Types at the specific level. I have mentioned at the beginning of this section some of the difficulties proto- zoologists face in designating type-specimens for all of their species. Xevertheless, indolence and ignorance have also played a part in keeping us in the dark ages of taxonomy, and we should make every effort to sta- bilize our science in this important respect. Students of the Foraminiferida. and of certain other groups rich in fossil material (e.g., consider the heroic efforts of Deflandre in this last regard), have long been desig- nating type-specimens. Of course their organisms gen- erally lend themselves neatly to preservation in a per- manently recognizable form. Scattered workers in other areas of protozoology, particularly in recent years. have also been conscientious in labelling speci- mens as type-material and carefully depositing them in proper institutions. Parasitic as well as free-living forms have been successfully so treated. One of the most energetic leaders in stressing the great need for types in protozoology and for their being gathered into central collections is Dr. Jaroslav Weiser of Prague ( e g . , see 28). The value of subsequent investigators’ being able to make direct comparisons of material in their tawnomic revisory work cannot be emphasized too strongly.

All types in the species-group are to be regarded as “the property of science” and are to be kept safely preserved, clearly marked, and accessible to any com- petent worker needing temporary use of them for le- gitimate research purposes (Art. 7 2 ) .

There is no reason why many of us-includinc the writer-should not take this matter of type-specimens more seriously in the future than we may have done in the past. A number of modern cytological-protozoo- logical techniques (see 8. 20, 29) result in excellent preparations of considerable permanency: various sil- ver methods, for example, lend themselves admirably to the cause in the case of many flagellates and ciliates, both free-living and parasitic. If certain conveniently located institutions‘” can be established-and widely publicized-as freely available international deposi-

10 I am hoping to be able to establish a “collection center” in the United Slates for all type-material specifically related to protozoa belonging to the subphylum Ciliophora. Any deposi- tory for protozoological specimens should be associated with a person. or persons, not only reasonably knowledgeable in the field but also actively willing to take proper care of the mate- rial; to make it available, often by post, to specialists engaged in raisions of particular groups; and to act in consultation x i th such experts concerning pertinent nomenclatural details, n hich may be numerous in certain instances.

NOMENCLATURAL PRACTICES AND THE NEW CODE 32 1

tories, in line with Weiser’s(28) good suggestions, then it only remains for us to make the individual effort to designate and label our type specimens whenever we describe new species. In certain instances older mate- rial should be given types (neotypes: see below) as well. First of all, we must understand what types are, in the case of species; the brief descriptions which fol- low should be supplemented by careful study of the provisions of Articles 71-75.

(1) Holotype. A single specimen properly desig- nated or indicated as “the type-specimen” in the origi- nal paper by the author(s) of the description of a new species. Outside-but seldom, one trusts, within-the field of protozoology there often may be only one origi- nal specimen on which the description of a new species is based. But even when the author bases his species on study of many specimens (the “type-series”), it may be advisable for him to designate one of them as holotype: in fact, the Code recommends such action.

( 2 ) Syntypes. All specimens in a type-series of a species none of which has yet been expressly singled out as a holotype or a lectotype.

( 3 ) Paratypes. All the remaining specimens in a type-series of a species after the original author has singled out and labelled one as the holotype. In other words, paratypes are the syntypes left over after desig- nation of a single specimen as holotype.

(4) Lectotype. One of the syntypes designated by any zoologist as the type-specimen of a species, subse- quent to the original publication on the subject, if no holotype was established by the author of the first paper in his description of the new species. Lectotypes must be designated individually, not by any wholesale methods. [ I t is interesting to note that a figure may, in effect, be designated as lectotype (Art. 74).]

(5) Paralectotypes. All of the original syntypes in a type-series of a species which are remaining after a Iectotype has been selected by some worker.

( 6 ) Neotype. A single specimen properly designated as the type-specimen of a species for which, through loss or destruction, there no longer exists a holotype or lectotype or any paratypes or syntypes. In other words, an entirely new type subsequently selected from fresh or different material because of irretrievable loss of the specimens belonging to the original type-series, the organisms originally studied and used by the au- thor of the new nominal species. The provisions of Article 75 strongly emphasize that neotypes are to be designated only under exceptional circumstances, even in revisory work: certainly never as a matter of “cura- torial routine” or for species whose names are not in general usage. And it is mandatory that a neotype become the property of a recognized scientific or edu- cational institution with proper facilities for preserving and making universally accessible such research ma- terial.

MISCELLANEOUS CONSIDERATIONS Many relatively minor provisions of the new Code,

or other data related to it, have not been specifically cited in the present work, in keeping with my aim to discuss the application only of articles of greatest im- portance in protozoological and parasitological taxo- nomic researches. But a number of more or less mis- cellaneous though nonetheless significant points which were not entirely appropriate for inclusion in the sev- eral major sections in preceding pages of this paper still need to be mentioned. These are considered briefly in the following paragraphs.

Suprafamilial categories. The Colloquium on ZOO- logical Nomenclature meeting in London in July, 1958, decided not to undertake regulation of the names of taxa belonging to categories above the family-group level. Thus no provisions of the new Code contain directives relative to names of such groups as orders, classes, and phyla. This is a reasonable decision, al- though it may come as a surprise to protozoologists familiar with Hemming’s ( 12 ) Copenhagen Decisions, published in 1953, or Bradley’s ( 1 ) 2 74-page prelimi- nary draft of the new Code, published in 1957 in the Bulletin of Zoological Momenclature. Before the Lon- don meetings, several articles, in whole or in part, had been planned which were to have imposed quite rigid control over various nomenclatural aspects of the su- prafamilial categories. Chaos might well have resulted.

Nevertheless, I believe that it behooves protozoolo- gists and parasitologists to pay a little more careful attention to the names of higher-level taxonomic group- ings within the phylum Protozoa and to treat them with more consistency in the future. Sometimes the principles of nomenclature which are applied at the lower levels might well be extended to these supra- familial taxa with universally agreeable results. For example, no one, I believe, will deny that recognition of Stein’s family “Ophryoscolecina” as Ophryoscoleci- dae Stein, 1859, is anything but fitting and proper. Yet a number of recent workers have not seemed to hesitate to publish ordinal names as their own, as new names, even if they have only changed the speIling of the suffix of names previously in existence for the same groups.

Homonyms, always a source of potential confusion, also exist, unreplaced and virtually unnoticed, among names of suprafamilial zoological categories.

I agree with those zoologists who claim that to re- quire uniform endings for names of higher taxa throughout the entire animal kingdom would wreak havoc with stability.ll On the other hand, for a rela- tively small group of organisms such as those compris- ing the phylum Protozoa it does not seem unreasonable to adopt a mild amount of uniformity: for example,

”And who would determine exactly what those endings were to be?

322 XO~ZIENCLATURAL PRACTICES AND THE NEW CODE

-ida for ordinal endings and -ina for subordinal. There is clearly a trend in this direction (see 7 and citations therein). The slight orthographical repairs which would be needed to establish such uniformity in end- ings should in no case. in my opinion. cause abandon- ment of the original authorship and date of the names so altered.

The principal reason why names of higher categories should not be subjected to rigid rules of nomenclature derives from the fact that the very instability of syste- matics a t such levels is indicative of the healthy “free- dom of taxonomic thought and action” so well worth preserving. In protozoology the suprafamilial groups are certainly in a state of Hux at the present time(7), a situation which may be viewed as indicating unham- pered progress toward an ultimately desirable degree of taxonomic stability.

Sumber of times a name can be “new.” Surely everyone would agree that a name of any taxon. whether covered by the Code or not, can be strictly new only the first time i t is published. Yet i t is not a t all uncommon in the protozoological and general parasitological literature to find a new species. a new genus, a new family. or even a new order being called “new” in two-or sometimes even more!-of the author’s own publications. For purposes of deter- mining synonymy or homonymy. if nothinq else. it is most distressing to find new tasa with dates of origin ranging over a period of several years!

Jn the earliest publication the new name should be followed by “gen. n..” “sp. n.,” or whatever is appro- priate. The name of the author and the date of the publication should not appear after the new name any- where it is used in that initial work. I n later. often fuller, publications the author should not modestly desist from placing his own name. and the oriqinal date, after the name of the taxon under consideration, now no longer new. If-the practice is not recom- mended-the name first appeared in a published ab- stract then that abstract must be treated as represent- ing as much a genuine publication as any other paper, long or short.

The expression “nomen nommt” (abbreviated as nom. n., n. nom.. nom. nov., or n.n.) should be used to denote only a name which i s being proposed as a re- placement !or a preoccupied name (a junior homo- nym). It is all too commonly employed where “fam. n.,” “gen. n.+” etc., should have been used. In subse- quent publications, however. the author and date-if it is necessary or desirable to cite them a t all-are placed after the scientific name in a manner exactly the same as in the case of names of taxa newly erected.

The Code treats the above matters only as “general recommendations” (Appendix E ) , along with a num- ber of other important items of advice. But they be- come involved in mandatory provisions, such as in the

directives concerned with correct dates of publication (Arts. 21, 22) or those having to do with matters of priority (Arts. 23. 24).

Subcategories in protozoology. The international rules provide clear directives concerning the names and naming of subfamilies, subgenera, and subspecies. As Articles 36, 43, and 46 state, all categories within the family- or the genus- or the species-group are co-ordi- nate: they have equivalent status in nomenclature, are subject to the same rules and recommendations. There must always be a “nominate” sub-category for any group-tauon which is subdivided. With a few notable exceptions (e.g., subfamiIies in the ciliate order Tin- tinnida) ~ subdivisions are generally but rarely seen in the protozoological literature. This is particularly true a t the species-level, and understandably so. The gam- ma stage of taxonomy, embracing the study of sub- species as “evolutionary units,” has seldom been reached by protozoologists. due to the very nature of the material they have to work with(7). It is thus sensible that such precise infraspecific divisions have seldom been attempted for the protozoan “morphologi- cal species.”

Inappropriate and incorrectly formed names. It is very important to realize that generic or specific names cannot be changed or rejected because of inappropri- ateness (-4rt. 18) or because of improper latinization or incorect transliteration (Art. 32) , even by the origi- nal author of the poorly chosen or poorly composed name. Dobellf 10) h a d - o r , at least, would have today -no right to endorse the change of “Terarzympha” to “Terutonympha” (see p. 308 of the present paper). [On the other hand, fami2iaZ names can be-and, in- deed. must be-changed , as discussed earlier, if either the base or the suffix needs emendation to conform with the appropriate rules (,4rts. 32, 33).1

A name which is inappropriate or undesirable in practically any way will nevertheless “stand through the centuries as a monument to the intelligence, taste, judgment. and ethics of its author”(22). The recom- mendations comprising appendices D and E of the new Code offer splendid suggestions for future guidance. ml th rare exceptions nothing can be done concerning past nomenclatural performances: for example, with such “monstrous Latin parasites”( 10) as Teranym- pha; with such lengthy concoctions as Brachyuropusk- kyodermatogammarzts greivlingwmnemnotus’2 or B . anteromediobasalinzagnofasc~~tipenizis ; with such non- euphonious names as Aages or Zyzzyva; with such

“Not a protozoon! Our record for length is perhaps to be found merely among such names as Eimeria ninakohlyakimo- oae, Entorhipidium multiinuronucleatum, Eumonodontophrya kijenskiji, Murospirotrichonympha porteri, Monocercomonoides quadrifunilis, or Tontonia appendiculariformis. I understand that the crustacean names cited in the text, above, have been formally rejected by the International Commission on Zoologi- cal Nomenclature.

NOMENCLATURAL PRACTICES AND THE NEW CODE 323

facetious samples as Amphionycha knownothing, Ta- banus balzaphyre, or the classically infamous Peggi- chisme (“Peggy kiss me”) , Polychisme, Dolichisme, etc.

Language of publication. It is recommended (Ap- pendix E) that the description of a new taxon be writ- ten in English, French, German, Italian, or Latin. If it is not, the suggestion is made that the account should be accompanied by a translation (summary adequate?) into one of those languages and that the explanation of figures should definitely be so given.

These recommendations may work hardship on some of our Russian protozoological and parasitological col- leagues, for example, among whom are many active taxonomists. But it may he recalled that no recom- mendation in or associated with the Code has the force of a mandatory provision. From one point of view perhaps the botanists have the most sensible solution after all, in their persistent requirement of a Latin diagnosis for every new plant taxon before its name can be considered to have been validly published ( 15).

Petitions to the Commission. Any zoologist may submit a case involving a nomenclatural problem to the International Commission on ZoologicaI Nomen- clature for resolution; in fact, he is required to do so in certain instances. The plenary powers of the Com- mission enable it to suspend the application of any provision of the Code “if such application to a particu- lar case would in its judgment disturb stability or universality or cause confusion” (Art. 79). The Com- mission’s interpretations of the Code are generally published in the form of “Declarations” (provisional amendments to the Code) or “Directions” or “Opin- ions” (Arts. 77, 78). All such statements are to be followed by zoologists in any given case, whether ple- nary powers as such are involved or not. Another duty of the Commissioners pertinent to the present discus- sion is to compile the Official Lists of accepted names and the Official Indexes of rejected names and works in zoology (Art. 7 7 ) .

From a practical point of view it is discouraging to the conscientious taxonomic protozoologist to even con- template making use of petitions to this authoritative international body of nomenclaturists. Reasons: we have so many hundreds of cases requiring treatment; composing a petition is often a long and difficult task (the Commission itself is only a fact-reviewing not a fact-finding board); and so few cases can be dealt with annually by the Commissioners (but through no particular lack of diligence on their part). With re- spect to the last point the figures given by Mayr et al. (22) are truly appalling: from 1907 to 1936, only 133 cases (involving use of plenary powers to suspend pro- visions of the Law of Priority) were considered by the Commission; from 1936 to 1950, 218 decisions were made (still only averaging 14 per year); in 1951, 268

cases were on the docket and new applications were being received a t the rate of about 100 per year! And one can imagine that the rate probably has increased considerably in still more recent years.

Very few names of protozoological taxa appear in the Official Lists or Official Indexes. Although there are hundreds of “candidates” for these Lists. it is- again-a matter of time and interest on the part of protozoan systematists whether or not much will ever be done about it. Such problems might well be con- sidered by the Committee on Taxonomy of the Society of Protozoologists.

Responsibilities of editors and editorial boards. I t is reasonable to conclude that essentially all published material, perhaps exclusive of certain books, contain- ing information of a nomenclatural nature has passed not only through the hands of a publisher but also, generally, across the desk of an editor. The editor, whether or not he has the assistance of members of an editorial board, should have assumed a certain degree of responsibility for the accuracy of the manuscript placed in his hands. If he is not personally acquainted with the provisions of the international rules of nomen- clature he should not hesitate to ask for help from per- sons with such knowledge. I t seems to me that exercise of just a little more editorial care in such matters would have reduced impressively the number of improper names now in need of mandatory emendation, the hun- dreds of inappropriate and undesirable names in the literature which cannot be changed, the many incor- rectly cited dates and authorship, erroneous combina- tions, double publications of a new name as new, and myriads of inaccurate or incomplete bibliographic cita- tions of taxonomic and nomenclatural importance. In addition, now, to the publication of the new Code itself (16)) editors have available such helpful guides as the recently appearing Style Manual jor Biological Jour- nals (3) , the revised Schenk & McMasters ( 18), and the invaluable systematics textbook of hlayr, Linsley & Usinger(22).

Of course no amount of editorial concern alters the fact that it is the author of a paper himself who is ultimately solely responsible for the accuracy of his work, nomenclatural or otherwise.

One specific point concerning truly editorial respon- sibility warrants special mentioning: that of the precise dating of a publication. December copies of journals, for example, which often are actually issued some time in the following year, or parts of works issued sepa- rately and later bound into single volumes, should clearly bear-somewhere within the publication-an indication of the exact date of their particular appear- ance. This point is stressed in recommendations at- tached to Article 21 of the Code. For reasons of pri- ority-as well as just plain accuracy, anyway-it is very important that the actual date of appearance of

324 SOMEXCLATVKAL PRACTICES AND THE KEW CODE

publications containing taxonomic papers be clearly ascertainable by any subsequent worker (s) needing such information.

R E F E R E S C E S

1. Bradley, J . C. 1957. Draft o i the English text of the Zn- trrnational Codr o f Zoological Sonienclature as amended by the Paris (1948) and Copenhngen (1953) Congresses. Bull. Zool. .?.onzed. 14, 11-285.

2 . Cleveland. L. R . 1938. Morphology and mitosis of Tera- npinpha. ilrch. Protistrnk. 91, 442-51.

3. Conierence of Biological Editors. Committee on Form and Style. 19613. S t y l e Manual jor Biologicul Journals. .Am. Inst. Biol. Sci., Washington.

4. Corliss, J . 0. 1957. Somenclatural history of the higher tnxa in the subphylum Ciliophora. ; f r r h . Protistenk. 102. 113- 46.

3. __ 1960. The problem of homonyms among generic names oi ciliated protozoa. u i th proposal o i several new names. J , Proto;ool. 7. 269-78.

6. ~ 1961. The Ciliated Protozoa: Charactrri:ation, Cla.ssijcrrtion, and Guide. to the Litrralure. Pergamon Press. London and S e w York.

7. 1962. Taxonomic procedures in classification of protozoa. .S!inp. Sor.. g r n . Jfirrobiol. 12, 37-67. 8. -__ 19623. Firinl: and staining of protozoa, in Lacy,

I). 8r Palay, S . O., The .Ilirrotoiiiist’s I ~ a d e - M e r u i i i (Rolles h e ) . 12th ed.. Churchill, London. ( In press.)

9. Corliss, J . 0. & I)ou;!hert>. E. C. 1962’. An appeal for stabilization of certain names in the iamily Tetrahymenidae (Protozoa Ciliophora) . Ivith special reference to the generic name Tetrahgnrena Furgabon, 1940. ( I n preparation.)

10. Dobell. C. 1939. On “Trrangrnpha” and other monstrous Latin parasites. Parasilolog)( 31. 25.5-62.

11. FaurC-Fremiet. E. 1961. Remarques sur la morpholopie comparee et la systbmatique des Ciliata Hypotrichida. Cornpt. rrnd. 252. 3515-9.

12 . Hemming. F. 1953. C o p ~ n h n g r n Decisions on Zoologiral -Yonirnrlature. London.

1.3. - 1956. Opinion 418. \.alidation under the plenary powers of the generic name Strntor Oken. 1815 (Class Cilio- phora) , In Opinions and Drclarations Rriidrrrd b! t k e Inter- national Corninission 011 Zoologira! Sontrnrlatrrre 14. 13-6s.

14. 19.58. Official text oi the “Riyles Internationales de la Somenclature Zoologiquc” as it esisted up to the open-

ing of the Paris Congress in 1948. Bull. Zool. Nomencl. 14, i-xxviii.

15. International Code of Botanical Nomenclature adopted by the Eighth International Botanical Congress, Paris, July 1931. 1956. Utrecht.

16. International Code of Zoological Nomenclature adopted by the XV International Congress of Zoology, London, July 1958. 1961. London.

17 . Kahl, .A. 1930-1935. Urtiere oder Protozoa. I: Wimper- tiere oder Ciliata (Infusoria), eine Bearbeitung der freilebenden und ectocommensalen Infusorien der Erde, unter Ausschluss der marinen Tintinnidae, in Dahl, F., Die Tierwelt Deutschlands, Teil 18 (year 1930), 21 (1931), 2 5 (1932), 30 (1935) , pp. 1-886, G. Fischer. Jena.

18. Keen. A. M. S- Miiller, S. W. 1956. Schenk & McMas- ters’ Procedure in Taxonomy , 3rd ed. Stanford Univ. Press, Stanford.

19. Kirby, H. 1944. Une faute de transcription, d’ortho- graphe. ou d’impression. Science 100, 425-7.

20. ~- 1950. Materials and Me thods in the Study o f Proto-oa. Univ. Calif. Press, Berkeley.

21. Kudo. R. R. 1959. Pelomyxu and related organisms. A n n . IT. 1.. dcad . Sci. 78, 474-86.

2 2 . Map-. E., Linsley, E. G. S- Usinger, R. L. 1953. Methods and Prinriplrs o f Sgstrnzatic Zoology. McGraw-Hill, New Tork.

2 3 . Seave , S. .%. 1939-1950. fiomenclator Zoologicus. Vols. 1-1.. Zoological Society of London, London.

2 1 . Soirot-Timoth&, C. 1960. Etude d’une famille dc ciliCs: les Ophryoscolecidae. Structures et ultrastructures. Ann. Sci. nut., Zool. (skr. 1 2 ) , 2, 527-718.

25. Poche, F. 1913. Das S3stem der Protozoa. Arch. Pro- t i s tenk . 30, 125-321.

26. Riley, S. D. 1961. Pcrsonal communication concerning the date of publication of the Code. [Note added in proof: this information has now appeared in print, in Par t 6 of Vol. 18 of Bull. 2001. S o n i r n c l . , p. 353.1

27 . Schulze, F. E. et al. 1926-1954. Nomenclator ilnimalium Grnrrrrrn e t Subgrnmi tn . Vols. 1-1’. Preuss. Akad. Wiss., Ber- lin.

2s. Weiser, J . 1962. Type collections of protozoa and tax- onomy. Proc. 1st int. C o n f . Protoiool., Prague, August 1961. ( In press.)

29. [Yenrich, D. H. S- Diller, M’. F. 1950. Methods of pro- tozoology, in Jones, R. McC., McClung’s Handbook o f Micro- scopical Technique, Hoeber, S e w York, 432-74.

30. \Yenyon. C. M. 1926. Protozoology, a Manual for Mrdi- cal M e n , l’rfrrinarians and Zoologists. I7ols. I and 11. Bailliere, Tindall and Cos, London.