The Problem of Homonyms among Generic Names of Ciliated Protozoa, with Proposal of Several New Names

J. PROTOZOOL. 7(3), 269-278 (1960). 2 69

The Problem of Homonyms among Generic Names of Ciliated Protozoa, with Proposal of Several New Names”

JOHN 0. CORLISS

1 k p n r t w r . l l t of Zoology, University of Illinoih, {Jrhann, Illi111)i~

SYNOPSIS. Over ninety Kenera of ciliatcd protozoa have been given names preoccupied by the generic names of other animal organisms. To date some sixty of these junior homonyms ncver have been corrected. I n half of these cases the homonymous ciliate names have been buried as synonyms of one kind or another, however, and need be of no further practical concern. For the remaining genera, thirty in num-

TLTATE genera now number well over eight hun- C dred, having nearly doubled their number in the past thirty years; counting only those considered as legitimately constituting independent taxa and whose names are valid( 17). Several hundred others described in the literature may be considered identical with previously established genera, their names thus falling as synonyms. Among the recognized taxo- nomically distinct genera is a group, involving a larger number of ciliates than commonly realized. comprised of forms whose names fall as homonyms of generic names properly proposed at earlier dates for other animals. The present work is concerned specifically with the nomenclatural problem existing with regard to this last-mentioned group of some ninety generic names. I am in agreement with the widely held be- lief among modern systematists that taxonomic stability never can be achieved or maintained through preservation of identical generic names both currently in use for two quite separate organisms.

GENERAL CONSIDERATION OF HOMQNYJIS

In cases of homonymous generic names for ciliated protozoa the following four possible solutions may be considered.

1 . Preservation of the junior homonym Occasionally a generic name may have become so

well-known or so firmly entrenched in the literature that it is desirable to have it officially preserved. even though it falls as a junior homonym by strict applica- tion of automatic provisions in the International Rules of Zoological Nomenclature. In such cases use of the same name for the other organism must be sup-

* The work reported here has becn supported by funds from the National Science Foundation. I t is a pleasure to mention the assistance of Miss Louise Weisberg in carrying out some of the library research involbed. I am indebted to Dr. Hobdrt M. Smith for critical reading of the manuscript and to Dr. Donald P. Rogers for assistance in formation of certain of the new generic names proposed.

ber, replacement names are proposed in the present paper in accordance with provisions of the International Rules of Zoo- logical Nomenclature. Such action is considered to be in the best interest of ultimate taxonomic stability among the groups of organisms involved. New names also have had to be proposed for five families or subfamilies whose typc genera have lost their original names through homonymy.

pressed. A petition must be prepared and submitted to the International Commission on Zoological No- menclature for its opinion.

This has been done for the generic name of the ciliate Stentor‘ Oken, 1815, preoccupied by Stentor Saint Hillaire, 1812, a little-used name of a genus of mon- keys. This widely acclaimed action has resulted in the long overdue preservation of Stentor as a nomen conscrvatunz of the ciliate genus(29,42). But I believe that none of the other genera under consideration in the present paper is worthy of similar treatment.

2 . Racognition of tJic junior Jtonzonym as a junior synonym

When the junior homonym involved has fallen into disuse, I believe that there is nothing to be gained by proposing a replacement name for it. In some twenty cases implicating genera of ciliates the preoccupied names have fallen as junior synonyms. Making a nomenclatural correction in these instances, although authorized by the Rules, seems entirely unnecessary. Junior homonyms which have fallen as junior synonyms include the following (see Table I for author- ship and date of the names):

Clypeolum, synonym of Strombidium; Cordyloso- ma, of Glossatella; Crateronaorpha, of Euplotes; Drepanostoma, of Loxodes; Iduna, of Dysteria; Lada, of Ptychostomum;2 Leptodesinus, of Homalozoon; Metopides, of Metapus; Morgania, of Kidderia (in turn a junior homonym itself; and both names may be synonyms of Conchophthirzrs: see(4.17.40) : Oospira, of Gymnodinioides; Paraglaucoma (of IVar-

1 Interestinglq enough, replacements for the name of thih protozoan genus have been proposed in the past , c.g, S h - tor rlla Reichenbach, 1828 But they never have been adopteti in authoritathe protozoological works.

2 Ptychostomiim Stein, 1860, itself appears to be preoccupied by Ptycho~tomlrm Agassiz, 1846, an ernendation oi Ptygo~toiniinz Ehrenberg, 1841, even though the emendation ma? be considered invalid. But names proposed in Iiterature- recording series or generic indices have no official status, anti thu5 ,\gassiz’s name offers no threat to Stein’? narne(l,49.60).

___

2 70 NEW GENERIC NAMES OF CILIATES

ren, not Kahl) , of Tetrahymena (in need of preservation itself because i t is a junior synonym: see( l8) : Rhabdodon, of Orthodonella; Rhynchophrya (of Raabe, not Collin), of Gargarius; Spathidiopsis (of Kahl, not Fabre-Domergue) , of Spathidium; Telosto- ma, of Platyophrya; Torquatella, of Strombidiuni; lrichorhynchus, of Mycterotlzrix; Triloba, of Teu- tophrys; Trochella, of Metopus; Turbinella, of Uro- centrum.

Neave(50) or Schulze(58) m a i be consulted by the reader interested in knowing the identity of the organisms whose generic names comprise the senior homonyms involved here or in any of the cases listed in Table I.

3. Use of a junior synonynz as the replacemeizt name When species believed to be congeneric have been

brought together in such a way that two or more generic names are thrown into synonymy, the next oldest synonymous name must be used if the oldest generic name of the combination has fallen as a junior homonym. This is why, as has been intimated elsewhere (49), it would be inadvisable for an “armchair nomen- claturist” to sit down with a copy of Neave and propose new names for all apparent cases of homonymy. Among the genera of ciliates under present consideration several have new names created unnecessarily, generally because the authors failed to study the taxonomy of the species involved. I n fairness it should be mentioned, however, that in some cases the older synonyms have not been in existence (i.e., they were considered as names of separate genera) at the time of proposal of the new name. The following genera of ciliates may be listed here; others for which no replacement names have ever been proposed may be found in Table I:

Arachnella Kent, 1882, and Ararhnidiumia Strand, 1928, for Arachnidium, fall as synonyms of Strobilidi- um; Aulaxella Strand, 1928, for Aulax, falls as synonym of Cristigera; Craspedonotana Strand, 1928, for Craspedonotus, as synonym of Homalozoon; Prooxy- t r kha Poche, 1913, for Trichogaster, as probable synonym of Urostyla, if the organism involved is recognizable at all; Trirhoderum Strand, 1942, for Tricho- pclma, as synonym of Leptopharynx. Orthodonella, for Orthodon, would fall as a synonym of Rhabdodon were not the latter a homonym itself; thus Ortho- donella is left as the oldest available synonym.

Hallezia Sand, 1895 (suctorian), preoccupied by Hallezia Kalenati, 1848 (trichopteran) and Hallezia Girard, 1893 (nemertean), car, be made to fall as a synonym of Paracineta Collin, 1911, by proper choice of type species, none having been chosen to date. Since this genus of Sand’s(55. p. 161) has had an uncertain taxonomic status for over half a century, I am here

deliberately designating as type the first and the most recognizable of the originally included species, Hal- lezia multitentaculata Sand, 1895 (= Paracineta mul- titeiztaculata in Collin). Specialists in suctorian taxonomy must determine the fate of the only species described as a member of the genus Efallczia since 1899, H . scottocalani Sewell, 1951 (59) .

Three additional cases are of interest because they involve controversial interpretations of the degree of taxonomic interrelationship among the several species involved in each instance. Kahl(36,37) considered Belonophrya AndrC, 1912, to be a synonym of ilctino- bolus Stein, 1867; Odontochlawzys Certes, 1891, a synonym of Chilodon Ehrenberg, 1834; and Spayo- tricha Entz, 1879, as a possible synonym of Lcnzbus Cohn. 1866. If Kahl’s taxonomic conclusions are rigidly followed, Actinobolina Strand, 1928, replacement name for Actinobolus, would have to fall as a synonym of the little-known name Belonophr?la; Chilodonella Strand, 1928, for Chilodon, would have to fall as synonym of the generally unheard of Odon- tochlamys; and possibly Cohnilembus Kahl, 1933, for Izemltirs, would have to fall as synonym of the rare name Sparotrkha. I have reexamined the original papers involved and believe that there is a possibility of legitimate taxonomic independence of the first two of these genera(2.10). And I agree with Kahl con- cerning the doubtfulness of the Sparotrkha-Lei?iltzis synonymy (3 7 ) .

Further pursuit of these particular problems is beyond the scope of the present paper; but, as long as there is reason to consider the independence of the genera involved, it is not in the best interests of stability to reduce to synonymy the quite well-known replacement names universally accepted to date.

4. Creation of a new name jor the preorrupird nanir When the junior homonym has not itself fallen as a

junior synonym, or no other recognized synonym is available, the homonym should be replaced. for the sake of clarity and stability. I n this category are some five dozen genera of ciliates whose names proposed during the past century and a half have, with- out doubt quite inadvertently, been identical to names already in use for various metazoan or, occasionally. other protozoan groups: the genera involved are those whose new names are marked with an asterisk in Table 1. About half of these preoccupied names have been discovered and replaced from time to time in the past: the other half, to the best of my knowledge, are being treated nomenclaturally for the first time i n this paper.

Before proceeding with the formal proposal of new generic names, two additional comments pertinent to the over-all subject of homonymous ciliate names will be made:

New C;~:NERIC NAMES 01’ CILIATES 2 7 1

(1 ) Some ciliate genera bear names which are senior homonyms. In such cases the organisms bearing the junior homonyms, only, are in need of new names, a responsibility beyond the scope of the present work unless both names are of ciliates. Interestingly enough, however, this is the situation in regard to seven names : Balnntidiopsis, Buetschliella13 Disco- p h y a , Paraglauconaa, Rhynchophrya, Spathidiopsis, and Spirnfiluwz (see 17 , and Table I ) . Members of other protxoan subphyla are implicated in several additional cases, and there are at least three instances of the involvement of metazoan genera. Two of the latter cases may be mentioned as examples: Cleve- landella Resser, 1938, name of a genus of fossil trilo- bites, is preoccupied (by only sixteen days!) by the name of a heterotrich ciliate, Clevclandella Kidder, 1918 (itscll a replacement name: see Table I ) ; and OnJlc haspis Raymond, 1925, another trilobite genus, i > preoicul)ietl by the hypotrich Onychaspis Stein, 1859.

( 2 ) Occasionally the generic name of a ciliate has been changed in error. A subsequent reviewer has t onsidered either thst the originally proposed name was a junior homonym when, in fact, it was not; or that it had been originally spelled a certain way when, in reality, it had not. Or a substitute name has been proposed when an earlier published replacement name for the oriqinal homonym already was available, un- known to the third party. A brief listing of such cases may be of value, since some fairly well-known genera of ciliates are Involved and certain of the errors are still being maintained in the current literature. These cases are in addition to those considered above, in section 3.

d4ncistrunza Strand, although popularly used, is an unnecessary substitution for the perfectly legitimate name . lncistrum Maupas (48,50,65). Charonella Hhatia is an unnecessary replacement name of the preoccupied Charon, since Charonina Strand predates 1Shatia’s publication( 5,65). C‘onrhophthims is the original. and therefore correct, spelling of this generic name of Stein’s( 61 ) , rather than Conchophthirius, Strand (65) notwithstanding. Cryptochilidiurn never should have been considered as a substitute for C’ryptochilum, since the latter name, proposed by illaupas(48), is not a junior homonym, as erroneously supposed by Kahl(39) and many subsequent workers. Kcntrophoros never should have been replaced by (‘cntrophorus (itself preoccupied) and, later, by Ccn- tuophorella, although Kahl(3 7,40) believed otherwise. Biitschli’s( 9) emendation, Lionotus, for Wrzesniow- ski’s( 70) Litonotus not only was unjustified but it has

Customarily this name is written “Bzitschlirlla,” but dia- critical marks are no longer to be maintained in zoological nomenclature(6). A “u” is to be replaced by ‘‘ue.”

to fall anyway because of preoccupation, in spite of its popular usage today. Sieboldiellina, Collin’s ( 13) substitute name for Discophrya Stein, 1860, was out of order, since Haptophrya Stein, 1867, already had been proposed as replacement (62 ) ; yet today many protozoologists even use Sieboldiellina as type of a subfamily, a taxonomic matter beyond further discussion here (but see 17). Spathidiella is not needed, as Kahl (35,36) supposed, as a substitute for his Spathidiodes ( 3 3 ) , since the latter, which must remain unaltered in spelling, is only close to being a homonym of Brod- sky’s (7) Spathidioides.

All unnecessarily proposed names fall as synonyms of the nomenclaturally correct names. But they serve as a source of confusion and as a continuing threat, even though an illegitimate one, to the stability of the proper generic names for the ciliates involved.

I’ROI’OSAL OF NKW N A M E S

In accordance with provisions of the lnternational Rules of Zoological Nomenclature (see 6, 30, 49) , Articles 2 5 and 34-36 of the code still in force a t the time of this writing, substitute generic names are for- mally proposed for thirty preoccupied names of ciliated protozoa. In this work, the preparation of which has extended over a period of years, the “code of ethics” has been adhered to. Conscientious attempts have been made to notify protozoologists, living or presumed to be living, who have proposed preoccupied names in their original descriptions of new genera, of such a fact, in order that they may publish replacement names themselves at their convenience. I n several instances the workers concerned have authorized me to rename the genera in their behalf. In four cases repeated attempts to get in touch with the original authors have failed, and I have proceeded to propose the new names with, I believe, adequate justification. Incidentally, Dr. J. Kormos (personal communication) will have a substitute name (Caracatharina) for the suctorian genus Catharina in publication by the summer of 1960, so he has kindly permitted me to include it here as well. A similar situation exists with respect to the gymnostome Faurea, being re- named Ciliofaurea by Dr. J. Dragesco (personal communication). Thus replacement names for these two genera appear in Table I.

I n general my new names are of one of two kinds. Either they are partially or wholly patronymic in na- ture, i f appropriate, euphonious, and not already preoccupied. Or they are very much like the original name, generally with a change only in the suffix, for sake of easy recognition by persons already acquainted with species bearing the preoccupied generic name. I purposely have avoided proposing any name which, to

272 NEW GENERIC NAMES OF CILIATES

ThCTiE I. Geiicra of eili:rtcd protozoa whicli havc had preoccupied iiaiiies, with iiidiratioii of t t ic proper rc~1)I:iecwiciit: citlier a iiew iiainc [riiarked with a11 asterisk ( * ) I or t i v:rlid syiioriym, ~ I i c i i available.

Order to which geuus belongs Preorcupied iianie Rcplnceineiit iiaiiic

ASTOM AT 1 I IA4

APOHTOMAT1I)A

THIGhIOTRI('H1DA

PERTTRICHIDA

NLW GENERIC NAMES 01' CILIATES 2 73

my knowledge, already is in use in the plant kingdom. Becker's (3) comments regarding generic homonyms in bacteriology and protozoology are pertinent in this regard.

Jn the present section the genera are arranged alphabetically with respect to the new names being proposed. In Table I they are arranged alphabetically according to their preoccupied names, and are placed in groups based on a sequence of ciliate orders recognized in a new classification scheme of the Ciliophora (14-17). Each account is kept as brief as possible, but necessary citations to original works are made. 41~0, in accordance with Article 25 and with common practice among modern workers in vertebrate and hiqher invertebrate taxonomy, the type species, in Imth its old and new combination, is given in every case. In fact, names of all included species recognized today are given here in their new combination, for the sake of completeness. Interestingly enough, 90% of the genera were monotypic at the time of their original description, and 60% of them stilI contain but a single species.

Familial names are involved in five cases in which the junior homonym happened to be the name of the type genus. New names are properly proposed for these taxa, three families and two subfamilies.

Full bibliographic references to the senior generic homonyms, if desired, may be found in such nomenclatural indices as Neave( 50) and Schulze( 58). I have checked all of them myself, but their citation is omitted here in the interest of brevity.

Actinocyathula n. nom. for Actinocyathus Kent, 1882, suctorian(41, p. 848), preoccupied by Actino- cyathus d'orbigny, 1849, coelenterate. Type species, by monotypy, Actinocyathzis cidaris Kent, 1882 (= :lctinocyathula cidaris4 n. comb.).

Ascampbelliella n. nom. for Craterella" Kofoid B Campbell, 1929, tintinnid (44. p. 194), preoccupied by Craterella Schrammen, 1901, sponge. Type species, by original description, Craterella urceolata (Os- tenfeld, 1899) Kofoid Pr Campbell, 1929 (= Ascanzp- brlliella urceolata n. comb.). The new generic name

-I An earnest plea is made here that future workeis not write the name of this species as "Actinocyathzila (Act inocyathn,) tidaric." The position (and placement of the name in parentheses) bctween the generic and specific names is preempted by the subgeneric name, and the subgeneric name only (Article 10; see 30). But many protozoologists, including Kahl and other well-known taxonomists, consistently have broken this important nomenclatural rule, placing here, in parentheses, the name of the genus to which the species formerly belonged or from which i t was transferred.

" N o t to be confused with Craterella Dons, 1942, foramiiii- feran, replaced by Crateriola Strand, 1943(67).


is proposed, with explicit permission via personal communication, in honor of Dr. A. S. Campbell, well- known authority on the bionomics and systematics of the Tintinnida.

In addition to the type, nine other species are included in this genus. In the new combination their names may be listed as follows: Ascampbelliellz acuta (Kofoid Pr Campbell. 1929): A . armilla (K. Pr C., 1929); A . atlantira (Busch, 1948); A . obscura (Brandt, 1906) : A . oxyura (Jorgensen, 1924) ; A . parifica (Busch, 1948) ; .4. protuberans (K. Pr C., 1929); A . retusa (Hada, 1935); A . torulata (Jorgen- sen, 1924).

This genus is type of a subfamily(44) the name of which must be changed accordingly, since family- group names based on generic names falling as junior homonyms must be replaced. Therefore I here propose Ascampbelliellinae n. nom. as replacement for Chterellinae Kofoid Pr Campbell, 1929.

Bizonula n. nom. for Bizonc Lepsi, 1926, hymenostome(46, p. 764), preoccupied by Bizonc Walker, 1854, lepidopteran, and Bizone Simon, 1903, arachnid. Type species, by monotypy, Bizone parva Lepsi, 1926 (= Bizonula pnrva n. comb.) The new generic name is proposed with explicit permission, via personal communication, from Dr. J. Lepsi, Roumanian protozoologist who has long been interested in studies on the morphology and ecology of ciliated protozoa.

Bursellopsis n. nom. for Bursella Schmidt, 192 1, trichostome( 57, p. 1 ) , preLccupied by Bursella Tur- pin. 1828, rotifer. Type species, by monotypy, Bur- sdla spumosa Schmidt, 192 l (= Bursellopsis spumosa n. comb.). Other included species are: Bursellopsis gavgameEEac (FaurC-Fremiet, 1924) ; B. oligobursa (von Gelei, 1954) ; B. truncata (Kahl, 1927).

Buschiella n. norn. for Zmperfecta Busch, 1948, tintinnid(8, p. 132), preoccupied by Zmperfecta Mlhely, 1935, hemipteran. Type species, by monotypy, Znaperfecta subantarcticum Busch, 1948 (= Buschiella subantarrtica n. comb.). The new generic name is proposed in honor of Dr. W. Busch, authority on several major groups of spirotrich ciliates. I have been unable to get in touch with Dr. Busch, although repeated attempts have been made to do so.

Cardiostomatella n. nom. for Cardiostoma Kahl, 1928, hymenostome(34, p. I O I ) , preoccupied by C'ardiostoma Sandberger, 1872, mollusc. Type species, by monotypy, Cardiostoma vernaiforme Kahl, 1928 (= Cardiostoinatella aermiformis n. comb.).

Dellochus n. nom. for Lorhus Delphy, 1938, heterotrich( 19, p. 66) , preoccupied by Lochus Douglas & Scott, 1868, hemipteran. Type species, by monotypy, Lochus nzucicola Delphy, 1938 (= Dellochus muckola n. comb.). The new generic name is proposed in honor of Dr. Jean Delphy, describer of a

variety of new genera and species of ciliated protozoa. I have been unable to get in touch with Dr. Delphy, although repeated attempts have been made to do so.

Diceratula n. nom. for Diceras Eberhard, 1862, gymnostome (2 1 ) preoccupied by Diceras Lamarck, 1805, mollusc. Type species, by monotypy, Diceras viridans Eberhard, 1862 (= Diceratula viridans n. comb.). Second included species, much more ade- quately described (35.36) : Diceratula Birornis (Kahl, 1930).

Discomorphella n. nom. for Disromorpha Levan- der, 1893, odontostome(47, p. 5 5 ) , preoccupied by Discomorpha Dejean, 183 5, coleopteran. Type species, by monotypy, Discomorpha pectinata Levander, 1893 (= Discomorphella pectinata n. comb.).

Much as in the case of z4scampbelliella, treated above, Discomorphella i s type of a family( 53) the name of which must be changed accordingly, since familial names based on generic names falling a5 junior homonyms must be replaced. Therefore I here propose Discomorphellidae n. nom. as replacement for Discomorphidae Poche, 1913.

Discosoinatella n. nom. for Discosoma Swarczew- sky, 192S, suctorian(68, p. 52), preoccupied by Dis- cosoma Rueppel Pr Leucliart, 1828, coelenterate. Z h - cosoma Perty, 1833, arachnid, and Discosoma I'aetal, 1875, mollusc. Type species, by monotypy, Zhsco- soma tenella Swarczewsky, 1928 (= Discosomatr~lla tenclla n. comb.).

Epalxella n. nom. for Expalxis Roux, 1893, odontostome( 54, p. 596), preoccupied by Epalxis Coss- mann. 1889, mollusc. Type species, by monotypy, Expalxis mirabilis Roux, 1899 (= Epalxella mirabilis n. comb.).

In addition to the type, six other species are included in this genus. In the new combination their names may be listed as follows: Epalxella antiquorum (Penard, 1922) ; E . bidens (Kahl, 1932) ; E. elliptica (Penard, 1922); E. exigua (I'enard, 1922); E . striata (Kahl. 1926); E . triangzila (Kahl, 1932).

As in the case of Disromorphella, treated above, Epalxclla is type of a family(69) the name of which must be changed accordingly, since familial names based on generic names falling as junior homonyms must be replaced. Therefore I here propose Epalxel- lidae n. nom. as replacement for Epalcidae Wetzel, 1928.

Grandoria n. nom. for I,agcnclla R. & L. Gran- dori, 1934," trichostome(26, p. 244), preoccupied by Lagenella Ehrenberg, 1834, flagellate, Lagenclla Farre,

6 Curiously enough, Lagenella and several other ciliate genera, as well, were described as new in a paper which obviously was written first bu t which appeared in print a year later than a much larger, monographic work. The nomenclature involved must date from the publication which appeared first, of course (cf. 26 and 2 7 ) .

NEW GENERIC NAMES OF CILIATES 275

183 7 , bryozoan, and Lagenella Rehberg, 1880, sporo- zoan. Type species, by monotypy, Lagenella aculeata R. Pr. L. Grandori, 1934 (= Grandoria aculeata n. comb.). The new generic name is proposed, with explicit permission via personal communication, in honor of Ilr . Luigia Grandori and her husband, the late Dr. Kemo Grandori, whose joint contributions to our knowledge of soil protozoology are well known.

As in the cases of Disromorphella and Epalxella, treated above, Grandoria is type of a family the name of which must be changed accordingly, since familial names based on generic names falling as junior homonyms must be replaced. Therefore I here propose Grandoriidae n. nom. as replacement for Lagenellidae R. Pr L. Grandori, 1935, a name which, itself, had already been published as a replacement for Cen- trostomatidae R . & L. Grandori, 1934( 17,26,26a).

Grube re l l a n. nom. for Stylocoma Gruber, 1884, hypotrich( 28, p. 525) , preoccupied by Stylocoma Lioy, 1864, dipteran. Type species, by monotypy, Styloroma ovijormis Gruber, 1884 (= Gruberella ovijormis n. comb.). The new generic name is proposed in honor of A. Gruber, in recognition of his significant contributions to protozoology seventy-five years ago. Second included species: Gruberella adriatira (Kies- selbach, 1937).

H i s t r i cu lus n. nom. for Histrio Sterki, 1878, hy- patrich( 63, p. 56) , preoccupied by Histrio Fischer, 1813, fish, Histrio Gristl, 1848, coleopteran, and Histrio I’feiffer, 1854, mollusc. Type species, by monotypy, Histrio steinii Sterki, 1878 = Histrio hirtrio (0. 1:. IIiiller, 1786) Kent, 1882 (= Histricu- lus histrio n. comb.).

In addition to the type, eight other species are included in this genus. In the new combination their names may be listed as follows: Histriculus acumina- fus (Marshall, 1886) ; H . cornplanatus (Stokes, 1887) ; I I . erethisticus (Stokes, 1887) ; H . hyalinus (V6rGsvary, 1950) ; H . musrorum (Kahl, 1932), possibly a synonym of the following species; H . similis (Quennerstedt, 1867); H . sphagni (Stokes. 1891); I t . vorax (Stokes, 1891).

J epps i a n. nom. for Chattonella Jepps, 1937, apos- tome( ? ) (32, p. 610), preoccupied by Chattonella Biecheler, 1936, flagellate. Type species, by monotypy, Chattonella ralani Jepps, 1937 (= Jeppsia ca- lani n. comb.). The exact taxonomic affinities of this ciliate are uncertain( 17,32). The new generic name is proposed, with explicit permission via personal communication, in honor of Dr. Margaret Jepps, noted authority on the protozoa, particularly the sarcodinids.

Kahliella n. nom. for Kahlia Horvath, 1932, hypotrich(31, p. 424). preoccupied by Kahlia Ashmead. 1900, hymenopteran. Type species, by monotypy, Kahlia acrobates Horvath, 1932 (= Kahliella arro-

bates n. comb.). I have been unable to get in touch with l)r. Janos Horvath, Hungarian protozoologist who is author of the preoccupied name, although repeated attempts have been made to do so. Additional included species: Kahliella barilliformis (von Gelei, 1954); K . costata (Kahl, 1932).

Kopper ia n. nom. for Malarosoma Kopperi, 1937. gymnostome(45, p. 21) precccupied by Malarosonia Huebner, 1820, lepidopteran, Malarosoma Dejean. 1835, coleopteran, and Malacosoma Man, 1879, crustacean. Type species, by monotypy, Malarosoma in- testinale (= Kopperia intestiizalis n. comb.). The new generic name is proposed in honor of Dr. A. J. Kopperi, Finnish protozoologist who lost his life in 1943 during the war.

Niemarsha l l ia n. nom. for Marshallia Nie & Ch’eng, 1947, tintinnid( 51, p. 69)? preoccupied by Marshallia Zittel, 1879, sponge, and Marshallia Doherty, 1889, lepidopteran. Type species, by monotypy, Marshallia aperta (Marshall, 1934) Nie & Ch’eng, 1947 (= Xiemarshallia aperta n. comb.). The new generic name is proposed in honor of Dr. Dashu Nie, a Chinese protozoologist whose present whereabouts I have been unable to determine. Letters sent to former addresses have been returned unopened.

Opisthostoinatella n. nom. for Opisthostonzum Ghosh. 1928, trichostomr( 25, p. 383), preoccupied by Opisthostomum Schmarda, 1859, turbellarian. Type species, by monotypy, Opisthostomum bengalcnsis Ghosh, 1928 (= Opisthostomatella bengalcnsis n. comb.). The exact taxonomic affinities of this ciliate are uncertain(5,17,37).

Polydiniella n. nom. for Polydinium Kofoid, 1935, entodiniomorphid (43, p. 502), preoccupied by Poly- dinium Kofoid & Swezy, 192 1, dinoflagellate. Type species, by monotypy, Polydinium mysoreurn Kofoid, 1935 (= Polydiniella mysorea n. comb.). I t might be argued, especially by the phycologist, that the senior homonym is the name of an algal plant (dinoflagellate, a phytomastigophoran) and thus does not preoccupy the use of Polydinium as an animal (entodiniomorphid ciliate). But also any systematic biolo- gist has the right to consider both of these groups as members of the single animal phylum Protozoa; in which case the homonymy must be relieved.

As in the case of Ascampbelliella, treated above. Polydiniella is type of a subfamily(43) the name of which must be changed accordingly, since family- group names based on generic names falling as junior homonyms must be replaced. Therefore I here propose Polydiniellinae n. nom. as replacement for I’oly- diniinae Kofoid, 1935.

Polymorphel la n. nom. for Polymorpha Dogiel. 1929, gymnostome(20. p. 322), preoccupied by Poly- morpha Soldani. 1791, foraminiferan, and l’olynorpha


Burgeff , 1826, lepidopteran. Type species, by monotypy, Polymorpha ampulla Dogiel, 1929 (= Poly- morphella ampulla n. comb.). Second included species: Polymorphella pellucida (Hsiung, 1936).

Pottsiocles n. nom. for Pottsia Chatton & Lwoff, 1927, suctorian(l1, p. l ) , preoccupied by Pottsia Ryder, 1885, coelenterate. Type species, by monotypy, Pottsia infusoriorum Chatton & Lwoff, 1927 ( = Pottsiocles infusoriorurn n. comb.). The new generic name (in translation meaning “to the glory of Potts”) is proposed with explicit permission, via personal communication, from the original junior author, Dr. AndrC Lwoft‘. Second included species: Pott- siorlps parasiticus (Nozawa, 1939).

Pyxidiella n. nom. for Pyxidium Kent, 1882, peri- trich(41, p. 666) , preoccupied by Pyxidium Leuckart. 1856, coelenterate. Type species, more or less by original designation, with subsequent affirmation herein, Pyxidium cotkurnoides Kent, 1882 (= Pyxidiella cothurnoides n. comb.).

In addition to the type, twenty-two other species are included in this genus. I n the new combination their names may be listed as follows: Pyxidiella acaula (Kahl, 1935): P. arboricola (Biegel, 1954); P. aselli (I’enard, 1922) : P . asymmetrica (Biczbk, 1956) ; P. bosminae (Kahl, 1935) : P. canthorampti (Penard, 1922); P . rollare (Kahl, 1935); P . curvicaula (Pen- ard, 1922) ; P. fabrei (FaurC-Fremiet, 1905) ; P. hebes (Kellicott, 1888) ; P. henneguyi (FaurC-Fremiet, 1905): P. inclinans (de Fromentel, 1874); P. in- aaginnta (Stokes, 1887) : P . Rahli (Nenninger, 1948) ; 1’. longicolla (Biegel, 1954) ; P. minuta (Lust, 1950) ; P. nutans (Stokes, 1889) ; P. pisficolae (Nenninger, 1948); P. stammeri (Lust, 1950); P . urceolata (Stokes. 1886) : P. ventriosa (Nenninger, 1948) ; P. vernalc (Stokes, 1887).

Sathrophilus n. norn. for Saprophilus Stokes, 1887, hymenostome( 64, p. 247), preoccupied by Saprophi- lus Streubel, 1839, coleopteran. Type species, by monotypy, Saprophilus agitatus Stokes, 1887 (= Snthrophilus agitatus n. comb.). The new generic name possesses exactly the same meaning as the former name.

In addition to the type, six other species are included in this genus. I n the new combination their names may be listed as follows: Sathrophilus chlo- rophagus (Kahl, 1931) ; S. mobilis (Kahl, 1926) ; S. musrorum (Kahl, 1931): S. ovatus (Kahl, 1926); S. ozriformis (Kahl, 1926), probably a synonym of Tetra- hymena pyrijornzis; S . putrinus (Kahl, 1926).

Schewiakoffia n. nom. for Maupasia Schewiakoff, 1892, gymnostome(56, p. 552), preoccupied by Mau- pasia Vignier, 1886, polychaete. Type species, by monotypy, Maupasia paradoxa Schewiakoff, 1892 (1 Srhewiakofia paradoxa n. comb.). The new ge-

neric name is proposed in honor of W. Schewiakoff, whose monumental work, in Russian( 56a) , on the biology and systematics of the holotrich ciliates will always remain a classic in its field.

Sciurula n. nom. for Sriurella Kopperi, 1937, gymnostome(45, p. 24), preoccupied by Sriurella Allman, 1883, coelenterate. Type species, by monotypy, Sriurella intestinalis Kopperi, 1937 (= Sciurula intestinalis n. comb.).

Spirofilopsis n. nom. for Spirofilum von Gelei, 1944, hypotrich(23, p. 144; see also 24, p. 324), preoccupied by Spirofilum von Gelei, 1929, a hypotrich ciliate also. Type species, by monotypy, Spirofilum tubic-ola natans von Gelei, 1944 (= Spirofilopsis tubi- colonatans n. comb., with the specific name here emended to conform to the Rules).

Von Gelei’s(22, p. 165) first Spkofilum falls ;I

synonym of Hypotrichidium Ilowaisky, 192 1, if one agrees with the taxonomic conclusion reached by Kahl (38) . Von Gelei himself(23,24) concurred with Kahl’s decision. His resurrection of the genus Spiro- filum, however, to contain a new species not congeneric with the single species of his first paper contravenes the Rules ( c . , ~ . , Article 30). His second use of Spiro- filum(23) must, then, be considered as erection of a new genus. But. since the name is preoccupied by his own earlier employment of it, it falls as a iunior homonym and must be replaced.

Trichopodiella n. nom. for Trichopus C1apari.de Pr Lachmann, 1859, gymnostome(l2, p. 338). preoccupied by Trirhopus Shaw, 1803, fish, Trichopus C. M., 1834, arachnid, and Trichopus Haan, 1835. crustacean. Type, by monotypy, Trichopus dysteria Clapari.de 8 Lachmann, 1859 ( L Trirhopodiello dysteria n. comb.). Second included species: Trirho- podiella larhmanni ( FaurC-Fremiet, 1957).

Turaniella n. nom. for Turania Brodsky, 1925, hymenostome( 7, p. 42 ) , preoccupied by Turania Ragonot, 1890. lepidopteran, and Turania Bethune- Baker, 1914, also a lepidopteran. Type, by monotypy, Turania vitrca Brodsky, 1925 (= Turaniella vitrea n. comb.).

Urospinula n. nom. for Urospina von Gelei, 1943. hypotrich( 23, p. 137), preoccupied by Urospina Sell- nick, 1932, arachnid. Type, by designation herein, Urospina biraudata von Gelei, 1944 (= Urospinula biraudata n. comb.). Second and third included species: Urospinula ra/rihia (von Gelei. 1944), ll. sinis- troraudata (von Gelei, 1944).

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2 78 ASTOME CILIATES FROM OLIGOCHETES

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J . PROTOZOOL. 7 ( 3 ) , 278-289 (1960)

Observations sur quelques Cilihs astomes des OligochLtes du lac d’Ohrid (Yougoslavie) . 11. Familles des Hoplitophryidae (Hoplitophryinae et

Maupasellinae) , des Intoshellinidae et des Anoplophryidae.

PIERRE DE PUYTORAC

Laboratoire de Zoologie, Biologie cellulaire, I , Avenue Vercinge‘torix, Clermont-Ferrand, France, et Statton Biologique, Ohrid

SYNOPSIS. Seven species of astomatous Infusoria from the alimentary tract of the Oligochaeta from Ochrida Lake are described. One of them, belonging to Hoplitophrya, shows many transitional characteristics between luxtaradiophrya and Hoplitophrya and proves the coherence of the group Radio- phryinae-Hoplitophryinae-Mesnilellinae. Two species are Mau- pasellinae, parasites from Glossoscolecidae. Buchneriella and Maupasella, both parasites from Criodrilus lacuum, a cosmo-

Sf. des Hoplitophryinae. Hoplitophrya georgevitchi de Puytorac, 1958.

ETTE espkce ne nous a CtC rCvClCe que par trois C exemplaires endoparasites de I’intestin moyen d’un Lumbriculus sp. provenant du marais qui cein- ture le lac.

De caractkre typique du genre, cylindroide, mesur- ant 540-600 p de longueur, chez les individus non en division, pour 72 p de diamktre, ce CiliC a un macro- nucleus grtle, CtirC d’un bout & I’autre du corps, un micronucleus petit, sphkrique et mCdian, et il porte une rangCe longitudinale de 12 A 15 vacuoles con- tractiles.

Le cytosquelette (Fig. 1) surtout, mkrite une atten- tion particulikre et en fait une espkce d’un interkt

politan worm, are also present in C . ohridensis, endemic species coexistent with the preceding, a t Ohrid. Two other species belong to Intoshellina. A discussion about the actual systematic state of Intoshellinidae is given, affinities of this family remaining uncertain. The two last species described are a typical Anoplophrya and a representative cf a new genus, Corlissiella, having many morphological and biological simi- larities to the primitive thigmotrichs. Heterogeneity of the Anoplophryidae is discussed.

certain. Son Ctude, bien qu’incomplkte, faute de ma- tCriel, nous a nCanmoins montrC qu’il est constituk d’un appareil en forme de “V” dissymittrique, situk dans le plan sagittal, coiffant l’apex de la cellule et dont la branche la plus longue (52 p ) est formke par l’assemblage de 6 spicules, situCs presque bout A bout, coalescents a I’une de leurs extrkmitks, et se relayant vers la partie postkrieure de telle sorte que leurs terminaisons sont respectivement dCcalkes les unes par rapport aux autres. Le spicule le plus antCrieur, paraissant CdifiC hi-mtme par deux filins torsadCs, atteint a peu prks la moitie de la longueur de l’en- semble, auquel se raccorde, en angle 1Cgkrement obtus, l’autre branche du “V” trks courte (6 p ) . Des fibres, insCrCes d’un seul c8tC de 1’ClCment en T,” (le droit par seule convention), et non point rkparties de part

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