Resolution of Persistent Taxonomic and Nomenclatural Problems Involving Ciliate Protozoa Assignable to the Astome Family Haptophryidae Cepede, 1923

TAXONOMY AND NOMENCLATURE OF HAPTOPHRYIDAE 265

interrelation in various microorganisms, in McElroy, W. D. & and nitrogen metabolism of Tetrahymena geleii. J . B i d . Chent. Glass, H. B , Antino rlcid Metabolism, Johns Hopkins Press, 198, 753-64. Baltimore, 335-46. 24. ___ 1956. Free and nonprotein amino acids in Tetra-

2 2 . Warnock, L. G. & van Eys, J. 1962. Normal carbohy- hymena pyrifornzts. Arch. Biochem. Biophys. 62, 70-3. drate metabolism in Tetvahymena pyriformis. J . Cellular 25. Youngburg, G . E. 1921. The removal of ammonia from Coinp Phyhiol. 60, 53-60. urine preparatory to the determination of urea. J . Biol. Cherrt.

45, 391-4. 23. Xu. C. & Hog::, J . F . 1952. The amino acid composition

J PROTUZOOL 12(?) , 263-273 (1965)

Resolution of Persistent Taxonomic and Nomenclatural Problems Involving Ciliate Protozoa Assignable to the Astome Family Haptophryidae Cepede, 1923*

JOHN 0. CORLISS, PIERRE DE PUYTORAC, JIRI LOM

Department of Biological Sciences, University o j Illinois at Chicago Circle, Chicago, Illi?tois. C S.A ; Laboratoire de Zoologie, L‘niversitt de Clermont-Ferrand, Clermont-Ferrand, France;

Protozoological Laboratory, Czechoslovak Academy o f Science, Prague, Czechoslovakia

SYSOPSIS. Confusion has long existed in the literature concerning both the taxonomy and the nomenclature of ciliates belongin:: to the astomatid family Haptophryidae. Most of the controversy has centered around the names and concepts of “Haptophuya” and “Sieboldiellina.” The latter name, widely used for certain species found as parasites (endocommensals) of turbellarians, must fall as a junior synonym of the former, which has commonly been restricted to the astomatous ciliate parasites (endocommensals) of various amphibians But the species of “Haptophrya” are then left without a generic vehicle, since Haptophrya Stein, 1867, must, in effect, be used to replace Sieboldiellina Collin, 1911, as the proper generic name of the turhellarian parasites Fortunately a name is available for the amphibian species: Cepedietta Kay, 1942.

The problem is further complicated because of the recognition of subfamilial groups, the name of one of these having

MONG mouthless ciliates presently assigned to A the holotrich order Astomatida are members of a curious group commonly found as endoparasites of certain turbellarians and amphibians. The group com- prises a family. the Haptophryidae. Although these large and quite distinctive astomes have been the subject of numerous studies( 1-10,18-20,22-24,26-43) since their discovery 125 years ago, most workers have been concerned with their general morphology, detailed cytology, or their parasitism, treating rather super- ficially their taxonomy and neglecting almost entirely the nomenclatural problems involved. Recently two comprehensive papers have been published (26,33) which have added greatly to our understanding of the general taxonomy of the whole group; a note has also appeared( 14, p. 142) concerning certain pertinent nomenclatural aspects. However, a deep-seated confusion still exists which the present paper will attempt to resolve in view of the increased interest today in

% T h e support of a National Science Foundation Grant ( to the senior author) is gratefully acknowledged. We also appred- ate the aid of Miss Alice Boatright in arrangement of the figures.

been formed from Sieboldiellina and having become associated with certain species parasitic in turbellarians. With the realiza- tion that, by the international rules of zoological nomenclature, Haptophrya must be used with reference to these particular turbellarian parasites, the subfamilial name associated with the amphibian forms must be changed. We propose Cepediettinae n. nom. in solution of this particular problem. We recognize a third subfamily, Ckpkde’s Lachmannellinae, to contain the three remaining acceptable genera comprising the family Hapto- phryidae: Anndophrya, Lachmanndla, and Steinella.

At the generic and specific levels numerous errors of a nomenclatural nature have been committed in the older literature, many unwittingly perpetuated in recent papers. These are all corrected in the present work. For the sake of future clarity we have included a series of figures, both original and from the literature, and have designated neotypes of the two principal species involved in the overall controversy.

these remarkable ciliates and before possibly even further complications are introduced into the literature.

TAXONOMIC PROBLEMS

I n this section we shtall deliberately avoid treatment of nomenclatural aspects in the history of the involved organisms, reserving such considerations for a subsequent section oi our paper. One of the principal sources of confusion in past publications on the Haptophryidae has been the attempt by workers to discuss simultaneously the two interrelated but separate problems of taxonomy and nomenclature. For the sake of clarity as well as brevity only the most important and most significant events in the systematics of the group will be considered here.

The Turbellarian Parasites

I n 1839, von Siebold allegedly1 described a new species of Opdina, as 0. planaviarum, from the digestive tract of the fresh-water triclad Planaria torva. Presumably no figures were included. In 18452 and 1848, von Siebold made several refer- ences to characteristics of this endoparasitic ciliate without illustration. Stein(40), in 1860, erected a new genus to contain von Siebold’s organism, properly removing i t from the genus Opdina. The name he proposed, Dhcophrya, was preoccupied,

266 TAXONOXY AND SOMENCLATURE OF HAPTOPHRYIDAE

TAXONOMY AND NOMENCLATURE OF HAPTOPHRYIDAE 26 7

Figs. 1-3a, b. Drawings of Haptophrya planariarum (von Siebold, 1839) Stein, 1867 (subfamily Haptophryinae).

Fig. 1. Various views of the ciliate in photographic reproduction of original figures by Schultze(35) published 114 years ago. Note distinctive features: broad thigmotactic area (adhesive sucker), contractile vacuole canal, macronucleus, unequal fission without chain-formation. Fig. 2 . Reproduction of draw-

so he renamed the genus Haptophrya in a later work(41: and see following section of the present paper).

In 1851, Schultze(35) described two new species of astomatous ciliates from turbellarians. The first, Opdina polymorpha from Planaria torva, has been considered identical with von Siebold’s species by subsequent workers. The second, 0. uncinata, from the marine worm Procerodes ulvae (= Planaria ulvae), has found a ,taxonomic home in CCpkde’s(5) genus Steinella, as the sole member of that distinctive taxon. As the earliest known figures of ciliates belonging to the familial group under discussion in the present paper, Schultze’s drawings of Haptophrya planariarum (von Siebold) and Steinella uncinata (Schultze) are reproduced in our Figs. 1 and 7. (The most recently published figures of the latter species, those of Sikora(38), are presented for comparison in our Figs. 8a, b.)

In 1859 (this date is commonly, but emoneously, given as 1858 in the literature), Clapartde & Lachmann(9) published a description of a unique species from the marine turbellarian Planaria limacina which they named Opalina recurva. CCptde ( 5 ) subsequently included this species as the only member of his genus Lachmannella. Clapartde & Lachmann’s original figure of L . recurva is reproduced in our Fig. 9.

The Amphibian Parasites

In 1879, Maupas(28) offered the first description of an

1 A frustrating mystery exists with regard to von Siebold’s first paper on his Opalina planariarum. Several authoritative early workers-such as Kent(Z)-ite “von Siebold, 1839” as the source of the original description. Yet others-such as Butschli(4)--claim that the first work on this particular organism appeared later, in 1845. With considerable difficulty we have obtained and perused a “von Siebold, 1839”(36) and have found no mention of his “Opdina.” On the other hand, the possibility remains that a second, still rarer “von Siebold, 1839” work, bearing the same or very similar title, also once existed and contained the “Opdina” account. Throughout the present paper we are taking the liberty of assuming the validity of the 1839 date, although the matter remains open to dispute.

2 A second mystery surrounds the date of publication of von Siebold’s celebrated “Lehrbuch der Vergleichenden Anatomie der Wirbellosen Thiere.” We have never been able to discover an earlier edition than one dated 1848(37), yet many works of the 19th century cite the proper date as 1845 (e.g., Biitschli’s most reliable treatise, 4). In the 1848 edition the name Opa- lina planariarum occurs several times, with no indication that it is a new species (see our preceding footnote); nor is there any indication of an earlier edition of the “Lehrbuch.” I n fact, in the Preface von Siebold(37) writes [English transla- tion], “The elaboration of this work . . . commenced in 1845, but its completion [has been1 delayed by my change of resi- dence from Erlangen to Freiburg, and partly by a pretty long sojourn of mine on the Adriatic Sea . . . ” and it is signed with the date and place as “Freiburg (in Breisgau), Feb. 27, 1848.” On the other hand, one of us (J.O.C.) has been told by a research librarian that a rare 1845 copy exists which is identical to the 1848 one!?! It should be pointed out that the principal difficulty in determining the existence and/or whereabouts of such works stems from the incomplete (or even totally lacking) citation of them in other (subsequent) older publications, a circumstance compounded by hasty copying habits of still later writers who have not bothered to check original sources first-hand.

ing by Bishop(l), showing organism in side view. Fig. 3a. Outline of body, schematically showing the general course of ciliary rows on the dorsal surface (original, by J.L.). Fig. 3b. Schematic drawing of anterior end of body, ventral surface, showing lines of ciliature in the thigmotactic area with details of their silverline system portrayed in central strip of six rows (original, by J.L.).

astomatous ciliate bearing a certain resemblance to von Sie- bold’s turbellarian parasite but found in the rectum of amphibian vertebrates, frogs of the species Discoglossus pictzls and Rana esculenta. He named the organism Haptophrya gigantea, but included no figures in his account. Later in the same year Certes( 7 ) redescribed Maupas’ interesting ciliate, leaving it in the same genus (Haptophrya) as von Siebold’s invertebrate parrasite and offering drawings of it which are reproduced in our Fig. 4.

Relatively recently, in 1942, Kay(22) described an organism from the intestinal tract of the salamander Eurycea bislineata which she considered to differ sufficiently from the previously described ciliates from both planarians and amphibians to require a new generic designation. She proposed the name Cepedietta jibrillata for her newt astome; but very recently de Puytorac(33) has concluded that it is congeneric with Maupas’ frog parasite, thus not warranting a separate generic name. Several of Kay’s drawings are reproduced in our Fig. 5 .

Decisions at the Familial Level

CCptde(S), in 1910, set all the known species of the group under present discussion aside from other astome ciliates by creation of a family which he named the Discophryidae. Later (6) he substituted Haptophryidae for his earlier familial name and established three subfamilies: the Haptophryinae, the Lachmannellinae, and the Steinellinae.

Bishop( 1) subsequently split Ckptde’s third subfamily into two subfamilies, using Sieboldiellinae (sic) as the name of the new 0ne.3 In the most recent taxonomic treatments of the whole group, Lom(26) and de Puytorac(33) recognized three subfamilies: Bishop’s and the first two of CCp4de’s. Textbooks of protozoology, incidentally, have never included mention of any of these subfamilies. Corliss( 14), who purposely excluded all subfamilies from consideration in his concise taxonomic monograph on the Ciliophora, listed seven genera (including three recently established by Lom, 26), as comprising the family Haptophryidae: Annelophrya, Cepedietta, Clauslocola, Haptophrya, Lachmannella, Proclausilocola, and Steinella.

NOMENCLATURAL PROBLEMS

Although taxonomic decisions are, rightly, based principally on a worker’s knowledge-preferably first-hand-of the comparative morphology of the species implicated in a given situation, the International Code of Zoological NomenclaCure(Z1) must be adhered to in the matter of proper names to attach to the various taxa involved. In the present case many proto- zoologists have committed errors of a nomenclatural nature which, unfortunately, have frequently become compounded with the passage of time. In the following paragraphs only the nomenclatural decisions of most direct significance to the problem under present consideration will be treated, and even these will be mentioned as briefly as is consistent with clarity.

3 Because Bishop’s study represents the first comprehensive investigation of von Siebold‘s species since the turn of the century, her drawing of the entire organism is reproduced in our Fig. 2 .

268 TAXONOMY A N D NOMENCLATURE OF HAPTOPHRYIDAE


Figs. 4-6a, b. Drawings of species of Cepedietta Kay, 1942 (subfamily Cepediettinae) .

Fig. 4. C. gigantea (Maupas, 1879) n . comb. Various views of the ciliate in photographic reproduction of original figures by Certes( 7) published 86 years ago. Note distinctive features: well-defined sucker, slightly posterior to anterior extremity of body, long contractile vacuole canal, compact macronucleus. Fig. 5 . C. jibrillata Kay, 1942. Drawings reproduced from

Haptophrya versus Discophr)a

When Stein(40) erected his astome genus Discophrya in 1860 to contain von Siebold’s Opalina planariarum, he was evidently unaware of Lachmann’s(25) creation of the suctorian genus of exactly the same name-Discophrya-in the preceding year. Several years later, in 1867, Stein(41) replaced his inadvertent junior homonym himself, using Haptophrya as the new name. Absolutely no criticism can be made of this commendable and lezal act. It should be pointed out that the type-species of the genus remained the organism originally described by von Siebold in 1839.

Haptophrya versus Sieboldiellina

Coll in( l l ) , in 1911, proposed an unnecessary replacement name-Sieboldiellina-for Stein’s Discophrya. This small but important error generally went unobserved, with most subsequent workers accepting Collin’s name change. Most unfortunately, however, 20th century specialists on the group, in addition to accepting the name, came to think of Sieboldiel- lina as a genus dlstinct from Haptophrya. That is, Collin’s replacement name for Discophvya was considered to represent a separate taxonomic group from that of Stein’s own earlier replacement name for Discophrya: an impossible conclusion ! Szeboldiellina Collin, 1911, having the same type-species as Haptophrya Stein, 1867, must fall as a junior synonym of the latter name. I t might well be emphasized that the failure to recognize this incontestable fact is the fault not so much of Collin but of careless subsequent investigators.

Still more serious is the fact that with passage of time the name Haptophrya has come to be associated with the ciliates found in amphibians, and the name Sieboldiellina with species from turbellarian flatworms But Haptophrya-ven if the name Sieboldiellina had never come into existence-must re- main attached to its single originally included species, which was a ciliate from a planarian host!

It might be mentioned that if Maupas(Z8) had been less conservative in the taxonomic disposition of his ciliate-the first species to be described from a non-planarian host-all would have been well. That is, if he had proposed a new generic name for his amphibian parasite, rather than Hap- tophrya which was already associated with von Siebold’s turbellarian form, then all subsequent workers probably would have followed Maupas’ lead, utilizing his distinct generic name in describing new species from newts and frogs. Haptophrya and Sieboldiellina would then have remained in rivalry as names only for the planarian astomes, with Haptophrya in- evitably winning out on grounds of priority. However, as in the case of Collin cited above, i t is not a t all the fault of Maupas that subsequent workers became confused, but their own lack of care in properly searching out and understanding the original literature, publications generally not difficult to obtain.

The status of Cepedietta

De Puytorac(33) has recently concluded that Kay’s(22) generic name Cepedietta should fall as a junior synonym of

Kay’s(22) work, emphasizing strong fibrillar system associated with the sucker. Fig. 6a. C. gigantea, dorsal view, schematic drawing showing reproduction by chain-formation typical of members of this genus (original, by P. de P.). Fig. 6b. C. gigantea, ventral view, anterior end of body, schematic drawing showing ciliary rows (purposely omitted in central strip) with emphasis on the inconspicuous but taxonomically important anterolateral secant systems (original, by P. de P.).

Haptophrya, in the sense that “Haptophrya” represents the genus of frog and salamander parasites within the iamily Haptophryidae. We concur in this taxonomic-nomenclatural decision. As proposed in our discussion below, resurrection of the thus available name Cepedietta as the genus of astome ciliates found in amphibians represents an acceptable nonien- clatural solution to this important aspect of the overall problem.

Lachmannella and Steinella

The two genera Lachmannella and Steiizella, erected by CCpede(5) in 1910, have as their respective type species. by monotypy, Lachiiiannella recurva (Claparede B Lachmann. 1859) Cbpi.de, 1910, and Steinella uncinata (Schultze, 1851) Ctpede, 1910. There has never been any particular dispute over the proper nomenclature to follow in these two cases, although the form of the names (complete with authorship and date) has often been given improperly in the literature.

Names at the familial level

CCp&de’s(d) name for the entire family, Haptophryidae. is perfectly acceptable. His subfamilial names are also available to taxonomists who wish to acknowledge differences at this level among the genera associated with the entire group. Bishop’s(1) recognition of an additional, separate subfamil>- is proper enough, but her name Sieboldiellinae (sic)-even when corrected to Sieboldiellininae-is not available: it falls as a junior synonym of Haptophryinae Cbpede. 1923, when the latter is correctly recognized as the name of the group containing ciliates solely from planarian, never amphibian, hosts. A subfamilial name for the frog and newt species must be formed from any other generic name available. ,.Zs mentioned in a subsequent section of the present paper, Ct-pedietf~ Kay, 1942, represents such a name.

RESOLUTION OF PROBLEMS

For the curious group of astome ciliates which form the subject of the present paper we essentially endorse the taxonomic conclusions recently published by de P u ~ t o r a c ( 3 3 ) . ~ Many of the changes necessitated at the nomenclatural level are so significant, however. that we believe it is important now to discuss both aspects of the situation at the same time, having pre-

.I A problem remains concerning disposition of two genera, Clausilocola and Proclausilocola, whose species have been described from the body cavity of certain land snails by Lom(26) who placed them in the subfamily Sieboldiellininae (the Hap- tophryinae of the present paper). De Puytorac(33) disazreed with this disposition. I t is the sentiment of the present authors that knowledge of the exact relationship of these mollusc parasites must await more detailed cytological information of a comparative nature. Certain of their already known characteristics would seem to separate them from the better under- stood members of the family Haptophryidae, but the extent or degree of this separation is not altogether clear. They are tenta- tively excluded from the family in the present paper.

2 70 TAXONOMY A N D S O M E N C L A T U R E OF HAPTOPHRYIDAE

sented all pertinent factual data, albeit briefly, in The name Huptophryu must be associated with the preceding sections. ciliate originally discovered by von Siebold and with

Figs. 7-10. Drawings of species of the genera Steinella Ckpkde, 1910, Lachmznnella CCpkde, 1910, and Annelophrya Lorn, 1959 (subfamily Lachmannellinae).

Fig. 7. Steinella iirtcinata (Schultze, 1SSl) C;pi.de, 1910. Views of the ciliate in photographic reproduction of original figures by Schultze(35) published 114 years ago. So te distinctive features: relatively indistinct thigrnotactic area adorned anteriorly with two hooks, contractile vacuole canal, macronucleus, unequal fission without chain-formation. Fig. 8a. S. uncinata, dorsal and ventral views and drawings of fission, reproduced from recent work by Sikora(38). Fig. Sb. S.

uncinuta, ventral view, anterior end of body showing details of silverline system in vicinity of thigmotactic area and hooks; reproduced from Sikora(38). Fig. 9. Lachmannella recurva (Claparede 8: Lachmann, 1859) Ckp&de, 1910. Reproduction of original figure by Claparede & Lachmann(9), published 106 years ago, the only drawing available in the listerature concerned with this species. Note the single hook, characteristic of the genus. Fig. 10. Annelophrya sphaevonucleata (Georgkvitch, 1950) Lom, 1959. Reproduction of original figure by Georgk- vitch(l9). Note the numerous hooks in the thigmotactic area, characteristic of the genus.


subsequent congeneric species (such as those described by GeorgCvitch( 18) and de Puytorac(31,33)). Sie- boldiellina falls as a junior synonym of Haptophrya. In other words, Haptophrya, the senior synonym, must replace Sieboldiellina, as the latter name has generally been employed in the literature. Similarly, Haptophry- inae must replace Sieboldiellininae at the subfamilial level. The single familial name, Haptophryidae, remains unaffected.

The ciliates of this family which are found in such amphibian hosts as salamanders, species probably more commonly known than those from the invertebrate planarian hosts, may not utilize the name Hap- tophrya. The only name already published which is available for them-concluding, with de Puytorac( 33), that they are all congeneric-is Cepedietta. The subfamily for this genus needs a new name; in effect, replacement of the formerly misused “Haptophryinae” is demanded. We therefore here propose Cepediettinae n. nom. as the replacement name for the “preoccupied” Haptophryinae.

Since several characterizations in the older literature must be rather extensively shifted about to fit our proposed taxonomic revision, we shall employ the word “emendation” after the names of groups so affected in the scheme given below. Other names involved in the taxonomy of the entire family need only minor repair. Such corrections will be incorporated in the overall taxonomic-nomenclatural revision given below without, for the sake of conciseness, special discussion of each case.

TAXONOMIC SCHEME FOR T H E FAMILY

In the following abbreviated synopsis of the family Haptophryidae emphasis will be placed on: succinct characterizations (limited to supra-specific taxa) ; cor- rect nomenclature, including authorship, dates, and principal synonyms ; clarification of types ; presenta- tion of key figures and designation of type-material for the three species whose genera are types of the three reocgnized subfamilies. The taxonomic-nomenclatural principles embodied in recent papers by one of us( 13- 16) are followed here without comment.

Family HAPTOPHRYIDAE CCpGde, 1923 ( 19 10) (for Discophryidae CC@de, 1910 [non Collin, 19121)

Astomatous ciliates with numerous rows of cilia which converge anteriorly a t horseshoe-shaped suture line, with two inconspicuous antero-lateral secant systems. Suture line marks anterior margin of ventral thigmotactic area or adhesive sucker well-defined a t its posterior border by either break in or distinctive

““Double-dating” is done here (and for the generic name Haptophrya) in accordance with Article 39a(i) of the Inter- national Code of Zoological Nomenclature(21) ; thus for pur- poses of priority the earlier dates prevail.

curvature of the kineties. Contractile vacuole repre- sented by long canal provided with several pores. Macronucleus oval to elongate. Unequal binary fission, with or without chain-formation. Parasitic (i.e., endocommensal) in digestive tract of amphibians and fresh- and salt-water turbellarians. Type-genus Hap- tophrya Stein, 1867 (1860).

Subfamily (1) HAPTOPHRYINAE CCpGde, 1923, emend.

(Syn. Sieboldiellininae Bishop, 1926) Suture line, at apex of body, marks anterior border

of broad and shallow thigmotactic area. Unequal binary fission without chain-formation. Found only in fresh-water planarians. Type-genus Haptophrya Stein, 1867 (1860).

Single6 genus: Haptophrya Stein, 1867 (1860), emend. (for Discophrya Stein, 1860 [non Lachmann, 18591 ;

syn. Sieboldiellina Collin, 191 1) With characters of the subfamily. Type-species: H .

planariarum (von Siebold, 1839) Stein, 1867, by monotypy (Figs. 1-3a, b) . Syns. Opalina planariarum von Siebold, 1839; Discophrya planariarum (von Sie- bold, 1839) Stein, 1860; Sieboldiellina planariarum (von Siebold, 1839) Collin, 1911.

Designation of neotype. Because of the importance of this species, type-specimens of which have apparently never existed, and in the interest of stability of nomenclature, we hereby designate as neotype the specimens on a single slide? prepared according to the Chatton-Lwoff silver impregnation technique ( 12) and deposited in the International Collection of Ciliate Type-Specimens recently established at the University of Illinois (1 7) . Such deposition in a recognized insti- tution is mandatory for neotype material(21, Art.

Other included species: H . acetabulifera (GeorgC- vitch, 1941) n. comb.; H . ohridensis (Georgkvitch, 1941) n. comb.; H . stankovici (de Puytorac, 1957) n. comb.; H . omeni (Gillespie in de Puytorac, 1963) n. comb.

75c).

Subfamily (2) CEPEDIETTINAE n. nom. (for Haptophryinae CCp&de, 1923, et auctt.)

Suture line, slightly posterior to apex of body, does not coincide with anterior border of thigmotactic area;

6 The genera Clausilocola and Proclairsilocola were also included in this subfamily by Lom(26) and were listed as members of the family by Corliss(l3), but they are excluded from the classification scheme given here. See footnote number 4 for brief discussion of this problem, full treatment of which is beyond the scope of the present paper.

7This slide is from the personal research collection of Dr. Jiii Lom and contains material taken by him from the intestinal tract of the fresh-water turbellarian Planaria torva, collected near Klatovy, Czechoslovakia during the month of April, 1955.

2 7 2 TAXOXOMY A N D SOMENCLATURE OF HAPTOPHRYIDAE

latter in form of definite sucker surrounded by thick- ened ecto-endoplasmic layer giving rise to a well- developed fibrillar system. Reproduction by chain- formation. Found only in amphibians (frogs and salamanders). Type-genus Cepedietta Kay. 1942.

Single genus: Ct7pcdzetta Kay. 1942. emend. (for Haptophiya Stein. 1867, Q S rizisuscd by

niaii? uwikel-s) IYith characters of the subfamily. Type-species: C.

gigantea (Maupas. 1879) n. comb + by subsequent designation (Figs. 4. 6a. b ) . Syn. H a p f o p h i y a gi- qanten Maupas. 1879.

Lk!!iXnation o j neotype. Because of the importance of this species. type-specimens of which have apparently ne\er existed, \ve hereby designate as rieot?ipe the specimens on a single alide’ prepared according to thil Chatton-Lwoff silver impregnation technique (12 ) and deposited in the International Collection of Ciliate Type-Specimens a t the rniversity of Illinois ( 1 7 ) . in accordance with the rules of nomenclature (21).

Other included species: C. michigunensis (IYood- head, 1928) n. comb.: C. iliiginiensis (Jfeyer. 1938) n. contb.; C. f i b d a t a Kay. 1942 (Fig. 5 ) ; c‘. plethodonis (Lipscomb in de Puytorac, 1963) n. comb. Type-specimen material is known to exist for two of these species: Kay(22) deposited a slide of C. fibril- lata in the United States Xational Museum ( S o . 36802): and de Puytorac has now deposited a slide of C. plethodonis in the International Collection at the University of lllinois.

Subfamily (3 ) LACHMASNELLISAE CCpi.de. 1923, emend.

(Syn Steinellinae C6pi.de. 1923) Suture line, near apes of body. marks anterior bor-

der of restricted, somewhat shallow thigmotactic area provided with variable number of well-developed skeletal hooks. Vnequal binary fission without chain- formation. Found in marine or fresh-water turbellarians Type-genus Lachmanrrella CCpi.de, 19 10.

n’ith characters of the subfamily, but only single hook and known only from marine host (Planaria limacina) off coast of Norway. Type-species: L. recurva (Clapari.de & Lachmann, 1859) C6pi.de. 1910, by monotypy (Fig. 9 ) . Syns. Opalina recuraa Clapa- rede E;r Lachmann, 1859; Hoplitophrya recurva (Claparkde & Lachmann, 1859) Kent, 1882.

I t is unfortunate that the type-species has never been seen again since its discovery in 1859. I n the absence of original type-specimens or of possible ~-

SThis slide is from the personal research collection of D r Pierre dc Puytorac and contains material taken by him from the intestinal tract of the frog Discoglossus pictiis collected in Tunisia during the month of rlpril, 1960.

neotype material we believe that the only published figure (the drawing by Clapari.de Pr Lachmann reproduced here as our Fig. 9 ) might well be considered as a ‘ $ type-specimen,” in our interpretation of Article 74b (concerning lectotypes) of the recently revised International Code of Zoological Nomenclature ( 2 1, p. 79).

Other included genera, also each with a single species: Stf inel la Cepi.de, 1910 (with two hooks of unequal size) ! type-species S. unciizata (Schultze, I85 1 ) Cepi.de. 1910 (Figs. 7 , 8a, b ) ; Aniaelophrya Lom, 1959 (with numerous, small spines surrounding the thigniotactic area) , type-species ‘4. sphaeronucleata (GeorgeLritch, 1950) Lom, 1959 (Fig. 10).

R E F E R E S C E S

1. Bishop, -4. 1926. Sotes upon Sirboldiellina planariarum (Siebold), a ciliate parasite o i Planaria torva. Parasitology 18, 187-94.

2 . Bush, M. 1933. The morphology of the ciliate Hap- tophyru nrichiganensis iyoodhead and its relation to the other members of the rlstomatea. Trans. A m . Vicroscop. Sor . 52, 2 2 3 ->3 2 .

3 . __ 1934. The morphology of Haptophrya michi- gunemis \Voodhead, an astomatous ciliate from the intestinal tract of Hentidactylinin scirtatzoii (Schlegel) . Unit! . Calif. Prtbls. 2001. 39, 251-76.

4. Biitschli. 0. 1887-1889. Protozoa. Abt. 111. Infusoria und System der Radiolaria, in Bronn, H. G., editor, Klassen wid Ordnirng des Thiers-Reichs, C. F. Winter, Leipzig, 1, 1098-2035.

5 . Cepi.de, C. 1910. Recherches sur les infusoires astomes. .inatomie. biologie, ethologie parasitaire, sptematique. .4rch. :ool. exptl. et g i n . (ser. 5 ) 3, 331-609.

6. - 1923. S o t e taxonomique sur 1es iniusoires astomes ( Haptophryidae Cepkde noni. nov. pro Discophryidae CCp6de 1910). B i d . soc. 2001. Fr. 48. 105-8.

7 . Certes, .4. 1879. h’ote sur 1’Haptophrya gigantea Maupas iniusoire parasite des batraciens anoures d’.4lgerie. Bull. soc. zool. Fr. 4, 240-4.

8. Cheissin. E. M. 1930. Morphologische und systematische Studien iiber -4stomata aus dem Baikalsee. Arch. Protistenk. 70, 531-618.

9. Claparede, E. 8: Lachmann, J. 1859. Etudes sur les iniusoires et les rhizopodes. M e m . inst. nut. Genivois 6, 261482.

10. Cohn. L. 1904. Zwei parasitische Infusorien aus Disco- glossus pictus. Arch. Protistenk. 4, 43-63.

11. Collin, B. 1911. Etude monographique sur les acinetiens. I. Recherches experimentales sur 1’Ctendue des variations et les iacteurs teratoghes. Arch. zool. exptl. et gin. 48, 421-97.

12. Corliss, J . 0. 1953. Silver impregnation of ciliated protozoa by the Chatton-Lwoff technic. Stain Tcchnol. 28, 97-100.

1.3. ~ 1960. The problem of homonyms among generic names of ciliated protozoa, with proposal of several new names. J . Protorool. 7 , 269-78.

14. ___ 1961. T h e Ciliated Protozoa: Characterization, Classification, and Guide t o the Li teratwe. Pesrgamon Press, London and S e w York.

15. _I 1962. Taxonomic procedures in classification of protozoa. S y n p . Soc. gen. Microbiol. 12. 37-67.

16. - 1962. Taxonomic-nomenclatural practices in protozoology and the new International Code of Zoological So- menclature. J . Protozool. 9, 307-24.

1 7 . __ 1963. Establishment of an international type- slide collection for the ciliate Protozoa. J . Protozool. 10, 247-9.

18. GeorgCvitch, J . 1941. [Infusoria Astomata of Ochrida Lake Triclada.1 Ghlas Srpsk. kralj. Akad. 94, 149-61. [ In Serbian1

19. - 1950. Nouvelles recherche5 sur les infusoires astomes des oligochktes et des triclades d u lac d’Ochrid. Bi~11. Bcad. serbc Sci. nzath. nut. (N.S.) 1, 63-73,

Eimcria aurati, N. SP.. FROM GOLDFISH 2 73

20. Hazard, F. 0. 1937. Two new host records for the protozoan Haptophrya michiganensis Woodhead. J . Parasitol. 23, 315-6.

2 1. International Code of Zoological Nomenclature adopted by the X V International Congress of Zoology, London, July 1958. 1961. London.

22. Kay, M. W. 1942. A new astomatous ciliate from the newt, Eurycra bislineata (Green). A m . Midl. Nat . 27, 422-7.

23. Kent, W. S. 1880-1882. A Manual of the Infusoria. Vols. 1-111. David Bogue, London.

24. Kijensky, G. 1926. Contributions to the amitotic divi- sion of the infusorian Discophrya planariarum Siebold. Sbornik Zool . Praci., year 1925, 75 ( I ) , 1-5. [ In Czech with English summary1

25, Lachmann, J. 1859. Ueber einige neu entdeckte Infu- sorien u. uber contractile Blasen bei den Infusorien. Verh. Naturhist. Ver. pueuss. Rheinlande 16, 66-8, 91-3.

26. Lom. J. 1959. Beitrage zur Kenntnis der parasitischen Ciliaten aus Evertebraten IV. Neue Ciliaten aus der Familie Haptophryidae Cepede 1923, nebst einigen Bemerkungen zum heutigen Stand dieser Gruppe. Arch. Protistenk. 104, 133-54.

27. MacLennan, R . F. 1944. The pulsatory cycle of the contractile canal in the ciliate Haptophrya. Trans. A m . Mi- croscop. Soc. 63, 187-98.

28. Maupas, E. 1879. Sur 1’Haptophrya gigantea opaline nouvelle de l’intestine des batriciens anoures d’Algirie. Compt. rend. 88, 921-3.

29. Meyer, S. L. 1938. Haptophrya virginiensis nov. sp., a protozoan parasite of the pickerel frog, Rana palustris Le Conte. (.4bstr.) Anat . Rec. 72, 54-5.

30. ~ 1939, Description of Haptophrya virginiensis, a protozoan parasite of the pickerel frog Rana palustris Le Conte. J . Parasitol. 25, 141-4.

31. Puytorac, P. de 1957. L’infraciliature de quelques ciliCs Haptophryidae. Comparaison avec celle de certains thigmo- triches. Compt. rend. 244, 1962-5.

32. - 1959. Structures et ultrastructures nuclkolaires et perinucliolaires du macronoyau intermitotique. Compt . rend.

33. - 1963. Contribution i l’itude des ciliis astomcs Haptophryidae C i k d e , 1903 (cytologie, ultrastructure, taxi- nomie). Ann. Sci. nat., Zool. ( s k . 1 2 ) 5 , 173-210.

34. Rankin. J. S. 1937. An ecological study of parasites of some North Carolina salamanders. Ecol. Monogr. 7, 110-269. 35. Schultze, M. S. 1851. Beitrage zur Saturgeschichte der

Turbellarien. Greifswald. 36. Siebold, C. T. von 1839. Beitrige zur Naturgeschichte

der Wirbellosen Thiere. N . Schrift. Nat7tr.f. GPS. Danzig. 3. 56-71.

37. __ 1848. Lehrbuch der Vergleichenden Anatomie der Wirbellosen Thiere, Heft 1 in Siebold, C. T . von & Stannius, H., edi’tors, Lehrbuch der Vergleichenden Anatornip, Berlin. 1, 1-679.

38. Sikora, J. 1963. Study on the parasitic ciliate S1rinella uncinata (Schultze). Acta Protozool. 1, 13-20.

39. ___ 1963. Morphology of parasitc ciliate: Sleinrlla uncinata (Schultze 1851) and Sieboldellina planariaruni (Sie- bold 1845), in Ludvik, J., Lorn, J . & Vavra, J.. editors, Progrrss in Protozoology, Proc. 1st In&. Cong. Protosool., Prague, . tug. 1961, pp. 101-2.

40. Stein, F . 1860. Uber die Eintheilung der holotrichen In- fusionsthiere und stellte einige neue Gattungen und Arten auf dieser Ordnung auf. Sitz.-ber. K . bohm. Ges. Wiss., pp. 56-62,

41. __ 1867. Der Organismus der Infusionsthiere nach eigenen Forschungen in systematischer Reihenfolge bearbeitet. 11. Leipzig.

42. Woodhead, A. E. 1928. Haptophrya michiganrrtsis sp. nov. a protozoan parasite of the four-toed salamander. J . Parcz- sitol. 14, 177-82.

43. ivoodhead, A. E. & Kruidenier, F. J . 1936. The prob- able method of infection of the four-toed salamander with the protozoan, Haptophrya michiganensis. J . Parasitol. 22, 107-8.

249, 1709-11.

J. PROTOZOOL. 12(2), 273-275 (1965).

Eimeria aurati n. sp. ( Protoz0a:Eimeriidae) from Goldfish (Carassius auratus) in North America

GLENN L. HOFFMAN

Bitreau of Sport Fisheries and Wildlife, Eastern Fish Disease Laboratory, Kearneysville, Wes t Virginia

SYSOPSIS. Einzeria aurati n. sp. is described from the intestine of the goldfish, Carassizts auratus, from Pennsylvania, U.S.A. Ooc>sts were very numerous in long, whitish, opaque

N March, 1962 this laboratory received some gold- I fish for disease diagnosis from Blr. Richard Rice, Mercersburg, Pennsylvania. Seven of 29 fish eyamined were infected with Eimeria. These fish had long, whitish, opaque fecal casts hanging from the anus. The casts were composed almost entirely of oocysts. When shed, they sank to the bottom and looked somewhat like “spent” tapeworms (Figs. 1, 2 ) . The infected fish were very lethargic and their intestines were swollen with contained fluid. Although I believe that the Eimeria is pathogenic, a heavy infection of other parasites made this difficult to determine. These fish had apparently accumulated a heavy load of

fecal casts. They were 16 to 24 by 14 to 17 p (mean, 20.1 by 16.3) and were non-sporulated when passed but sporulated 2 to 5 days later.

parasites during the winter; Gyrodactylus, Trichodina, Chilodonella, Glossatella, Ichthyophthirius and Uroph- agus were present. Some also had bacterial infections of iieromonas liquefaciens.

Eimeria aurati n. sp.

Description. The developing oocysts of Eiineria were abundant in the intestinal mucosa, and round oocysts were present in the lumen of the anterior intestine; seven such non-sporulated oocysts were 12- 14 p (mean, 13.6 p ) in diameter (Fig. 5 ) . The sporont extended to the thin wall, which was about 0.25 p thick.

Documents

Resolution of Persistent Taxonomic and Nomenclatural Problems Involving Ciliate Protozoa Assignable to the Astome Family Haptophryidae Cepede, 1923