Freshwater Trie Lads (Turbellaria) of North America IV. the Polypharyngeal Species of Phagocata

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Freshwater Trie Lads (Turbellaria) of North America IV. the Polypharyngeal Species of Phagocata ROMAN KE Freshwater Trie lads (Turbellaria) of North America IV. The Polypharyngeal Species of Phagocata SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY • 1970 NUMBER 80 SERIAL PUBLICATIONS OF THE SMITHSONIAN INSTITUTION The emphasis upon publications as a means of diffusing knowledge was expressed by the first Secretary of the Smithsonian Institution. In his formal plan for the Insti- tution, Joseph Henry articulated a program that included the following statement: "It is proposed to publish a series of reports, giving an account of the new discoveries in science, and of the changes made from year to year in all branches of knowledge not strictly professional." This keynote of basic research has been adhered to over the years in the issuance of thousands of titles in serial publications under the Smithsonian imprint, commencing with Smithsonian Contributions to Knowledge in 1848 and continuing with the following active series: Smithsonian Annals of Flight Smithsonian Contributions to Anthropology Smithsonian Contributions to Astrophysics Smithsonian Contributions to Botany Smithsonian Contributions to the Earth Sciences Smithsonian Contributions to Paleobiology Smithsonian Contributions to ^oology Smithsonian Studies in History and Technology In these series, the Institution publishes original articles and monographs dealing with the research and collections of its several museums and offices and of professional colleagues at other institutions of learning. These papers report newly acquired facts, synoptic interpretations of data, or original theory in specialized fields. Each publica- tion is distributed by mailing lists to libraries, laboratories, institutes, and interested specialists throughout the world. Individual copies may be obtained from the Smith- sonian Institution Press as long as stocks are available. S. DILLON RIP LEY Secretary Smithsonian Institution SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY NUMBER 80 RomanKenk Freshwater Triclads (Turbellaria) of North America IV. The Polypharyngeal Species of Phagocata SMITHSONIAN INSTITUTION PRESS CITY OF WASHINGTON 1970 ABSTRACT Kenk, Roman. Freshwater Triclads (Turbellaria) of North America. IV. The Polypharyngeal Species of Phagocata. Smithsonian Contributions to Zoology, 80:1-18, 1970.—Two polypharyngeal species of Phagocata are recognized: P. gracilis (Haldeman), which includes P. subterranea Hyman, and P. woodworthi Hyman. The distinguishing characteristics of the two species are described and illustrated. P. gracilis has a backward loop of each sperm duct before it enters the penial bulb, an elongated, pointed penis papilla, a fibrous layer below the epithelium of the papilla to which longitudinal muscles attach, a non-glandular antechamber to the glandular seminal vesicle, and lacks a sphincter on the bursal canal. In P. woodworthi the sperm ducts enter the bulb without forming a back- ward loop, the penis papilla is short and truncate, its outer muscle layer consists of alternating sheets of circular and longitudinal muscle fibers, the penial lumen is not differentiated into seminal vesicle and ejaculatory duct, and the bursal canal is very wide and is provided with a terminal sphincter. The morphology of the reproductive system, geographic distribution, ecology, and life cycle of the two species are discussed in detail. Official publication date is handstamped in a limited number of initial copies and is recorded in the Institution's annual report, Smithsonian Year. UNITED STATES GOVERNMENT PRINTING OFFICE WASHINGTON : 1970 For sale by the Superintendent of Documents, U.S. Government Printing Office Washington, D.C. 20402 - Price 30 cents Roman Kenk Freshwater Triclads (Turbellaria) of North America IV. The Polypharyngeal Species of Phagocata Introduction been seen in several planarian species as an extraor- dinary occurrence, probably of traumatic origin. Polypharyngy, or the development of multiple It is also different from the transverse duplication pharynges, has been observed in three genera of or triplication of structures (pharynx, eyes) reported freshwater triclads: Phagocata, Crenobia, and by Ste"phan (1963) and by Tar and Torok (1967) Sphalloplana. The arrangement of the 3 to about for some clones of Dugesia tigrina. 40 pharynges within the common pharyngeal cavity From the taxonomical standpoint, polypharyngy generally is such that one pharynx occupies the is of very limited significance. The polypharyngeal normal position in the midline of the body, at- forms of the European genus Crenobia, i.e., C. tached to the anterior wall of the pharyngeal pouch montenigrina (Mrazek) and C. teratophila (Stein- and connected with the anterior ramus of the in- mann), agree so closely in other characters with the testine, while the remaining pharynges are inserted on the lateral walls of the pouch, more or less more widely distributed C. alpina (Dana) that they serially alternating on the right and left and con- are often considered subspecies of C. alpina. The nected to the corresponding posterior intestinal tripharyngeal C. anophthalma (Mrazek) differs rami. They may be separated from each other by from C. alpina chiefly by the lack of body and eye partial septa formed by tissue projecting from the pigment besides the presence of three pharynges. walls of the pouch. During the process of food in- In the American genus Sphalloplana, comprising gestion several pharynges are extended through the only subterranean planarians, polypharyngy is single, very distensible mouth opening. The origin known to occur in 5. mohri Hyman and related and phylogenetic significance of the polypharyngy forms discovered in caves of Texas, which in other have been amply discussed in the literature. The anatomical features entirely conform with the various hypotheses tendered on the basis of com- monopharyngeal species of this genus. The same parative morphological and experimental consid- may be said of the polypharyngeal forms of Phago- erations have been reviewed by Steinmann (1917: cata which, by the anatomy of their reproductive 3293-3304) and need not be repeated here. True systems, fit the design exhibited by other species of polypharyngy is a genetically stable condition, not the genus which have only one pharynx. It is a to be confused with the occasional development of coincidence that the generic (originally subgeneric) individual supernumerary pharynges which has name Phagocata was first applied by Leidy (1847: 248) to the polypharyngeal Planaria gracilis Halde- Roman Kenk, Department of Invertebrate Zoology, National man and that its derivation (from phagein, Greek, Museum of Natural History, Smithsonian Institution, Wash- to eat, devour) apparently alludes to the mode of ington, D.C. 20560. food ingestion. This seems to be one of the reasons 1 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY why some investigators, particularly in Europe, genus, Phagocata, on account of its polypharyngeal prefer the generic name Fonticola Komarek (1926: characteristic. With regard to its occurrence he 9) which, however, is superseded by the older name stated: "The animal I have only found in abun- Phagocata. dance in the neighborhood of Prof. Haldeman's Within the genus Phagocata, polypharyngy oc- residence, near Columbia, Pa. In a spring in front curs in the forms currently designated as P. gracilis of his house, thousands of them may be seen glid- gracilis, P. gracilis woodworthi, and P. subterranea. ing along the bottom. ." In a later paper, Leidy Examination of the anatomical characteristics of (1885: 51) gives more details about the locality: these three forms revealed, however, that their in- My attention was first called to the singular creature by the terrelationship has been misunderstood. late Prof. S. S. Haldeman, while I was a guest in his hos- pitable home at Chickis, on the Susquehanna river, in 1847. He had previously described it, in 1840, with the name of Acknowledgments Planaria gracilis. In front of the house is a beautiful spring This study was supported by National Research of water, which has its source in the neighboring cliff of Potsdam sandstone, and runs in a clear stream over a sandy Foundation Grant GB-6016 to the George Wash- bottom into the Susquehanna. ington University. I wish to express my thanks to the many persons who have contributed to my ef- Thus, we may consider the spring at Chickis (pres- forts by furnishing materials or information or have ent spelling Chickies), Lancaster County, Pennsyl- served as guides and collaborators in the field: Dr. vania, to be the type locality of Phagocata gracilis. J. T. Baldwin, Jr. (College of William and Mary), EXTERNAL FEATURES (Figure 1A-D).—Descriptions Mr. Ian R. Ball, Dr. Thomas E. Bowman (Na- of the external habit of Phagocata gracilis have tional Museum of Natural History), Mr. Harold S. been given by Haldeman (1840: 3), Leidy (1847: Feinberg (American Museum of Natural History), 248), Peaslee (1910: 5-6), and Hyman (1937a: 302). Dr. John R. Holsinger (Old Dominion College), Sexually mature specimens are generally 8 to 20 Mr. Leslie Hubricht, Mr. Lyndon Jon Mayers, mm long and 1.5 to 5 mm wide. Peaslee has ob- Dr. Marvin C. Meyer (University of Maine), Mr. served individuals measuring up to 30 mm in length Russell M. Norton, Dr. R. A. Raff (Massachusetts and 6 mm in width. As has been pointed out by Institute of Technology), Mr. Dale Webb, Mr. Law- Hyman, P. gracilis is a polytypic species, popula- rence Weingartner, and Dr. Eliot C. Williams tions from various localities differing particularly (Wabash College). in the shape of
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