ub A REVISION OF THE ADULT TAPEWORMS OF HARES AND RABBITS. 1 By OH. WARDELL/STILES, Ph. D., Honorary Custodian of the Helminthologieal Collection, U. S. National Museum. A DISCOVERY by Cooper Curtice in 1887 regarding certain young stages of rabbit tapeworms, observations which I was able to confirm and extend in 1894. immediately brought the leporine cestodes into a very important position , viewed from an economic as well as from a scientific standpoint, for Curtice's observation at first sight seemed to offer a guide to solving the puzzle as to the larval stage of the tape- worms of cattle, sheep, horses, and certain other animals. Curtice observed in the intestine of rabbits certain young cestodes which bore minute hooks upon the rostellum, and older stages were foTTiid which had lost these hooks. He determined the adult cestodes found in these rabbits as Tcenia pectinata andflooked upon the armed forms as the young of this species. .i With these observations before us, it lookedjas if some very radical changes would be made in the near future in Jhe classification of the tapeworms of the larger domesticated animals; it further seemed prob- able that the young stage of the tapeworms of cattle, sheep, and horses, would eventually be found to be an armed cysticercoid. In a paper 2 in 1894 I called attention to this bearing of the question and extended Curtice's observations. I found that the suckers 3 as well as the rostellum were armed, and that the head bore a remarkable resemblance to the head of Davainea. This complicated the question in so far as to render necessary the consideration of two further pos- sibilities i. e., were these young cestodes the early stages of avian 1 At the request of the author, it is here stated that he is not responsible for the insertion of commas between the names and the authorities, and for the absence of capitals at the beginning of specific names derived from personal names. These changes have been made in order to bring his paper into conformity with the usages at present followed in the U. S. National Museum. EDITOR. -Notes sur les Parasites 31 : line phase precoce des Te"nias du Lapin (Notice pre"- liminaire), Bull. Soc. zool. France, 1894, XIX, pp. 163-165. 3 Some old sketches of Curtice's show that ho also observed the hooks upon the suckers. PROCEEDINGS U. S. NATIONAL MUSEUM, VOL. XIX No. 1105. * 145 Proc. N. M. vol. xix - 10 ^*X2> fresh specimens. Plate VII, fig. 7, represents a segment measuring 2.08 mm. broad. The cirrus pouch with the enclosed inverted cirrus is the only portion of the male organs which is now visible. The female glands have also dis- appeared, but the receptaculum seminis is still present. The uterus fills the entire median field of the segment and appears as a reticulate structure, with dichotomous branches on the periphery. As the ova develop, the boundaries between the different branches of the network become more and more indistinct and the uterus assumes the form of a simple sac (Plate -VIII, fig. 1), in which no divisions, or at most extremely fragmentary and rudimentary divisions, are visible. The cirrus pouch and the receptaculum seminis are still present. Whether this uterus is primarily an actual network, or whether it is originally a simple tube with numerous proximal and distal branches which secondarily anastomose and then gradually disappear to form a common simple sac, can not be stated at present with certainty, but with the data now at hand, I incline decidedly to the latter view. The ova measure 52 to 60 /*; the pyriform body measures 12 ^ broad by 28 jj. long (horns included). In the younger segments no calcareous corpuscles are visible. When the ova in the uterus become more distinct and the partitions in the uterus become less distinct, a few calcareous corpuscles appear in the cortical portion of the segments. As the shells of the ova develop, the calcareous corpuscles become numerous. From Biehm's anatomical description and from my own study, I pro- pose the following as a revised specific diagnosis : Diagnosis. Andrya rliopalocepJiala (Eiehm, 1881), Stiles, 1895: Stro- bila attains 60 to 80 cm. in length by 5 mm. in breadth. Head unarmed, about 1 mm. in diameter, nearly quadrate in apex view. Neck about 1 mm. long. Segments 500 to 600 in number; active segments slightly broader than long; segments gradually increase in length so that posterior segments are as long as broad; they may attain 5 to 5.5 mm. in breadth. Genital pores single, near posterior corner of the segment, 158 TA PE WORMS OF HARES AND RA BBITS STILES. VOL. xix. for the greater part unilateral. Genital organs appear in about the one-hundredth segment. Male organs : Testicles dorsal, comparatively few in number, 75 to 80 JJL in diameter, and more numerous in the aporose than in the pore side of the segment 5 cirrus pouch 0.32 to 0.34 mm. long by 0.16 mm. broad; cirrus short, generally lies in two spirals within the pouch; prostata elongate. Female organs: Vagina slightly distal of cirrus pouch, swells to. a large receptaculum seminis median and dorsal of ventral canal; ovary, shell-gland, and vitellogene gland distal in pore half of median field: uterus ventral, appears as an apparent network with dichotomous peripheral branches, and finally forms a sac with indistinct partitions or without partitions. Ova 52 to 60 //; pyriform body 12 // broad by 28 /i long. Calcareous bodies appear in segments containing ova, and become numerous in segments in which the egg shells are distinct. Host. European Hare (Lepus timidus) by Biehm in Saxony. Cotypes.Nos. 1379, 1484, 1485, U.S.N.M.; collection of Leupkart; Vienna Museum. ANDRYA CUNICULI (R. Blanchard, 1891), Railli^t, 1893. Plates VIII, figs. 4-8; IX, fig. 1. 1881, Tceniarhopaliocephala [nee rhopalocephala] RIEHM, Zeitschr. f. d. ges. Naturwiss., 3 ser., VI, pp. 562-565, pis. v, fig. 2, vi, fig. 3. 1891, Anoplocephala cuniculi, R. BLANCHARD, Me'm. Soc. zool. France, IV, p. 447. 1893, Andrya cuniculi (R. BLANCHARD, 1891), RAILLIET, Traite d. Zool. me"d. et agric., I, p. 283. Eiehm described this form in 1881 under the following diagnosis: Kopf hakenlos, klein, aber gegen den sehr diinnen Halstheil stark keulenformig abgesetzt, wenn letztere nicht zu stark coutrahiert ist. Geschlechtsoffnimgen ein- fach, im dritten Viertheil des Proglottidenrandes gelegen ; Glieder trapezformig, etwa eben so lang wie breit. Lange im ausgestreckten Zustande bis 100 cm., Breite der reifsteu Glieder bis zu 8 mm. Wohnthier: Lepus cuniculus. Blanchard in 1891 changed the name to cuniculi on grounds of arti- cles 54 to 55 of the International Code, and placed the worm in the genus Anoplocephala. Eailliet in 1893 placed the form in the genus Andrya. I have not yet found this species in the United States, but have been fortunate enough, through the kindness of Geheimrath Leuckart and Dr. von Marenzeller, to obtain some of Biehm's original stock. One of the specimens, which was mounted whole, shows the following details: The anterior end is very narrow (0.4mm.) and segmentation is scarcely visible, so that only the head and a portion of the neck have been lost. Segmentation is noticed 0.64 mm. from the anterior extrem- ity, while 2 mm. from the end the segments are perfectly distinct, meas- uring 0.8 mm. broad by 0.24 mm. long. The anlagen of the female organs are indistinctly visible at about this point; they lie close to the pore side, of the segment, but owing to the poor condition of the mate- rial they can not be analyzed. Testicles could not be distinguished in NO. 1105. PROCEEDINGS OF THE NATIONAL MUSEUM. 159 these segments, but they appear about twenty segments later, soon becoming numerous and distinct and occupy the entire median field except the portion taken up by the female organs. Plate VIII, fig. 6, represents four segments about 15 mm. from the anterior end. About forty testicles are present. The anlage of the female glands is near the median line in the pore side of the median field; it is roundish and not very prominent. The genital canals are not very distinct, but may be traced to the lateral margin. Plate VIII, fig. 7, represents three seg- ments about 30 mm. from the anterior end, The testicles have increased in number and size ; the ovary and vitellarium are distinct, but the shell- gland is scarcely visible j the genital canals have become well differen- tiated, and the pore has developed. The greater majority of the genital pores are on one side of the strobila, and are situated in the distal half of the margin. In the specimen under discussion they range about as follows : 15. 16. 119. 44. 7. 5. 9. 4. 4. 6. 6. 2. 2. 18.' etc * About 50 mm. from the anterior extremity the structure of the seg- ment becomes complicated by the development of the uterus. My prep- aration does not permit a study of its gradual development, but this organ appears to be much more complicated than one would expect from Eiehni's description. At first no distinct limits can be made out, and one sees only numerous ova scattered through the segment. As the testicles atrophy, however, and the uterus comes more distinctly into view, the latter is apparently composed of a network of anastomosing tubes, much like the uteri found in Moniezia, arid totally different from the transverse uterus found in Cittotcenia, Bertia, and Anoplocephala. As in the case of the anastomosed testicles of Fasciola, it is here often impossible to distinguish whether we are dealing with anastomosing tubules or branched tubules which lie close together. My limited material will not warrant a more minute study of the uterus of this form. See also General Eemarks, p. 203. The cirrus pouch is seen to best advantage in segments in which the uterus has begun to develop. It is pyriform and measures 0.4 mm. long by 0.144 mm. broad. It is highly muscular, especially in its proximal portion, and its middle portion contains a prominent dilatation repre- senting a vesicula seminalis. Median to the pouch is found a roundish body, which evidently corresponds to the "prostata" described by Biehm. The vagina and large elongate receptaculum seminis lie distally of the pouch and vas deferens, as Eiehm has already described. The ova measure 48 to 60 // in diameter j the pyriform body measures 20 ^ broad by 32 to 44 p. long. The following is proposed as a revised specific diagnosis: Diagnosis. Andrya cuniculi (K. Blanchard, 1891), Kailliet, 1893: Strobila attains 100 cm. in length by 8 mm. in breadth. Head 160 TAPEWORMS OF HARES AND RABBITS STILES. VOL.XIX. unarmed, about 0.5 min. in diameter; rostellum not observed; neck filiform. Segments 500 to 800 in number, quadrate, broader than long; gravid segments may attain 8 mm. in breadth. Genital pores in about the middle of the margin or in distal half of the margin, irregularly alternate, but for the greater part unilateral. Male organs : Testicles scattered through the entire breadth of the median field ; cirrus pouch 0.4 to 0.48 mm. long by 0.14 to 0.16 mm. broad, quite muscular, and con- tains a distinct vesicula semiualis; prostata round. Female organs: Vagina distal of cirrus swelling to an elongate receptaculnm seminis ventral of vas deferens; ovary near median line in pore side of median field: uterus forms a network in median field, but as the ova develop the boundaries of the meshes become quite indistinct. Ova 48 to 60 IJL in diameter; pyriform body 20 // broad by 32 to 44 /.i long. Calcareous bodies become numerous in distal segments. JTostf. European wild rabbit (Lcpus cuniculus) in Saxony by Bielun ; ? European hare (Lepus timidus), see p. 155. Cotypes.~~Nos. 1377, 1378, U.S.N.M.; collection of Leuckart; Vienna Museum. Genus BERTIA, R. Blanchard, 1891. 1891, Bertia, R. BLANCHARD, Me'm. Soc. zool. France, IV, pp. 186-196. Type, Bertia studeri, R. Blanchard. Provisional diagnosis. Anoplocephaliuae, with segments broader than long. Genital pores regularly or irregularly alternate. Uterus (in all cases ?) a transverse tube with proximal and distal egg pouches ; geni- tal canals pass dorsally of dorsal and ventral canal and lateral nerve trunk, but in the two cases at least ventrally of dorsal longitudinal nerve; distinct prostatic gland wanting. Dorsal canal dorsal to dorso- lateral of ventral canal. Egg with well-developed pyriform body. Calcareous corpuscles present or absent. Hosts: Primates and rodents. Type.B. studeri, K. Blanchard, 1891. Two years prior to the publication of the genus Andrya by Eailliet, R. Blauchard, in 1891, proposed the genus Bertia for anoplocephaline cestodes with alternate genital pores, taking B. studeri from Antliro- popithecus troglodytes as type of the genus; as second species of the genus he described B. satyri. Unfortunately, on account of paucity of material, Blanchard was unable to give the anatomy of the type species, so that the generic diagnosis was based chiefly upon external characters. His original diagnosis reads as follows: Caput crassum, subsphaerieum, rostro aculeisque carens, acetabulis ellipticis, in dua paria valde distautia dispositis. Collum breve, propre tarn larguni quam caput. Corpus e permultis annulis brevissiinis latisque, imbricatis, constans. Pori geni- tales raarginales, tenuissimi, ab uno annulo ad alterum plus minus regulariter alter- nantes. In anuulo permaturo, ova in plures fasciculos regulares, transverse dis- positus, collecta. Oncosphaera pyriforme apparatu circumdata. Involutio ignota. From Blanchard's descriptions of the species the following may be taken as provisional specific diagnoses. NO. 1105. PROCEEDINGS OF THE NATIONAL MUSEUM. 161 BERTIA STUDERI, R. Bianchard, 1891. (Plate IX., figs. 2-3.) 1891, Bertia studeri, R. BLANCHARD, Me"m. Soc. zool. France, IV, pp. 187-190, figs. 1-4. 1894, Taenia studeri (R. BLANCHARD, 1891), BRAUN, Vermes, Bronn's Klassen und Ord- nungen, etc., IV, 36-37, p. 1143. Diagnosis. Bertia studeri, E. Bianchard, 1891 : Strobila attains 130 mm. in length, 15 mm. in breadth, 2.5 mm. in thickness; contains about 400 segments. Head subspherical, 0.65 mm. broad by 0.61 mm. long; suckers oblong, 0.34 to 0.345 mm. long by 0.27 to 0.28 mm. broad, two arranged on dorsal surface, two on ventral surface. Neck very short (0.3mm.); about as broad as head. Segments always much broader than long; maximum breadth 15 mm. at 45 mm. from head, maximum length about 0.35 mm. Genital pores very small, lateral, alternating very regularly. Male organs :? Female organs : Glands'? Uterus in fully developed stage composed of 30-35 polyhedral packages, 0.5 to 0.9 mm. by 0.1 to 0.8 mm., arranged in transverse row, occupying entire breadth and thickness of segments. Ova 53 to 60 /*, pyriform body 14 to 16 f.i broad, 23 to 30 // long, horns generally straight, oncosphere 10 to 12 //. Cortical layer of strobila supplied with numerous calcareous corpuscles 15 to 20 yu by 11 to 17 yu. Host. Chimpanzee (Anthropopithecus troglodytes (Linna3us) [Troglo- dytes niger])) by Studer. Type. Type and one paratype in Zoological Museum at Berne, Swit- zerland. Fragments in collection of 11. Blauchard. BERTIA SATYRI, R. Bianchard, 1891. 1891, Bertia satyri, R. BLANCHARD, Me~m. Soc. zool., France, IV, pp. 190-192. 1894, Tcenia, satyri (R. BLANCIIARD, 1891), BRAUN, Vermes, Bronn's Klassen und Ordnungen, etc., IV, 36-37, p. 1143. Diagnosis. Bertia satyri, E. Bianchard, 1891 : Strobila attains 245 mm. or more in length by 10 mm. in breadth by 2 mm. in thickness, and contains about 350 segments. Head and neck unknown. Segments always much broader than long, attaining a maximum length of 0.75 mm. Genital pores very small, lateral, irregularly alternate. Calcare- ous corpuscles numerous, attaining 30 by 20 yu. Dorsal canal lateral of ventral canal. Cirrus-pouch claviform, large and elongated. Uterus resembles somewhat that of B. studeri. Ovum 35 to 38 /* by 30 to 32 ^ pyriform body 12 to 17 JJL by 19 to 25 ^, oncosphere 13 //. Host. Oran-TJtan (Simia satyrus, Linnaeus). Type. Leyden Museum. From these descriptions it is impossible to come to any satisfactory conclusion as to whether the genera Andrya and Bertia should be kept separate or united. The form of the segments must surely be rejected as a generic character, and the fact that the pores of Bertia show a tendency to appear regularly alternate while those of Andrya show Proc. IN . M. vol. xix 11 162 TAPEWORMS OF HARES AND RABBITS STILES. VOL.XIX. a remarkable tendency to unilaterality can not, unassociated with other characters, be looked upon as establishing the genera as distinct. In the preliminary note to this revision, I described two American parasites as provisional members of the genus Andrya, reserving opin- ion as to the validity of the genus and calling attention to some important differences in the American and German leporine single- pored forms. Since the publication of my note, 1 I have received a copy of a recent paper by Meyuer describing two new species of cestodes, probably allied to the two forms of Bertia described by Blanchard, and through the kindness of Geheimrath Leuckart I have obtained one of the cotypes of Meyner's Tcenia (Bertia) mucronata. It will be necessary to consider these forms briefly in connection with the leporine parasites. Meyner evidently accepts Bertia only as a subgenus of Tcenia; as Bertia has, however, absolutely no generic relations with Tcenia, I now change his specific combinations from Tcenia (Bertia) mucronata and T. (B.) conferta to Bertia mucronata and B. conferta. Meyner discusses the anatomy of these two forms in detail, and from his account the following descriptions may be taken as specific diagnoses : BERTIA MUCRONATA (Meyner, 1895), Stiles, 1896. (Plate IX, figs. 4-5.) 1895, Tcenia (Bertia) mucronata, MEYNER, Zeitschr. f. Naturw., LXVIII, (5 ser., VI), pp. 1-86, pi. i, figs. 1-7. Diagnosis. Bertia mucronata (Meyner, 1895), Stiles, 1896: Strobila dagger-shaped, attains 140 mm. or more long by 8 to 10 mm. broad, serrate, imbricate, whitish yellow. Head, 0.34 to 0.714 mm. broad, apex nearly square, rostellum wanting; suckers oval 0.255 mm. broad, 0.2 mm. deep. Neck short, not sharply separated from head. Genital pores irregularly alternate. Male organs: Testicles appear in one hundred and twentieth segment, numerous, 75 to 100 p in 'diameter, crowded together in [antero Jdorsal portion of median field. Yas defer- ens dorso-anterior of vagina; cirrus-pouch not mentioned. Female organs: Glands in pore side of median field; vagina long; receptacu- lum seminis globular; uterus single, transverse, at first a simple tube appearing in about one hundred and thirtieth segment; eggs enter it in three hundred and fiftieth segment and blind pouches are formed. Ova 30 //, with three membranes; pyriform body 15 to 16 yu broad; oncosphere 13 to 14.4 //. Three longitudinal nerves each side of seg- ment, of which middle nerve is the largest. Dorsal canal dorsal of ventral canal. Genital canals pass dorsally of longitudinal canals and longitudinal ventral and main nerves, but ventrally of dorsal nerve. Calcareous bodies 4.9 to 18.4 //, more numerous in cortical layer; about 150 visible in transverse section of scolex, 10 to 12 visible 10 mm. from anterior extremity, 400 to 500 in transverse section of distal segments. 'Notes on Parasites 38 : Preliminary note to " A Revision of the Adult Leporine Cestodes," Vet. Mag., 1895, II, p. 341-346. NO. 1105. PROCEEDINGS OF THE NATIONAL MUSEUM. 163 Host. Black Howler (Alouatta caraya (Humboldt) [Mycetus niyer]} in Paraguay, collected by Neumeister. Types. In collection of Leuckart; one cotype No. 1483, U.S.N.M. In the cotype of this species at my disposal the uterus is not devel- oped. The worm presents an entirely different appearance from Andrya. The muscular cirrus pouch seems to be almost wanting, so far as I can distinguish, the end of the male canal appearing simply as a widened portion of the vas deferens with very weak muscles and much less prominent than the vagina. The prostatic gland, so characteristic for Andrya rlwpalocephala and A. cuniculi, is entirely wanting. Plate IX, figs. 4-5, give the general topographical anatomy of the segment. I do not believe that the division of the ovary into two wings as described by Meyner can be maintained, for in the cotype this division is extremely irregular. In some cases the ovary is not divided ; in other segments it is divided into two, three, or four wings. I further find the dorsal canals dorsal to dor so-lateral of the longitudinal canals. BERTIA CONFERTA (Meyner, 1895), Stiles, 1896. (Plate IX, fig. 6.) 1895, Tcsnia (Bertia) oonferta, MEYNER, Zeitschr. f. Naturw., LXVIII (5 ser., VI), pp. 86-103, pi. i, figs. 8-13. Diagnosis. Bertia conferta (Meyner, 1895), Stiles, 1896: Strobila attains 84 mm. or more in length by 6.5 mm. in breadth ; serrate. Head roundish, 0.68 mm.; rostelluin wanting; suckers oval. Neck about 2 mm. long, at first about as broad as the head, from which it is not sharply separated. Segments always much broader than long; sexual segments measure 5.1 mm. broad by 0.27 mm. long by 1.02 mm. thick. Genital pores irregularly alternate. Male organs : Testicles numerous in dorsal portion of median field (in figure, of transverse section con- fined to aporose two-thirds of median field at plane of genital pore), vas deferens dor so -anterior of vagina, its lateral portion quite wide; cirrus-pouch? Female organs: Glands in middle of pore side of seg- ment; receptaculum seminisoval; vagina with widened ciliated lateral portion. Uterus single, transverse, at first tubular, later with blind sacs. Ova begin to enter uterus 35 mm. from head in about the one hundred and thirtieth segment. Three lateral longitudinal nerves as in B. mucronata. Dorsal canal dorsal of ventral canal. Genital canals pass dorsally of longitudinal canals. Calcareous bodies numerous, may attain 19 /*; 20 to 25 visible on transverse section of sexual segments, 600 to 800 on transverse section of distal segments. Host. Bonnet Monkey (Macacus sinicus (Linna3us) [Macacus radiatus]). Type. In collection of Leuckart. While looking upon Meyner's paper as an interesting and impor- tant contribution to the knowledge of the worms of this group, I do 164 TAPE WOE MS OF HAKES AND EABB1TS STILES. VOL.XIX. not feel justified in utilizing liis work as basis for more than a provi- sional generic diagnosis for Bertia until the type species of the genus (B. studeri) can be studied more in detail. Nor do I consider the data at hand sufficient to justify a worker in suppressing Railliet's genus Andrya in favor of the earlier genus Bertia. From the very limited material and data at hand, I am inclined to believe that Anoplocepkala*, Bertia , and Andrya will all eventually be recognized as good genera, established upon well-recognized anatomical characters, but for the present, although Anoplocephala unquestionably stands, Bertia and Andrya can be accepted only as provisional genera, and as convenient means of classification. The final acceptance of the genera can follow only after examination of a larger series of specimens representing, if possible, more species 1 than are at present included under Andrya and Bertia. To utilize the generic terms Andrya and Bertia provisionally is cer- tainly better than to place the forms in the same genus with Tcenia solium. BERTIA PLASTICA (Sluiter, 1896), Stiles, 1896. 1896, Tcenia plastica, Sluiter, Centralbl. f. Bakteriol., Parasitenk. u. Infektionskr., I Abt., XIX, No. 24, pp. 941-946, figs. 1-6. June 30, 1896. During the proof reading of this revision an article has appeared by C. Ph. Sluiter, describing a new species of tapeworm (Tcenia plastica} from Galeopithecus volans. Sluiter is inclined to consider this new para . site as very closely related to Anoplocephala plicata, A. mamillana, and A. perfoliata, all of which he retains as members of the genus Tcenia. Unfortunately, several important points in the topographical anatomy have been omitted by the author, but his description and figures show that Tcenia plastica is much more closely related to Meyner's Tcenia (Bertia) mucronata than it is to the tapeworms of horses, on which account I transfer it to the genus Bertia. From Sluiter's description and figures, the following specific diagnosis is written : Diagnosis. Bertia plastica (Sluiter. 1896), Stiles, 1896: Strobilamore 1 Gottheil (1887) described two other cestodes, which may have some bearing upon the question, with the following diagnoses : ( Tcenia) No. 1. Length 20 cm., breadth at largest segment 3.5 mm. Head globular, four suckers, no rostellum, and no hooks. Neck extremely fine and filamentous 2.5 cm. long. Proglottides slowly increase in size, greatest breadth being only attained 15 cm. from head. Sexual orifices at the sides. Segments oblong, 3.5 mm. by 0.75 mm. From Macaque monkey (Macacus cynomolgm (Schreber) ). (Tcenia) No. 2. Length of largest specimen, 15 cm. Breadth at largest segment, 8 mm. Head large and clubbed, four suckers, no rostellum or hooks; neck short and thick; proglottides rapidly increase in size after the first inch and attain their full diameter from head, 7.5 cm. mature proglottides. They overlap each other at their posterior angles. Segments mature measure 8 by 4.5 mm. From Macaque monkey (Macacus cynomologus (Schreber) ) and Weeping Capuchin (Cebus capucinus (Linnseus) ). From the descriptions and figures it is very possible that these two forms are anoploceph aline cestodes, but Gottheil gives no characters which will aid in defi- nitely determining the question at hand. NO. 1105. PROCEEDINGS OF THE NATIONAL MUSEUM. 165 or less lanceolate, attaining 24 to 220 mm. in length by G to 11 mm. in breadth, and containing 80 to 400 segments. Head about 1 mm. broad by 1 mm. long; suckers round. Neck absent. Proximal segments increase rapidly in breadth; middle segments the broadest; distal segments decrease gradually to 5 min. in breadth, with more or less crenate and imbricate posterior edge, and measure 0.5 to 0.76 mm. long. Genital pores irregularly alternate in about the middle of the lateral margin; genital cloaca and organs developed in fortieth segment; eggs in uterus in fiftieth segment. Male organs : Cirrus large; cirrus-pouch muscular; testicles occupy chiefly the anterior portion of the segments extending the entire breadth of the median field. Female organs: Vagina distal to cirrus-pouch; receptaculum seminis elongate; vagina and receptaculum together extend about one-third across the segment; ovary very broad, extending nearly or quite to the aporose submedian line; shell-gland and vitollegene gland about in the porose submedian line; uterus tubular, transverse with proximal and distal pouches. Eggs 25 yw in diameter. Topography of nerves and canals'? Host : Flying Lemur ( Galeopitliecus volans), collected by Hubrecht in In India. Type: 1 *. Deposited in Amsterdam. Technique of type 1 ? BERTIA AMERICANA (Stiles, 1895), Stiles, 1896. (Plate X, figs. 1-10.) 1855, ? Tcenia laticephala, LEIDY, Proc. Acad. Nat. Sci. Phila., VII (1854-1855), Decem- ber, 1855, p. 443. 1895, Andrya americana, STILES, Vet. Mag., Phila., II, June, p. 344. Aug. 28, 1895. Leidy has given a short description of a tapeworm (Tcenia laticephala] from the Canada Porcupine which agrees in some characters with the form I described (Andrya americana) from the yellow-haired Porcupine. I am unable to find Leidy's types, but it seems to me very questionable whether the two parasites are identical. Leidy's description, which in a measure recalls the genus Davainea, reads as follows : T^NIA LATICEPHALA, Leidy. Head large; acetabula opposite, very prominent, large, hemispherical; mouth slightly prominent, unarmed. Neck short. Anterior segments of the body short, oblong square; posteriorly square. Generative aper- tures marginal, alternate. Protruding penes, elongate conical. Length of one speci- men 9 inches, greatest breadth f of a line. Breadth of head i a line ; of neck a line. Hob. The small intestine of Hystrix dorsata. B. americana was described in the preliminary note to the pres- ent revision as an Andrya, but a comparison of Meyner's excellent anatomical description of B. mucronata, and of his cotype, with the form under discussion shows that the American species is more closely related to B. mucronata than to A. rhopalocephala; on this account A. americana is transferred in the present paper to the genus Bertia. Several specimens of tapeworms were sent to the collection of the 166 TAPE WOK MS OF HARES AND RABBITS STILES. VOL.XIX. Bureau of Animal Industry by Professor Elrod, of Bloomington, Illi- nois, with the following label : " From Mesenteries of Canada Porcupine. Snake River, 1 near Nat. Park, Aug., 1894, M. J. Elrod." The following specific description will easily allow the recognition of the form : Diagnosis. Bertia americana (Stiles, 1895), Stiles, 1896: Strobila attains 33 mm. in length by 6 mm. in breadth and contains about 90 segments, the oldest of which are 0.8 mm. long. In some specimens the posterior segments become much narrower, longer, and thicker than the middle segments. Head, unarmed, measures 0.6 mm. broad by 0.38 mm. long by 0.32 mm. thick, and is nearly rectangular in apex view. The neck is absent, and the head is frequently retracted into the body, as in Drepanidotcenia laceolata. Suckers round, 0.176 mm. in diameter, open anteriorly. Genital anlagen visible in the earliest segments. Genital pores alternate in posterior half of margin. Male organs: Tes- ticles form a continuous band in the distal portion of the median field, extending on both sides to the longitudinal canals; about seventy testi- cles to a segment; vas deferens runs in the proximal portion of the segment; cirrus-pouch lies dorsal of the vagina, is very muscular, 0.48 mm. long by 0.144 mm. broad and extends to the ventral canal ; it con- tains a vesicula semiualis (0.19 mm. long) in its proximal portion, and the rather short, retracted, spinous cirrus in its distal portion. Female organs: The anlagen of the glands are seen immediately back of the head in or near the median line ventral of the testicles ; at first the glands are but little differentiated, but on their pore side a globular receptaculum seminis rapidly develops and becomes filled with sperma- tozoa; the glands develop rapidly, coming to lie right and left of the median line, the ovary becoming quite broad. The development of the uterus could not be followed in detail, but eventually it occupies the entire median field and becomes filled with ova 40 n in diameter; bulb of pyriform body 16 to 18//. Excretory and nervous systems: 2 Dorsal canals lie lateral of ventral canals and possess a thin lining. Genital canals cross the longitudinal canals and nerves dorsally. Calcareous corpuscles absent. Hosts. Yellow-haired Porcupine (Erethizon epixanihus], by Elrod; Canada Porcupine (E. dorsatus), by A. K. Fisher. 3 1 The Canada Porcupine (Erethizon dorsatm) does not extend so far west ; the host must have been the yellow-haired Porcupine (E. epixanthus). 2 The excretory system of this form will repay a careful study; in several anterior segments (transverse sections) I found the dorsal canals connected with the trans- verse canals, see pi. x, fig. 8. Since finishing this paper I have found some specimens in the Bureau of Animal Industry collection (No. 1502) which agree with Elrod's specimens, and bear the label, "E. dorsatm." These specimens were collected by Dr. A. K. Fisher at Lake George, New York. Fisher states that nearly every porcupine he has examined har- bors this worm. Cobbold, in 1862, examined some parasites from the same host-species and deter- mined them as "Tcenia pectin ata," with pores "all on one side." NO. 1105. PROCEEDINGS OF THE NATIONAL MUSEUM. 167 Types. Bureau of Animal Industry Cestode Series No. 1349 desig- nated as type, and deposited with sections of paratypes (Nos. 1350-1352) in United States National Museum. Paratypes distributed as follows: America: Collection of Bureau of Animal Industry; collection of Stiles (U.S.N.M.); collection of Hassall; collection of M. J. Elrod (Blooming ton, Illinois). Europe: Berlin Museum; Vienna Museum; collection of Bl an chard ; British (South Kensington) Museum (London). Geographical distribution. North America (Wyoming, by Elrod, and Lake George, New York, by Fisher). BERTIA AMERICANA LEPORIS (Stiles, 1895), Stiles, 1896. (Plate X, figs. 11-15..) 1895, Andrya americana leporis, STILES, Vet. Mag., II, June, p. 344. Aug. 28, 1895. Of this form I have but five specimens, all in exceedingly poor con- dition and unfit for any histological observations or detailed anatomical study. Enough can be seen on the preparations, however, to give a diagnosis which distinctly separates the parasites from all the other forms found in rabbits. The specimens were collected by Cooper Curtice; when they came into my possession there was nothing upon the label to give any clue to their origin, other than that Curtice collected them from Lepus. The specimens (Bureau of Animal Industry, Cestode Series Nos. 1170-1172, 1175-1176), all mounted, measure 23 to 47 mm. long, the widest segments attaining 5 to 6.5 mm. in breadth. Due allowance must here be made for the contraction of the specimens and the fact that they were subjected to pressure in mounting. The head is present on all specimens, and varies in measurement, as shown in the following table : Measurements of heads of Bertia americana leporis. No. (IT. S. N. M.). Breadth. Length. 1170 mm. 0.640 mm. 0.504 1171 .656 .448 1172 .656 (?) 1175 .688 (?) 1176 .640 .672 The form of the head can not be taken as a character of value, as it varies according to contraction. Plate X, figs. 12-13, show the heads of Nos. 1 170 and 1176, U. S. N. M. Eostellum could not be distinguished. The four suckers are powerful, and open diagonally forward; they are unarmed in the specimens at hand; their diameter varies between 0.22 and 0.24 mm., the muscular wall measuring about 64 ^ thick. There is no neck present, the segments being perfectly distinct immediately back of the head. The number of segments in the diiferent specimens varies from about 168 TAPEWORMS OF HARES AND RABBITS STILES. VOL.XIX. 60 to 95, but in no case was the strobila complete, as all of them had evidently shed proglottids. In some cases the head was retracted into the anterior segments, as is described for Drepanidotcenia lanceolata. In only one specimen (Plate X, fig. 13) was it extended. The first segment could not be satisfactorily measured; the second segment (No. 1176, U.S.N.M.) measured 0.448 mm. broad (slightly narrower than the head) by 64 /* long, and showed no trace of genital organs. The cirrus was distinctly visible in one specimen in the sixth segment, but no positive statement can be made for the earlier proglottids. In some of the other specimens the cirrus did not appear until several segments later in the eleventh in one case. The poor condition of the material must, however, betaken into consideration in this connection. Several segments after the appearance of the cirrus, a roundish body about 16 JJL in diameter appears in each proglottid, alternating a very short distance to the right and left of the median line. This body, which develops first from the eighth to the fourteenth segment, evi- dently represents the receptaculum seininis. In each succeeding seg- ment it is slightly farther from the median line than in the segment immediately preceding, so that we may conclude that the median line of the cestrode strobila is the seat of very active growth, a conclusion supported by observations on other species also, more particularly on the early genital anlageu of Thysanosoma giardi. 1 The testicles soon appear and extend in an irregular line across the median field of the segment, for the greater part distally of the female anlagen; their num- ber could not be ascertained exactly because of the poor condition of the material, but, as nearly as could be estimated from various portions of segments which were in better condition, there are about 50 testicles to each proglottid. The genital pores are alternate and situated in about the middle of the lateral margin ; the genital cloaca is generally quite deep. The cirrus and cirrus pouch are quite characteristic and allow an immediate determination of the form. The cirrus pouch assumes various shapes and proportions according to its contraction, but in general may be described as pyriform; it measures about 0.4 mm. long by 0.16 mm. broad; its proximal portion (0.192 to 0.224 mm.) is extremely muscular, being provided with an inner circular layer and an outer longitudinal layer, the two together, in some cases, measuring 16 // thick. The cirrus extrudes from the pouch in the majority of the segments ; when retracted, it appears spinous (wrinkles of the cuticle?) but extruded it is evidently smooth; the largest cirrus observed was 0.24 mm. long by 32 /n thick. The continuation of the cirrus canal in the proximal portion of the pouch is swollen into a vesicula seminalis and from the proximal extremity of the pouch the rather prominent vas deferens extends, somewhat sin- uously, through the anterior portion of the pore side of the segment to 1 A Revision of the adult cestodes of Cattle, Sheep, and allied animals; Bull. 4 Bureau of Animal Industry, p. 63, pi. xn, tig. 3. NO. 1105. PROCEEDINGS OF THE NATIONAL MUSEUM. 169 the vicinity of the female anl age farther than this it could not be traced. The female organs gradually undergo a change, but this process can not be described in detail because of lack of proper material. All that it is at present safe to say is that a dark body of cells arises in the vicinity of the receptaculum seminis and develops into the female glands very similar to those found in the Bertia of the porcupine; the uterus gradually extends over the segment suppressing the genital glands. Eggs globular with three membranes, but pyriform body not visible; outer membrane 40 to 42 u in diameter; middle membrane could not be studied; inner membrane immediately surrounding the oncosphere 21 JLI in diameter; hooks of oncosphere 9 ^ long. Numerous small calca- reous corpuscles present in the parenchyma. This worm has an entirely different appearance from the one found in the porcupine and it is possible that it will eventually be raised to specific rank. My material, however, does not warrant such a step at present. The following is proposed as a diagnosis of the variety: Diagnosis. Bertia americana leporis (Stiles, 1895), Stiles, 1896: Strobila 23 to 47 mm. long by 5 to 6.5 mm. broad, possessing from 60 to 95 segments. Head about 0.6 mm. broad by 0.4 to 0.6 mm. long, unarmed; suckers large, about 0.2 mm. in diameter, unarmed. Neck absent, stobilization beginning immediately back of the head. Genital pores alternate, situated in about the middle of the lateral margin. Genital organs develop very early, the male organs being visible some- times as early as the sixth segment. Cirrus large and ( ? ) smooth. Cirrus pouch about 0.4 mm. long by 0.16 mm. broad ; its proximal por- tion is very muscular and contains a vesicula seminalis. Vas deferens, large and prominent, extends from the region of the female glands to the cirrus pouch in the proximal portion of the segment ; testicles for the greater part in the distal portion of the segment, about 50 in number, arranged in an irregular row across the median field. Female glands first appear in the eighth to fourteenth segment alternately right and left of the median line; uterus spreads from female glands and occupies entire median field ; ova globular with three thin membranes; pyriform body apparently not present; outer membrane 40 to 42 }JL in diameter; middle membrane?; inner membrane 21 /* in diameter; hooks of onco- sphere 9 f.i long. Calcareous corpuscles present. Host. Lepus, sp. ?, by Curtice, development unknown. Types. Description based upon five specimens, of which Bureau of Animal Industry Cestode Series No. 1171 is designated as Type and deposited in the United States National Museum; paratypes Nos. 1170, 1172, 1175, 1176 U.S.N.M. Original material poorly preserved, alcohol ( ?) method, stained with ha3inatoxylin. Geographical distribution. United States of North America (State?) by Curtice. 1 70 TAPE won MS OF HARES AND RABBITS STILES. VOL.XIX. Genus CITTOTCENIA, Riehm, 1881. 1881, Cittotcenia, EIEHM, Zeitsclir. f. d. ges. Naturwiss., 3 ser., VI, p. 200. Type, Cittotcenia latissima Riehm, 1881 Tcenia denticulata Rudolphi, 1804. 1881, "Dipylidium, LEUCKART," 1863, p. p. of RIEHM, Zeitschr. f. d. ges. Naturwiss., 3 ser., VI., p. 200 ; p. 565. Type, Tce.nia canina Liunreus, 1758. 1891, Moniezia p. p., R. BLANCHARD, Mdm. Soc. zool. France, IV, p. 187; p. 444. Type, Tamia expansa Rudolphi, 1810. 1893, Ctcnotamia, RAILLIET, Traitd Zool. m6d. et agric., I, p. 278. Type, Tamia marmotce, Frolich, 1802. Diagnosis. Cittotcenia, Eiehm, 1881 : Anoplocephalinre with segments broader than long and longer than thick ; end segments in some cases showing a tendency to become longer and narrower. Two full sets of genital glands and two lateral genital pores to each segment; gener- ally one, in some cases two, simple transverse tubular uteri in each segment; uterus generally possesses simple proximal and distal diver- ticula. The vagina is ventral of the cirrus pouch on both sides of the segment. Dorsal canal varies somewhat in position, but shows a con- stant tendency to lie between the ventral canal and nerve, especially at the plane of the genital pores. Genital canals cross the longitudinal canals and nerves dorsally. Interproglottidal glands absent. Calca- reous bodies not yet recorded. Eggs with well-developed pyriform body, the horns of which are long, generally filamentous, crossing each other. Hosts. Eodents. Type. Gittotamia latissima, Eiehm, 1881, = Cittotcenia denticulata (Eudolphi, 1804), Stiles & Hassall, 1896. Eiehm proposed the genus Cittotcenia in 1881, with C. latissima as type and only species ; his description reads as follows : 4. Cittotcenia latissima. Die Gattung ist errichtet mit Riicksicht auf die merkwiir- dige Ausbildung des Excretionsorganes, welches nicht leiterartig, sondern mit je 3-5 vielfach omastomosirenden und in einander iibergehenden Hauptseitenstammen und zahlreichen, das ganze Thier netz-formig durchsetzenden Nebeniisten, ranken- artig die Tame dnrchzieht. C. latissima hat doppelte Geschleehtsorgane, welche auf den zizzenartig vorspringenden Hinterecken der Glieder nach aussen miinden. Der Kopf breiter als bei den vorigen erscheint vorn abgestutzt, hakenlos. Liinge der Strobila bis 2 Fuss, Breite der letzten Proglothiden bis iiber f Zoll. Wohnthier: Kaninchen. In the same paper Eiehm placed two other double- pored anoploce- phaline cestodes in Leuckart's genus Dipylidium, i. e., D. pectinatum and D. leuckarti, and later in the same year l he rejected his newly estab- lished genus Cittotcenia, uniting it with Dipylidium. In 1891 Blanchard 2 united in the genus Moniezia the double-pored anoploceph aline cestodes of rodents with those of ruminants. In 1893 1 Studien an Cestoden. Zeitschr. f. d. ges. Naturwiss., 3 ser., VI, 1881, pp. 565, 580. 2 Mem. Soc. zool. France, IV, 1891, p. 187. NO. 1105. PROCEEDINGS OF THE NATIONAL MI'S BUM. 171 Stiles ' excluded the known parasites of rodents from the genus Moniezia, while Railliet proposed for the double-pored leporine forms the genus Ctenotccnia with Tcenia marmotw as type. At first thought this genus does not appeal to helminthologists, who have not paid special attention to the Anoplocephalin*, as being well founded, but an anatomical study of the various forms does, I believe, fully justify the separation of the double pored cestodes of rodents from the double pored forms found in ruminants. The generic term Cittotcenia having priority must of course be accepted in preference to Ctenotamia. The characters of the three genera of double-pored anoplocephaline forms, according to my present interpretation, are as follows: Diagnosis. Moniezia, R. Blanchard, 1891: Anoplocephaline with segments generally broader than long and longer than thick ; end seg- ments showing a tendency to become longer and narrower. Two full sets of genital organs, with two (very complex) reticulate uteri and two lateral pores in each segment. On the right side the vagina is ventral, cirrus dorsal; on the left side vagina dorsal, cirrus ventral. Dorsal canal lies dorso-inedian of ventral canal. Genital canals cross the lon- gitudinal canals and nerves dorsally. Interproglottidal glands gener- ally present. Calcareous corpuscles absent from parenchyma. Eggs with well developed pyriform body, the horns of which generally (always ?) end in a disk. Hosts. Ruminants. Type. Moniezia expansa (Eudolphi, 1810), E. Blanchard, 1891. For diagnosis of Cittotcenia, see above, p. 170. Diagnosis. Thysanosoma, Diesing, 1835: Auoploeephalma3 with seg- ments generally broader than long and longer than thick; end seg- ments show a tendency to become longer and narrower. Two sets or one set of genital glands and two lateral genital pores, or one lateral genital pore to each segment; one transverse undulating uterus with ascon-spore or cornucopia-like egg-pouches. Genital canals cross the ventral canal and nerve dorsally, the dorsal canal ventrally. Interpro- glottidal glands absent. Calcareous bodies absent from the paren- chyma. Horns of pyriform body absent. Hosts. Rum inauts. Type. Thysanosoma actinioides, Diesing, 1835. From this analysis of characters it will be seen that Cittotcenia forms an excellent intermediate genus between Moniezia and Thysanosoma. The genus may be divided into two groups : Marmotce or Denticulata Group and Pectinata Group, the division being based upon the form of the cirrus pouch. In the first group this organ is pyriform, distinct and very muscular ; in the second group it is more elongated, resembling the nozzle to a hose, and is less distinct. 'Centralbl. f. Bakteriol. u. Parasitenk., XIII, 1893, p. 457; also, Bull. 4, Bureau Animal Industry, 1893, p. 54. 172 TAPEWORMS OF HARES AND RABBITS STILES. VOL.XIX. MARMOTS OR DENTICULATA GROUP. CITTOT^ENIA MARMOTS (Frolich, 1802) Stiles & Hassall, 1896. (Plate XI, figs. 1-8 ; Plate XII, figs. 1-2. ) 1802, Tcenia marmota, FROLICH, Der Naturforscher, XXIX, p. 77-79, pi. n, fig. 17-20. 1891, Moniezia marmota; (FROLICH, 1802), R. BLANCHARD, M6m. Soc. zool. France, IV, p. 187 ; pp. 444, 461-467, tigs. 33-35. 1893, Ctenotwnia marmota} (FROLICH, 1802), RAILLIET, Trait6 d. Zool. m6d. et agric., I, p. 278. 1896, Cittotcvnia marmota (FROLICH, 1802) STILES & HASSALL, Veterinary Magazine, III, p. 407. Gceze l in 1782 examined a worm which Blunienbach had collected from the Marmot; this parasite, which was probably Gittotamia mar- motce, Goeze considered identical with his Tcenia pectinata. Frolich 2 in 1802 found a cestode in the intestine of the Marmot (Arc- tomys marmota) and described it under the name Tcenia marmotce, as follows : [p. 77.] 43 : Taenia Marmoiae oblonga lanceolata antice attenuata utrimque serrata, articulis brevissimis, medio utrimque poro pilifero notatis, capite mutico. Tab. II, f. 17-20. MURMELTHIER-BANDWURM. Die Lange dieser "Wiirmer ist sehr veranderlich. Die grossten Individuen batten 6-7 Zoll in der Lange, ungefabr II'" in der Breite am Hintertheile, und beinabe 1'" in der Dicke. Ibre Gestalt ist iiberbaupt lanzettformisr, in die Liinge gezogen, am Vordertbeile allmalig, aber sebr betriicbtlicb, verscbmiilert, so dass das Vorderende am Gruude des Raises kaum etwas iiber V" breit ist. Aeltere Wiirmer, die durcb Geburten scbon mebrere Glieder abgesetzthaben, sind am Hinterende am breitesten, abgestutzt, oder balbmondformig ausgerandet; jiiugere Wiirmer, die nocb nicht gebobren haben, werden am Hinterende etwas schmacbtiger, uud das letzte Glied ist abgerundet. Der Kopf ist verhultnissmlissig sebr klein, rundlicb, riissellos, unbewaffnet, bell- weiss, mit vier deutlicben Saugblasen, die gepaart iiber einanderstebeu; [p. 78] an der Spitze ein undeutlicbes Knotchen, statt des Riissels. Der Hals ist zusammengedriickt rundlicb, sebr scbmiicbtig, etwas lunger als der Kopf, gliederlos, abwiirts breiter, und unvermerkt in den Vorderkorper iibergebend. Das Vorderende des Korpers scheint da, wo es an den Hals grenzet, fast gliederlos, wird von da abwtirts sanft uud unvermerkt breiter, und die Glieder, die vorwiirts nur durcb Querstreifen angezeigt waren, entwickeln sicb da deutlicher. Die Glieder des Korpers sind ganz sicbtbar ineiuander geschoben, wenigstens 20 mal breiter als lang, gleicbbreit, dicklicb, perlenweiss, zu beiden Seiten in eineu scharfen, bervorragenden Zabu auslanfend, in der Mitte der Oberfliiche eine einge- clriickte Langslinie. Jedes Glied bat in der Mitte des Seitenrandes ein duukleres, hervorragendes, wulstiges, in der Mitte durcbbobrtes Knotcben, aus welcbem ein baarfeiner, geradegestreckter, scbneeweisser, iiber V" langer Borsteii bervorragt. Diese Seitenborsten sind nicbt iinmer sichtbar, sondern sebr oft in den Rand der Glieder zuriickgezogen, wo sie denn an jeder Seite des Gliedes eiue bellere Querlinie bilden. Die bintern Glieder nebmen an Breite und Liinge etwas zu. 1 Versuch einer Naturg. der Eiiigeweidewiirmer tbieriscber Korper, Blankenburg, p. 363, footnote. 3 Beitrage zur Nat. der Eingeweidew., Der Naturforscbjer, XXIX, 1802, Halle, p. 77. NO. 1105. PROCEEDINGS OF THE NATIONAL MUSEUM. 173 Wolmort: In den diinnen Gedurmen des Murmeltbiers (Arctomys marmota Scbreb.). Anm. Icli traf diese scbone Wurmart in dem genannten Thiere, das in den all- gauischeu Alpen einbeimiscb ist, und von dem icb ein paar Stiicke am 28 Julius 1797 zu untersucben Gelegenbeit bekam, in ausserordentlicber Menge an. In den Gedilrm- en ernes iiltern Murmeltbiers fand icb ganze Nester von diesen Wurmern, so dass ibrc Zabl, im Ganzeu genommen, iiber 50 Stiicke betragen haben muss. Es lagen ibrer niancbmal 5-7 beysammen, und debnten den Darnikanal, indem sie in Knaulen iibereiuander lagen, ungemein aus. Icb sab Stiicke von diesen Wiirmern, deren bintere Glieder in der Mitte durch- locbert waren : andere Individuen batten sich am Hintertbeile durcb die Scblin<-e 7 O gescboben. Man siebt von selbst, dass die gegenwartige Art mit dem lanzetfdrmigen Baud- wurme 1 ungemein viele Aebnlicbkeit babe, da aucb dieser an den Randern der Glie- der kurze Seitenborsten zu naben pflegt ; demungeacbtet kanu icb mich bey genauer Vergleichung uicbt iiberzeugen, dass beide Wiirmer eiuerley Art seyn. Aucb die Tcenia pcctinata- ist mit dieser Art nabe verwandt. Eudolplii 3 in 1814 and 1819, Diesing 4 in 1850, and Baird 5 in 1853, mention Frolich's parasite under Tcenia pectinata, accepting Tcenia mar- motcu as synonym. E. Blanchard 6 in 1891 was evidently the second zoologist to find this species; he found 214 worms in four marmots at Briancon, in September, 1887. The specimens attained 112 mm. in length by 5 to 13 mm. in breadth. Head measured 0.55 to 0.63 mm. long by 0.80 to 0.84 mm. broad ; suckers globular, 120 ^ in diameter; neck absent; segments varied from 56 ^ to 0.75 mm. in length; genital pores double and opposite, in posterior half of lateral margin. Blanch- ard described a subcuticular excretory system, with two lateral longi- tudinal canals, connected by a transverse canal with the canals of the opposite side. Ova measured 48 to 60 yu; bulb of pyriform body, 23 /*. Stiles 7 in 1893 found that the dorsal canal lies between the ventral canal and nerve, and that the genital canals run dorsally of the longi- tudinal canals and nerves. He was unable to find Blanchard's acces- sory excretory system. Later he 8 figured the genital organs. Railliet 9 in 1893 took this species as type of his new genus Cteno- tcenia. From writings of Frolich and Blanchard and from my own studies I propose the following as revised specific diagnosis: Diagnosis. Cittotcenia marmotw (Frolich, 1802), Stiles & Hassall, 1896: Strobila attains 112 mm. (perhaps more) in length by 5 to 13 mm. in breadth. Head, 0.8 mm. broad by 0.5 to 0.6 mm. long ; hooks absent ; 1 Goeze, Eingeweidew., p. 377, pi. xxix, fig. 3-12. 3 Ebeudas, p. 363, pi. xxvn, fig. 7-12. 3 Erster Nacbtrag, etc., Ges. Naturf. Freunde Berlin, etc., VI, 1814, p. 108; Ento- zoorum Synopsis, 1819, p. 488. 4 Systema belmintbum, I, p. 498. 6 Cat. Specs. Entozoa or Intestinal Worms, Coll. Brit. Mus., p. 78. 6 Mem. Soc. zool. France, IV, pp. 444, 461-466, figs. 31-35. 7 Cent. f. Bakt. u. Paras., XIII, pp. '449-450, fig. 9. 8 Stiles & Hassall, 1893, Bull. 4, Bureau of Animal Industry, p. 71, pi. vn, figs. 6-7. 9 Traite de Zool. me'd. et agric., I, p. 278. 174 TAPEWORMS OF HARES AND RABBITS STILES. VOL.XIX. rostellum not visible. Keck absent, segmentation beginning immedi- ately back of the head, the proglottids rapidly becoming distinct; segments vary in length from 56 /t to 0.75 mm., and are always much broader than long, measuring in some cases 13 mm. broad; the poste- rior flap projects but a short distance over the anterior border of the next following segment. Genital pores double and opposite in poste rior half of lateral margin. Genital cloaca quite deep. Anlagen of genital canals and female glands visible in the first segments, testicles appear a little later. Male organs: Testicles appear about the thir- tieth segment, between 100 and 150 in number to a proglottid, scat- tered through the median field between the ovaries; cirrus-pouch, 0.5 mm. long by 0.17 mm. broad, very muscular, with vesicula seminalis in proximal portion and coiled smooth penis in distal portion. Female organs: Female glands nearly 1 mm. from the lateral border of the segment, some distance from the longitudinal canals; ovarian tubules appear about the thirty-fifth segment, reach their highest development from the forty-seventh to fifty-seventh segments and then rapidly atro- phy, disappearing almost entirely by the sixtieth segment; vagina ventral of the cirrus-pouch on both sides of the segment; uterus single, transverse, proximal to testicles, and possesses proximal and distal blind pouches, similar to those of Cittotcenia pectinata. Ova, 48 to 00 // ; bulb of pyriform body, 23 //. Excretory system : Dorsal canal between ventral canal and nerve. Longitudinal nerves rather close to lateral margin, near distal end of cirrus-pouch. Genital canals cross the longitudinal canals and nerve dorsally. Host. Marmot (Arctomys marmota) by Frolich and Blanchard. Types. Original type (?). Typical specimens with R. Blanchard (Paris), collection Bureau of Animal Industry (No. 1370, B.A.I.), and collection of Stiles (U.S.N.M.). Geographical distribution. (!) Frolich; France (Brianon), by Blan- chard. CITTOT^ENIA DENTICULATA (Rudolphi, 1804), Stiles & Hassall, 1896. (Plate XII, figs. 3-8; Plate XIII, figs. 1-3.) 1804, Tcenia denticulata, RUDOLPHI, Bemerkungen a. d. Gebiete d. Naturg., etc., p. 81. 1828, Alyselminthus denticulatus (RUDOLPHI, 1802), DE BLAINVILLE, Diet. d. Sci. nat., LVII, p. 607. 1853, Tcenia goezei, BAIRD, Cat. Entozoa Coll. British Museum, p. 78. 1881, Cittotcenia latissima, Reihm, Zeitschr, f. d. ges. Naturw., 3 ser., VI, p. 200. 1881, Dipylidium latissimum (Riehm, 1881), RIEHM, Zeitschr. f. d. ges. Naturwiss., 3 ser., VI, pp. 583-590, pi. v, figs. 5, 15, 17; vi, fig. 2. 1887, Tcenia latissima (RIEHM, 1881), NEUMANN, Trait6 des maladies parasitaires non- mi crobiennes, p. 426. 1891, Moniezia denticula \ta] (RUDOLPHI, 1804), R. BLANCHARD, Me"m. Soc. zool. France, IV, p. 187. 1891, Moniezia latissima (RIEHM, 1881), R. BLANCHARD, Me"m. Soc. zool. France, IV, pp. 187, 451. 1891, Moniezia goezei (BAIRD, 1853), R. BLANCHARD, M6m. Soc. zool. France, IV, pp. 444, 452-457, figs. 21-25. NO. 1105. PROCEEDINGS OF THE NATIONAL MUSEUM. 175 1893, Ctenotwnia goezei (BAIKD, 1853), RAILLIET, Trait<5 d. Zool. mc~d. et agric., T, p. 278. 1896 ; Ctenotcenia denticulata (RUDOLPHI, 1804), STILES & HASSALL, Vet. Mag., Ill, p. 6-9, and Centralbl. f. Bakt., Parasitenk. u. Irifektionskrankh., Erste Abt., XIX, 70-72. 1896, Cittotcenia denticulata (Rudolphi, 1804), Stiles & Hassall, Vet. Mag., Ill, p. 407. This species has had a most curious history, which should be a demon- stration to every helminthologist not yet convinced that in determining a cestode the microscope not the yardstick is his most important instrument, and that internal topographical anatomy not conditions of contraction of the external form furnishes the important characters for classification. At present we may be said to have two extreme parties in helmin- thological work; one party apparently considers minute histological details as all-important, and looks down upon the systematic work, especially that class of systematic work which studies into the biblio- graphic history of the species and demands a consistent application of the international rules of nomenclature; a second party appears to conceive the highest helminthology as consisting of determining and describing species chiefly upon external form, looking upon histological and anatomical details as "of interest to those who are interested in such matters," but of no importance to science. The history of C. denti- culata should, however, convince everyone that there is a middle ground upon which we should all unite, namely : First, careful determination and description of worms upon internal topographical anatomy; secondly, a thorough study of the history (both life history and bibliographic history) of each species; thirdly, histological details, and fourthly, a consistent application of such rules of nomenclature as will render all of our work international: International Rules. Rudolphi 1 in 1804 originally described Tcenia denticulata as follows : Unter den Eingeweidewiirmer [i. e., in the Hanover collection] war ein Stuck, das mieh sehr interessirte, namlich ein Bandwurm, der in Havemanns Gegenwart einem Kalbe abgegangen war; er war aber ohne Kopf.* *Ich habe diese Wiirmer im Museum zu Alfort wieder gefunden, und auch einige Exemplare geschenkt bekornmen. In seiiiem itzt ganz vergrifi'enen Werke iiber die Wiirmer hat Chabert ihn mit alien iibrigen Bandwurmem, unter dem Namen Tenia rubna6 [ruband] zusammengeworfen ; im Museum hingegeii war der Bandwurm aus Darm und Magen der Kuh ohne Noth in zwey Arten getheilt. Es ist eine wirklich neue, noch nirgends beschriebene Art, die ich Tcenia denticulata nenne, und an einem andern Ort ausfiihrlich beschreiben werde. Der Wurm ist 5 bis 16 Linien lang, die vordern Glieder sind 2 bis 5 Linien, die hintern beynahe einen Zoll breit; der kleine viereckige Kopf unbewaffnet, mit vier in Kreuz stehenden Saugmiindungen ; kein Hals, alle Glieder kurz, mit Oeffnungen an beyden Seiten, aus denen mehrentheils ein kleiuer spitzer Korper hervorsteht, wodurch der Wurm wie gezahnt erscheint. Ich selbst habe ihn nie beym Kindvieh gefunden. Later, 2 in 1805, he adds : 14. Tcenia denticulata mihi. T. ru bane's et lance"ole"s dans les intestins d'une vache. ^emerkungen aus dem Gebiete der Naturgeschichte, Medizin und Thierarzney- kuude auf einer Reise durch einen Theil von Deutschjand, Holland und Frankreich, I, Berlin, p. 81. 2 Bemerkungen, etc.. II, p. 39. 176 TAPEWORMS OF HARES A XL) RABBITS STILES. I ni Katalog steht : Deux especes de Tdnias rendues par la vaclie. Auf eiueiu anderu Glase steht: Tenias rubanes dans la caillette d'une vaclie. Ich muss aber alles fiir einerley balten, und liabe scbon vou dem Wurni eine kurze Besckreibuug gegeben wie icb von dem Kabinet der Thierarzneyschule in Hannover sprach. Still later Eudolphi 1 in 1810 continues the parasite under the same name and discusses it more in detail. His entire remarks are quoted here, as they are important, being based upon the original material. 2 [P. 79.] TAENIA J>ENTICULATA, K. Taenia: capite tetragono, collo nuUo, articulis bre- vissimis foraminibus marginalibus oppositis, lemuiscis dentiformibus. [P. 80.] Camper >o Beschiift. Berl. N. Fr., 4, p. 139. Gnielin, Syst. Nat., p. 3074, n. 55, Taenia ovina ft. bovis. Carlisle Transact. Soc. Linn. II, tab. 25, fig. 15, 16. Taenia ovina, bovis. Rudolphi bodoepor, Vol. I, p. 81 ; Vol. II, p. 39. Taenia denticulata. Hab. in Jlwe. Camperus a bove; Havemannus, Scliolae Vet. Hannov. Director meritissimiis- a vitulo (solitariani, capite destitutam); Cbabertus a vacca copiose, dejectam observaveruiit, hie etiani in vaccae ventriculo quarto reperit. Specimiua mea ex Mnseo Scbolae Veterinariae Altorfensis ditissimo. Descr. Vermes quindecim ad sedecim polliceslongi, antice duas ad quinque liueas, postice frve pollicem lati; coloris albidi, vel grisei. Caput ^xiguuin, tetragonum, latiusculum, osculis quatuor, anticis, subcontiguis, subglobosis, liorum apertuia exigua, orbiculari. Collum iiullum. Articuli aliquot capiti proximl angusti, mox vero latiores, tandem latissimi et subaequales fiunt, ita tamen, ut in uno alterove specimine media verinis pars paullo angustiores et simul longiore^ objiciat; omnes ce.teroquiu brevissimi, ut longitudo latissimorum vix liueam, Tplurimorum no dimidiam quidein excedat. Margines articuloruni postici crenati sive undulati superficieni insequentium qua partem tegnnt; latcrales (aute- rioribus nonnullis, rarius mediis quibusdam articulis exceptis) obtusiusculi [p. 81], foramine utrinque medio, opposite, insignes, e quo denticulus acutus, leviter reflex us et duriu^culus (lemniscus) exseritur. Substantia mollis, plus minus crassiuscula, ut crassities iuterdum lineam adaequet. Ova in substantia media cumulata, ovariis regularibus mibi lion visis. Obs. 1. Carlisle in figuris citatis canales articulorum laterales longitudinem versus decurrentes et ovaria in racemum brevisainium et latissimum digesta, babet ; baec omnia lie in tenuioribus quidein speciminibus vidi, vel sectione detegere potui. Denticuli validi liorunKiue foramina ab eodem minus bene sistuntur. Obs. 2. Chabertus in opere Bibl. n. 19.9. 3 indicate Taeniam hancce cum reliquorum mammalmm Taeniis, sub nomine Tnia ruban6, perperam conjunxit, in Musei Alfor- tensis autem catalogo duas in species (T<5nia rubane" et Tenia lanceole) praeter ullam necessitatem divisit, nani specimina niiuora quam reliqua ad formaui lanceolatana magis accedere nil refert. Obs. 3. A specie praecedente notis in obs. 2. ad eandem indicatis abunde differt quibus autem Taenia caprae n. 84. dicta distinguenda sit, hujus capite nou viso determinare non ausini; pars antica tanaeji quam in denticulata tenuior, licet quaiu in praecdente latior, videtur. That the citations from Camper, Carlisle, and Gmelin must be rejected from the bibliography of this species follows from the data given on page 177. ^ntozoorum sive vermium intestinalium. Historia naturalis, II, pt. 2, p. 39. 2 The original description of 1804 is evidently based upon the Alfort material and not upon tlie specimen seen at Hanover. Chabert's specimens must therefore be taken as types. 3 Traite" des maladies vermineuses dans les animaux. Paris, 1782; second edition, 1787 J German translation, Gottingen, 1789. NO. 1105. PROCEEDINGS OF THE NATIONAL MUSEUM. 177 Gurlt 1 in 1831 obtained Budolphi's specimens and gave two figures of the worm. Creplin 2 in 1842 also studied the originals and gave quite a, fair description of them (for details see Stiles 3 ). Since Creplin's time numerous authors have mentioned T. denticulata as a parasite of cattle and sheep, and quite a number of specimens have been determined as belonging to this species (for details see Stiles' 1 ). ^ In 1891 Blanchard 5 placed this species in his genus Moniezia. Later, in 1S!)3, 1 studied several end segments of Kudolphi's originals, compared them with several specimens determined as T. denticulata by various helminthologists, and called attention to the disagreement in the statements of different authors as well as TO the fact that none of their specimens agreed with J udolphi's original type. The few statements I made upon the originals were extremely meagre and guarded, showing that "we know practically nothing of the micro- scopic anatomy of M. denticulata." Upon examining originals of Baird's Tcenia f/oesei and Riehm's Dipylidium latissimum, Hassall and Stiles were surprised to find the great resemblance the cirrus bore to the cirrus in the few segments of Bndolphi's material deposited in the United States National Museum. Through the kindness of Geheimrath Mobius and Dr. A. Collin we obtained further material of liudolphi's originals and were able to prove 7 that Tcmia denticulata contained two distinct species of cestode, i. e., one agreeing Avith Biehm's I), lattssimum and T. goezcij the other with Cteuotcmia pectinata (Goeze, 1782 partiin, Biehm, 1881), Railliet, 1893. We then expressed the opinion that an error had occurred in the original label of Budolphi's specimens, and that they were in real- ity leporine rather than bovine cestodes. Baird's description 8 of T. goezei, in 1853, reads as follows: 19. Tcenia Goezii, Baird. Head wanting. Articulations of body very short, numer- ous. The inferior margin (straight. Genital orifices opposite, situated on or near the lower edge of each joint, the lemniscus being projected out in form of an ele- vated papilla, which curves downward. Greatest "breadth of body 6 lines, length of articulations about | a line. This species differs from expansa and denticulata in having the posterior border or edge of each joint smooth and rounded, instead of being crenulate or undulated, and having the genital orifices situated on the lower edge of the joint instead of in the middle. Hab. (!) From old collection. 1 Lehrbuch der pathologischen Anatomie der Haussiiugethiere, p. 381, pi. x, figs. 3-4. 2 Endozoologische Beitrage, Weigmanu's Arch. Naturg., I, pp. 315-327. 3 Revision of adult Cestodes of cattle, sheep, and allied animals, Bull. 4, Bur. An. Ind., U. S. Dept. Agric., 1893, p. 44. 4 Loc. cit., 1893, pp. 44-46. 6 Me"m. Soc. zool. France, IV, 1891, p. 187. 6 Loc. cit., 1893, pp. 42-47. 7 Vet. Mag., Ill, 1896, p. 6-9. 8 Cat. Specs. Ent. or Intestinal Worms, Coll. Brit. Mus., Lond., p. 78. Proc. K M. vol. xix 12 178 TAPEWORMS OF HA E ES A XD E A B BTTSS TILES. VOL. xix. Riehm 1 in 1881 studied the tapeworms of rabbits and hares, and in a preliminary account of his work described this same form as Citto- tcenia latissima, new genus, new species. Later 2 he placed this species in the genus Dipylidium with the following specific diagnosis : Kopf hakenlos liber f min. breit, rnit stark vorspringendeu Saugnapfen und da ent. Genital pores double, prominent, opposite in posterior half of margin, near corner of the segment. Male organs: Testicles 0.115 mm. in diameter, very numerous, scattered through the dorsal portion of the median field; cirrus-pouch large and prominent, 1.12 mm. long by 0.32 mm. broad. Female organs: Vagina runs ven- trally and distally of cirrus-pouch on both sides of segment; female glands very similar to those of Gittotcenia pcctinata, situated close to longitudinal canals ; uterus very complex. Eggs 52 to 63 //in diameter; bulb of pyriform body 16 to 24 /--. Excretory system : Very complex, numerous, thin-walled longitudinal canals (probably all belonging to the ventral system); dorsal canals with thick wall. Genital canals cross the longitudinal canals and nerves dorsally. Host. Common European wild rabbit (Lepus cuniculus). Types. Eudolphi's type in Berlin Museum; fragments in the collec- tion of Stiles, Nos. 17, 1492, U.S. Geomi/sf bnrsariua, Richardson; 1852, Geomys canadensis, LeConte; 1852, G.orcyonen- sis, LeConte. C. Hart Merriam, 1895. Monographic Revision of the Pocket Gophers, Family Geomyidje (exclusive of the species of Thomomus), p. 120. North American Fauna, No. 8, Division of Ornithology and Mammalogy, United States Department of Agri- culture. 184 TAPEWORMS OF HAKES AXD RABBITS STILES. 2. The second species, C. perplexa, is directly intermediate between the first and third, "but I am unable to find connecting links between it and the other forms. The cirrus pouch is about two thirds as large as that of C. pectinata and somewhat more distinct than that of C. rarl- abilis; the testicles are arranged in two triangles and extend laterally beyond the ovaries to the lateral canals. In general configuration of the body it resembles C. pectinata very closely, but is much shorter. 3. C. variabilis represents the last form of the series, presenting a cirrus pouch slightly smaller than that of G.perplcxa and testicles in a quadrangle which is confined entirely to the space between the ovaries. It occurs in three different forms as follows: (a) C. variabilis found in Lepus sylvaticus with characters described below, p. 192 7 measuring up to 10 mm. broad, the segments always much broader than long. (b) C. variabilis angusta differs from C. variabilis only in point of size; it measures only 2 mm. broad and the segments are about three times as broad as long. This form I have never found with ova, and it may be a case of arrested development. Those authors who take external form as a specific character would be obliged to give this parasite spe cific rank, but I am unable to admit this view. This variety is found in Lepus sylvaticus. (c) C. variabilis imbricata is the third variety, and is one due proba- bly entirely to host influence. It is found in Lepus palustris in Florida. The differences between this form and the type are but very slight ; the posterior margins are more scalloped, the segments more imbricate, and the cirrus pouch slightly more distinct. CITTOT^ENIA PECTINATA (Goeze, 1782 partim, Riehm, 1881), Stiles & Has- sall, 1896. (Plate XVI, figs. 1-2; Plate XVII, figs. 1-2.) ?1781, Tcenia acutissima, PALLAS, Neue nord. Beytrage, I, pp. 75-81, pi. in, fig. 25. 1eits, fast in der Mitte des Proglottidenrandes. Glieder kurz, trapezforrnig, anch im gestrecktesten Zustande miudestens 4 mal breiter als lang. Liiiige des ansgestreck- ten Wurmes niclit iiber 40 cm. rneist geringer. Breite der reifsteu Proglottiden bis 8 mm. Die Strobila ist oft durch Liingsfalten gestreift. Wohntliier: Lepus timid us. This diagnosis is hardly detailed enough to meet the requirements of the present day, but in the anatomical description Eiehm gives data which supply what is lacking in the passage just quoted. Many of the points he mentions can hardly be looked upon as specific characters, but should, I believe, be attributed to influences of technique, individ- ual variation, and possibly the specific influences of environment, namely, the host a subject to which the helminthology of the future must certainly give its most serious and careful consideration. The characters which appeal to me most in Eiehm's description may be briefly summarized as follows : Male organs: The testicles form a continuous band across the median field in the distal portion of the segment, and in Plate VI, fig. 4, they extend laterally close to the J Entozooruui sive Vermium intestinalium Historia naturalis, II, Pt. 2, Arnstelse- dami, pp. 82-84. 2 Icones Helminthum, pi. xiv, figs. 5-6. 3 Systema Helminthum, I, 1850, p. 498. 4 Loc. cit., p. 575. NO. 1105. PROCEEDINGS OF THE NA TIONA L MVSE UM. 187 longitudinal canals; the cirrus pouch is deserving of special notice. In the majority of the known Tseniidte the cirrus pouch is pyriform and rarely extends median of the ventral canal, but in C. pectin ata it is a long narrow structure reminding one of the noxzle of a hose; Riehm gives its average length as more than 1 mm., and on Plate VI, fig. 4, shows that it is more than twice as long as the distance between the genital pore and the longitudinal canal. This extreme length of the pouch is a character of great importance. Female organs: The vagina, according to Riehm, is about as large as the cirrus pouch. At a point corresponding to the proximal end of the cirrus pouch it becomes suddenly very thin, and leads to a second swelling, the receptaculum seminis. The female glands correspond in all essential characters to those of the genus Moniezia, except that the ovary is described as composed of two quite distinct halves. Their position, some distance from the longitudinal canals, is striking. The uterus is said to be similar to that of C. ctcnoides, namely "a common uterus for both sides, which extends the entire breadth of the segrnen, and is constricted only in the middle, so that in the gravid segments the lateral portion appears swollen by the ova in comparison with the rather thin median portion. Its volume is also considerably increased laterally [namely, laterally to the uterus, longitudinal in reference to the worm] by apparently uubranched tubes which extend anteriorly and posteriorly. As a matter of fact, however, these tubes resolve themselves as the optical sections of a corresponding number of circular wideniugs of the uterus." Excretory system: The dorsal canals become obliterated some distance from the head. The transverse canals are connected with one another, not only by the ventral canals, but also by numerous smaller longitudinal canals. Topographically, Riehm figures the genital canals as dorsal of the nerve and longi- tudinal (ventral) canal. The parasite is said to occur only in the fall and first half of the winter, and only in hares. It was especially common around the Roblinger See, but rare on the higher plateaux of Saxony. Blanchard 1 in 1891 states that he found this species in several hares of unknown origin. He has never found it in hares in the central part of France or around Paris, but found four specimens at Briancon in Lepus variabilis, killed at a height of 1,500 meters. His description, based upon these specimens, may be summarized as follows: The largest specimen was 18 cm. long; maximum breadth, 7 to 10 mm. ; head 315 to 340 /it broad ; neck, 285 to 325 jti broad ; in contracted specimens the neck may measure 1 mm. broad at the first segment; suckers elliptical, 142 // long by 135 JLI broad; opening, 80 // long by 53 // broad ; mature segments, 7 to 10mm. broad by 1.1 to 2mm. long; penis smooth, 40 to 45 f.i in diameter, extrudes 175 to 200 u from pore; eggs generally polygonal from reciprocal pressure, but become elliptical or subspherical when pressure is removed; 80 to 90 JJL by about 75 /.i ; outer membrane 1 to 2 u thick; diameter of bulb of pyriform body 25 to 30 JLI ; length 40 to 50 JLI ; horns terminate in a long filament; hooks of oncosphere 8 ju. Railliet 2 in 1893 places this species in his genus Ctenotwnia. Through the kindness of Dr. Brandes, I have obtained one of Riehrn's original specimens of Dipylidium pcctinatum for comparison with the American forms, and Blanchard has placed his forms from L. variabilis at my disposal. In my private collection I find several specimens ot tapeworms from Lepus tlmidus which I collected in Leipzig in 1890, and which agree perfectly with Rielmrs form. With this material at hand, together with one specimen from von Linstow and several from Moniez. 'Mem. Soc. zool. France, IV, pp. 445, 452, 457-460, figs. 26-30. 2 Traite Zool. uied. et agric., I, pp. 278-279. 188 TAPEWORMS OF HAKES AXD RABBITS STILES. I can add a few points of importance to Riehm's diagnosis and lay greater stress upon other points which 1 think should be brought out more emphatically. The anlagen of the female glands and genital canals appear in the earliest segments, while the testicles begin about 7 mm. back of the head. Male organs: The testicles are confined entirely to that portion of the segment which lies distally of the uterus, and they extend on both sides laterally of the ovaries to the longitudinal canals ; this latter character is one of considerable importance when we compare the Ameri- can and the European forms. Riehm has already directed attention to the loughoselike cirrus pouch and vagina, but I venture to call particu- lar notice to their size and form. Female organs: The excessive development which the female glands may attain is worthy of note: Riehin's statement that each ovary was divided into two halves I was unable to confirm. A peculiar feature of the female glands is that they develop suddenly and atrophy suddenly. In Riehm's cotype, for instance, the tubes of the ovary be- come visible 8 mm. from the anterior end (head lost) ; they then develop rapidly to a maximum and suddenly disappear, so that 22 mm. from the point where the ovarian tubes appear (or about 30 mm. from the anterior extremity) the ovary can no longer be seen; the testicles persist very much longer. The first trace of the uterus is seen about 12 mm. from the anterior end of specimen (about 14 mm. from anterior end of individual, if we allow 2 mm. for the lost head and first segments); the uterus passes on the ventral side of the ovary, and in all segments which I have exam- ined there is a single uterine anlage extending across the segment. The genital canals and uterus run dorsally of the nerve and ventral canal as Eiehm figures them, and on transverse sections the cirrus is shown to lie dorsally of the vagina on both sides of the segment. The nerve is very close to the lateral margin. Kiehm gives no measurements for the ova; Blanchard (1891, p. 460) states that the ova measure to 90 /; by 75 /v, and that the bulb of the pyriform body measures 25 to 30 IJL in diameter. The measurements in Riehm's specimen are somewhat smaller, the ova varying from 72 to 84 ^ the bulb of the pyriform body 14 to 16 y. My own specimens agree with Riehm's, but many of the ova do not exceed 56 ^ in diameter. The following is proposed as a revised specific diagnosis: Diagnosis. Cittotcenia pectinata (Goeze, 1782, partim, Riehm, 1881), Stiles & Hassall, 1896. Strobila may attain 400 mm. in length and 8 mm. in breadth, anterior portion usually lanceolate. Head small, about 0.25 mm. or less in diameter, and about 0.125 mm. thick; rostelliui and hooks not observed; suckers small. Neck very short, segmentation beginning almost immediately back of the head; the proglottids rapidly become distinct and are always much broader than long, the length being about one-seventh of the breadth; gravid posterior segments measure NO. 1105. PROCEEDINGS OF THE NATIONAL MUSEUM. 189 about 10 mm. broad by 1.5 mm. long. The anlagen of the genital organs appear very early; the female anlagen are found near the longitudinal canals immediately back of the head; testicles appear about (> mm. back of the head; genital pores double, in posterior half of margin. Male organs: The testicles are about O.G4 mm. iu diameter; they are numerous, about 150 in number, confined to distal half of segment, posterior to uterus, and extend across the entire median field passing the ovary on each side to the longitudinal canals; cirrus pouch unusu- ally large, attaining 1 mm. in length, extending some distance median of the longitudinal canals. Female organs: Ovary, shell gland, und vitellogene gland situated some distance median of the ventral canal, about 1 mm. or more from the lateral margin; a common transverse uterus to both ovaries; it passes the ovary ventrally, is generally larger in its lateral portions than in its median portion, and increases its volume by proximal and distal branches. Excretory system: Dorsal canal not observed; ventral canal median of nerve; transverse canals connected by secondary longitudinal canals. Longitudinal nerves close to lateral margin. Cirrus pouch, vagina, and uterus pass from median field into lateral field dorsally of nerve and longitudinal canals. Cirrus pouch dorsal of vagina on both sides of the segment. Ova globular 56 to 84 //. in diameter; bulb of pyriform body 14 to 1C //; horns long, curved. Hosts. Common European hare (Lepus timidus) by Goeze, liiehm, von Linstow, and Stiles; Mountain hare (L. variabilis) by E. Blanch- ard. Development unknown. Types. Original types 1 Cotypes of Eiehm in the collection of Leuckart; No. 1411, U.S.N.M. Typical specimens in collection of Blanchard; collection of Moniez; collection of Stiles (Xos. 116, 1234, 1238, U.S.N.M.). Geographical distribution. Europe: Germany (Saxony by Eiehm and Stiles; ? by Goeze); France (Biiancon by It. Blanchard; ? Lille by Moniez). CITTOT^NIA PERPLEXA (Stiles, 1895), Stiles & Hassall, 1896. (Plate XVIII, figs. 1-3.) 1895, Ctenotcsnia perplexa, STILES, Veterinary Magazine, II, June, p. 345. Aug. 28, 1896. 1896, Citlotmiia perplexa ( STILES, 1895), STILES & HASSALL, Veterinary Magazine, II I, p. 407. The specific name perplexa was proposed because it was so difficult to decide what to do with the form under discussion. Some half a dozen specimens were collected by Hassall from Lepus sylvaticus in Bowie, Maryland. They are all contracted and measure up to 57 mm. long by 10 mm. broad. The parasite resembles C. pectinata in general form in the early appearance of the genital anlagen and in the fact that the testicles extend beyond the ovaries to the lateral canals. It differs from C. pec- tinata radically in the size of its cirrus pouch and vagina, and in the 190 TAPEWORMS OF HAKES AXD RABBITS STILES. fact that the testicles are almost absent from the median portion of the field, being arranged in two triangles. C. perplexa agrees with C. variabilis in the general size of cirrus pouch. It differs from C. variabilis in the earlier appearance of the genital anlageii in the position and arrangement of the testicles. The following is proposed as specific diagnosis : Diagnosis. Cittotcenia perplexa (Stiles, 1895), Stiles & Hassall, 1896. Strobila attains 57 mm. long by 10 mm. broad. Head unarmed, small, about 0.32 mm. broad, not distinctly separated from body; rostellum iiol observed; suckers 0.112 mm. in diameter. Keck extremely short, segmentation beginning almost immediately back of the head. Genital pores double, and opposite, in about the middle of the margin. Anlagen of female glands and canals visible within 0.64 mm. of the anterior extremity of the head. Male organs: Cirrus pouch similar to that of C. variabilis, but smaller, about 0. -'88 to 0.32 mm. long, extending to or slightly beyond the lateral nerves; testicles arranged in two groups in each segment, one triangle being around each ovary and extending laterally to the longitudinal canals. Female organs : Agree essentially with those of C. variabilis, as does the general topography; uterus single or double. Host. Cottontail Babbit (Lcput> sylvaticus] by E assail in Bowie, Maryland. Types. Bureau of Animal industry, Cestode series, No. 1126, desig- nated as type, and deposited in the United States National Museum. Paratypes in Bureau of Animal Industry; Nos. 1110, 1131,1137-1139, U.S.N.M.; collection of Stiles; collection of Hassall. Other typical specimens will not be distributed until more material is obtained. CITTOT^ENIA VARIABILIS (Stiles, 1895), Stiles & Hassall, 1896. (Plate XIX, figs. 1-14; Plate XX, figs. 1-5.) 1892, "Tcenia pectinala, GOEZE," 1782, ex parte of CURTICE, Journ. Comp. Med. Vet. Arch., XIII, pp. 232-233. 1895, CtenotceniavariabiUs, STILES, Veterinary Magazine, II. June, p. 345. Aug. 28, 1896. 1896, Cittotcenia variabilis (&TILKS, 1895), STILES & HASSALL, Veterinary Magazine, III, p. 407. As stated in the introduction, helminthology is not so far advanced that it is possible for us to determine what limits should be given to genera, subgeuera, species, and subspecies, and for some time to come all classification into groups must be looked upon as experimental, the ideas of every author being subject to change from day to day as new facts in the comparative anatomy of cestodes are published or observed. The American form which I now describe as Cittotcenia variabilis is one of the parasites which can equally well be considered as a distinct species, or as a subspecies, possibly peculiar to given hosts. It is so perfectly distinct from the European C. pectinata that no specialist could fail to recognize the differences when he has the two forms side by side for comparison. Yet it is so closely allied to the European NO. 1105. PROCEEDINGS OF THE NATIONAL MUSEUM. 191 C. pectinata that the question forces itself upon us whether these dif- ferences can not be considered as subspecific, possibly due to some extent to a difference of conditions found in different hosts and in different countries. I frankly admit that during an examination of a large series of specimens 1 have changed my mind a dozen times in regard to tliis particular species. I am now of the decided opinion, however, that this form should be given specific rank. The specimens upon which this description is based were collected at Bowie, Maryland, by Hassall from the cottontail (Lepm Ni/lraticus). The strobila attains 100 to 180 mm. in length and 10 mm. in breadth. The head is very small, measuring 0.32 to 0.56 mm. broad; it may or may not be sharply denned from the neck; no rostellum or hooks visible; the suckers are small, measuring 0.12 to 0.28 mm. long by 0.112 to 0.24 rnm. broad; the opening of the sucker is directed diagonally forward. The neck is very short or absent, segmentation beginning very early and the proglottids rapidly becoming distinct. The anterior portion varies greatly in shape, according to contraction. The genital anlagen appear rather early; at about 5 to 10 mm. from the head two roundish bodies appear in each segment, one on each side of the median field near the longtndinal canals; these anlagen soon change in form to the pistol- shaped anlagen described for Moniezia; at about 25 mm. from the head the first testicles are visible. The genital pores are double and opposite, and situated in about the middle third of the margin. Segments 125 mm. from the head measure 0.56 mm. long by 6.5 mm. broad by 0.48 mm. thick. Male organs: The testicles number about 65 to 90; they are confined to the dorsal half of the segment distally of the transverse uterus, and do not extend laterally of the ovaries. The cirrus pouch lies dorsally of the vagina on both sides of the segment; it is long, narrow, and quite indistinct, coloring in carmine much more lightly than the vagina. Female organs: The vagina runs ventrally of the cirrus; it is long and narrow. From the margin of the segment for a distance of about 0.48 mm. it is surrounded by a deeply coloring- layer of cells, and then it is reduced to a thin narrow canal which later swells into the receptaculum seminis. The ovary, shell gland, and vitellogene gland resemble those of Moniezia and C. pectinala and lie 1.28 mm. from the lateral margin. The uterus may be double or single; in some segments a single uterine aulage extends across the entire seg- ment, running through the ventral portion of the ovary, and passing to the lateral fields dorsally of the longitudinal canals and nerves; in the majority of segments there are two distinct uterine anlagen, one to each set of female glands. The ventral canal runs about midway between the ovary and the lateral margin; the dorsal canal lies dorso- median of the ventral canal and is bound by a heavy cuticular lining. The longitudinal nerve lies laterally of the ventral canal, and ventrally of the genital ducts, about 0.64 mm. from the lateral margin. In the older segments the topography described above is preserved, 192 TAPEWORMS OF HARES AXD RABBITS- STILES. but gradually the uterus develops to such an extent that it suppresses all the genital glands. The cirrus pouch becomes indistinct, bat the vagina can be traced in nearly all segments. As the uterus increases in size it gives rise to proximal and distal branches, but the latter are far less regular than those of the European C. pectinata. In the posterior segments it is generally difficult to dis- tinguish the two uteri. The ova measure CO to 04 // in diameter: the bulb of the pyriform body, 12 to 16 /,/. This is the species (var. angusta] upon which my paper. "A double-pored cestode with occasional single pores 77 was based, and since writing that note another case of the same variation was noticed. On account of the numerous variations in the position of the genital pores, uterus, etc., noticed in the specimens studied, I proposed to name the worm Ctenotcenia rariabilis. Larval stage. The young specimens described on page 201 were col- lected in the same locality as the species here described, and from the same host; and it seems probable that the unarmed young mentioned on page 201 are the young of either C. rariabilis or C. perplexa. Diagnosis. Cittotcenia variabilis (Stiles, 1895), Stiles & Hassall, 1896. (American representative of C. pectinata.) Strobila attains 100 to 180 mm. in length and 10 mm. in breadth. Head small, about 0.3 to 0.6 mm. in breadth ; it may or may not be denned from the neck accord- ing to contraction. Neck very short, segmentation beginning almost immediately back of the head; segments always much broader than long. Genital anlagen appear very early, about 5 to 10 mm. from the head. Genital pores double, in about the middle third of the lateral margin. Male organs: Testicles about 60 to 100 in each segment, con- fined between the ovaries to. the dorsal portion of the distal half of the median field; cirrus pouch about 0.4 mm. long, narrow and very indis- tinct, lying dorsally of the vagina on both sides of the segment. Female organs: Ovary, shell gland, and vitellogene gland resemble the corre- sponding organs of Moniezia; they lie median of longitudinal canals, about 1.3 mm. from the lateral margin; the vagina is rather distinct, lies ventrally of the cirrus pouch, and for about 0.5 mm. from the pore it is surrounded by deeply staining cells; uterus may be double or single (in the same specimen) and may produce proximal and distal branches; ova 60 to 64 /* in diameter; bulb of pyriform body 12 to 16 //. Longitudinal nerve about 0.6 mrn. from the lateral margin; ventral canal large with thin lining; dorsal canal much smaller than ventral canal, dorso-median of ventral canal, with thick lining; transverse canals connect ventral canals [no injections made for secondary longi- tudinal canals]. Genital ducts and uterus pass from median to lateral field on the dorsal side of the longitudinal canals and nerves. Host. Cottontail Babbit (Lepus syh'aticus), by Hassall; Marsh Hare (L. palustris), by Mills. Types. Bureau of Animal Industry, Cestode series No. 117 desig- nated as type and deposited in the United States National Museum. Paratypes distributed as follows: Collection of Bureau of Animal NO. 1105. PROCEEDINGS OF THE NATIONAL MUSEUM. 193 Industry- collection of Leidy (University of Pennsylvania); collection of Harvard Museum of Comparative Zoology; collection of Ward; col- lection of Stiles ; collection of Hassall. Europe : Berlin Museum ; collec- tion of Leuckart; collection of Max Braun ; Halle Zoological Institute; Vienna Museum; collection of Stossich; collection of Parona; collec- tion of Monticelli; collection of Zschokke; collection of R. Blanchard; collection of Eailliet; collection of Neumann; collection of Moniez; British (South Kensington) Museum. Asia : Imperial University of Japan, Tokyo. Technique of types : Corrosive sublimate, acetic acid, acid carmine. Unarmed young stages deposited in collection of Bureau of Aiiimal Industry; col- lection of U.S.N.M. ; collection of Stiles; collection of Hassall; collection of Berlin Museum; collection of Leuckart; collection of R. Blanchard. Geographical distribution. Maryland (by Hassall), Florida (by Mills), Long Island (by Peters), ? Puget Sound (collection of Leidy). Varieties. To classify the forms at my disposal, I am compelled to recognize three varieties : (a) C. variabilis. B. A. I. Cestode series No. 117, type of the species is designated as type of this variety. The posterior liaps of the seg- ments are nearly straight; genital pore in about the middle of the lat- eral margin. Habitat: Lepus sylvaticus. (b) C. variabilis angusta. B. A. I. Cestode series No. 1119, designated as type and deposited in the United States National Museum. This variety is only about 2 mm. broad, the posterior flap is straight and does not overlap prominently, the genital pore is generally in the posterior half of the lateral margin. Pores occasionally single. (c) C. variabilis imbricata. No. 1246, U.S.N.M., borrowed by B. A. I.; B. A. I. Cestode series No. 1246, designated as type and returned to United States National Museum. At first sight it seems almost like splitting hairs to create a variety for these specimens the technique of which was different from that of C. variabilis. The worms were col- lected by Kobert Mills, an enthusiastic collector at Chuluota, Florida, and kindly presented to the United States National Museum. They were placed in 95 per cent alcohol, accordingly they are somewhat con- tracted. The posterior border of the segments is lobed, a character which is quite constant, and overlaps the next following segment about one-third of its length so that the segments have a general campulate appearance. Subfamily DliPYILIDIIlST^E, Railliet, 1896. 1850, Section Ehynchotcenia, 1 DIESING, Sy sterna Helminthum, I, p. 497. 1858, Subf. Malacolepidota (Soft-shell Tapeworms), WEINLAND, Human Cestoides, p. 52. 1863, Cystoidew, R. Leuckart, Die menschlichen Parasiten, I, p. 389. 1864, Subg. Rhynchotwnia, 1 DIESING, Sitzber. Akad. Wiss. Wien, XLIX, 1, p. 365. iKhynchotwnia, Diesing, 1850, a " section" of Toenia, may be interpreted as a sub- genus; by the law of priority, however, it falls as a synonym of Fimbriaria, Frolich, 1802, taking of course the same species as type, i. e., Tceuia malleus, Goeze, 1782. In order to meet objection to this ruling, should objection arise, I here definitely pro- pose Tcenia malleus, Goeze, as type of the subgenus. Proc. N. M. vol. xix 13 194 TAPEWORMS OF HARES AND RABBITS STILES. VOL.XIX. 1884, Subg. Microtcenia, 1 SEDGWICK, in GLAUS & SEDGWICK, Elementary Text-book of Zoology, I, p. 336. 1886, Cystoidei, K. LEUCKART, Die Parasiten des Menschen, 2d ed., I, p. 825. 1886, Group Cystoidotcenice, RAILLIET, filaments de Zool. m6d. et agric., p. 253. 1896, DipyUduncc, EAILLIET, Recueil de Me~d. vdt., 8 ser., Ill, 5, p. 159. STILES, Bull. 12, Bureau of Animal Industry, U. S. Dept. Agric., 1896, p. 28. Diagnosis. Taeniidse with rostellum which is generally armed ; geni- tal pores lateral (marginal), single or double; eggs with thin transparent shells, frequently arranged in egg sacs, in some cases scattered through the segments ; larval stage a cysticercoid ; adults in birds and mam- mals. Type. Dipylidium, E. Leuckart, 1863. Genus DAVAINEA, R. Blanchard & A. Railliet, 1891. 1891, Davainea, R. BLANCHARD & RAILLIET, Me~m. Soc. zool. France, IV, p. 428. Type, Tcenia proglottina, Davaine, 1860. ?1893, Chapmania, MONTICELLI, Naturalista Siciliano, XII (7-8-9), pp. 16-19. Type, Tcenia ar. richarcU on the other is, in iny opinion, another case of homoplasy. The development of double-pored forms in two distinct families (the Bothriocephalkhe Blanchard's genus Krabbea, and a number of cases in which a few double-pored segments have been found in Bothrio- cephalm latm and the Taeniidue forms a fourth case of the same nature. One of the greatest criticisms upon the present classification of para- sites seems to rne the fact that systematists have not taken into con- sideration the principle so well established in paleontology and other branches of zoology, namely, that the same structure may develop independently in several parallel series of animals. The sooner this prin- ciple is acknowledged the sooner we shall have a natural classification. Zone of lateral growth. A comparison of the older and younger seg- ments of numerous Ta3uiida3 shows that as the segments grow broader, the median field namely, the space between the two ventral canals increases much more rapidly in proportion than the lateral fields namely, the space between the ventral canals and the lateral mar- gin. This establishes the median field as the greatest zone of lateral growth. No particular narrow zone of the median field can be looked upon as the zone of lateral growth for all cestodes, as is shown by a comparison of a large number of forms. In Tccnia (type T. solium], for instance, the entire median field appears to participate in a more or less uniform lateral growth, for the genital glands retain their rela- tive position, increasing in size in proportion to the increase in size of the segment; the same appears to hold for Davainea. In the anoplocephaline forms we find some interesting variations in the zone of growth, as is shown by the relative position of the ovaries to the ventral canals or to the median line. In Thysanosoma yiardi the ovary bears an almost constant relation to the ventral canal, while the dis- tance between the ovary and the median line constantly increases as the segment grows broader; in this case, therefore, there is but little growth between the ovary and the canal, while the growth in the median side of the ovary is very marked. The same holds true to a lesser degree in Bertia americana. In Moniezia expansa and M. pianissimo, also the relation of the ovaries to the ventral canals remains almost constant, while the chief zone of growth is in the median field between the ovaries. 'The anatomy of the large American fluke (Fasciola magna] and a comparison with other species of the genus Fasciola s. st. vide p. 221, Jour. Comp. Med, Vet. Arch., 1895. 206 TAPEWORMS OF HARES AND RABBITS STILES. Iii Cittotcenla marmotce (Plate XI, fig. 5), C. prcecoquis, and C. pecti- nata (Plate XVI), on the other Land, there also is a marked lateral growth between the ovary and the ventral canal while the lateral field remains nearly the same width throughout the entire length of the strobila. In C. variabilis (Plates XIX-XX) there is a marked growth in the lateral fields. Abnormalities in the segments. Eiehm has already recorded segments of Ciitotcenia denticulata with three sets of female glands, and I have recorded segments of C. variabilis angusta with a single pore. New cases of both of these variations have been noticed in the preparation of this paper, namely, triple sets of female glands in C. denticulate and single pores in C. variabilis. I can look upon these cases, however, only as variations, similar to the occasional double pores in Bothrioceph- alus latus and Thysanosoma (jiardi, and can not ascribe to them any particular value from a systematic standpoint. The value of the genital pores in classification. Objection has arisen in some quarters to adopting the genital pore as the basis of classifica- tion. I both agree and disagree with this objection. It must not be forgotten that when Blanc-hard used the genital pores as basis for his classification, he naturally used the pore as representative of the geni- tal system; thus, if the pores are single, the entire female system is generally single; if the pores are double, the entire system of female glands is generally double. Viewed from this standpoint (which is the only logical interpretation I can give to Blanchard's classification, pro- posed in 1891), I must insist upon the great value of the pores in classi- fying cestodes. At the same time I fully agree and have stated so more than once before that the pores un associated with other charac- ters can not be relied upon as basis for a natural classification of cestodes. Internal topographical anatomy must, in my opinion, form the basis of the natural classification for both Oestoda and Trematoda. The size and form of the segments are characters which we should use with the greatest caution, always making due allowances for technique and contraction ; as generic characters I can under no circumstances admit their validity, and must therefore reject Sonsino's recently pro- posed genus Panceria (based essentially upon double pored segments which are longer than broad) unless other characters are brought for ward to place the genus on a firmer footing. In systematic work in helminthology we must not lose sight of another principle which is well acknowledged in other specialities, namely, that a given character which may be of great importance in classifying the species or genera of one group does not necessarily hold as a taxo- nomic character in all genera or higher groups of the same order or class. Thus the unilateral! ty of the pores in Hymenolepis and Anoplo- cepliala appears, so far as investigations have gone, to be a very impor- tant and constant character, while the same character must be used with the utmost precaution in the genus Davainea. I have shown NO. 1105. PROCEEDINGS OF THE NATIONAL MUSEUM. 207 above that the pores of I), salmoni, for instance, may be either unilat- eral or irregularly alternate; the same holds for J). tetragona, and apparently also for Andrya. The characters to be used in classify- ing the species of any given genus must be determined separately for every genus by an examination of a large series of specimens from dif- ferent localities and from different hosts. The unsatisfactory condition in which I have been obliged to leave the genera Bertia and Andrya is directly attributable to the limited amount of material at my disposal, and the fact that what few, and for the most poorly preserved, specimens I have examined represent only a few localities and a few hosts. The influence of a host upon its parasites. Considerable has been written upon the influence of various parasites upon their frosts, but very few authors have ventured to publish upon the influence of the hosts upon the parasites. This is probably due to two reas.ons: first, to the tendency on the part of many helmiuthologists to take the host species as a specific character of the parasite, and secondly the fact that the influence exerted by the parasite upon its host is a matter of economic importance, while the influence of the host upon the parasite would be looked upon by most workers as purely of scientific interest. To deny the economic importance of the host influence upon the parasite is, however, to my mind a short-sighted policy, for it must be this very host influence (i. e., environment) which has played an important role in the evolution of species, and which must have resulted in differentiating species and varieties, each of which after a time becomes in some cases at least dependent upon a single host. The genera Demodex, Psoroptes, and Cfiorioptes form excellent examples of the point under considera- tion; in these genera we find varieties which resemble each other so closely that it is often difficult and even impossible to determine the variety without knowing the host; experiments to breed some of the varieties of these genera upon other than their regular hosts, even though the experiment animal harbors a very closely allied variety of the same species, have been totally negative. This same variation is noticeable among worms. Specimens of Moniezla expansa taken from Ovis aries are for instance totally different from the specimens of the same species found in Ovis laticauda; the strobilse from this latter host recently studied by Setti, which I have been able to examine through the kindness of my Italian colleague Prof. 0. Parona, can not I believe be separated specifically from the European form, yet they certainly represent a distinct variety. Lonnberg has called attention to the variation in the hooks of avian cestodes taken from different hosts. I have also repeatedly noticed a variation in the spicules of the same species of nematode (Strongylus contortus, for instance) taken from cattle and sheep. To clearly define the varieties peculiar to certain hosts touches directly upon the question of transmission and infection, and on this account I submit that the study is of great economic as well as scientific importance. 208 TAPEWOEMS OF HAKES AND RABBITS STILES. The division of the genus Tcenia. In connection with the citations, p. 162, from Meyner's work, I take this occasion to reply to his remarks regarding the efforts which E. Blancliarcl, Eailliet, Hassall, and I have recently made to bring some order into the genus Tcenia. Upon page 8 of his thesis he says : Die Versucbe von Blauchard, Stiles und Railliet die Uebersichtlichkeit de'r Familio der Tamiaden durcli Zerlegung in eine Auzahl von Unterfainilien zu erleieb- tern, siud uiclit als besonders gliicklicbe zu bezeicbiien, namentlicb mit Riicksicbt darauf, dass die, von verscbiedeuen Gesicbtspunkten ausgehende, stets nur an einer bescbrankten Anzabl von Species vorgenonimene Eintbeilung notbgedrungeu aucb zu verscbiedenen Resultaten fiibreu musste. Bevor nicbt durcb eiugebende For- schungen wenigstens die typiscben Hanptforinen als feststebend angeseben wer- deu koimeri, diirften derartige Experimeute wobl uicbt auf allgemeine Anerkennung zu recbnen baben, bis dabiu aber verfeblen sie ganz entscbiedeu ibren eigentlichen Zweck, niimlicb Klarbeit nnd Eiufacbbeit in die Systematik zu briugen. In writing this critique, which is of course welcomed both by my French colleagues Blanchard and Eailliet, and by Hassall and myself as showing the view which Dr. Meyner takes of our work, the author has unfortunately overlooked several very important points. First of all, he has overlooked that very important principle so well expressed by Bacon in the words, "Truth emerges sooner from error than from con- fusion." iNone of us look upon our efforts to classify these parasites as perfect, free from criticism, and final beyond revision. But errors which we may have committed in these attempts to aid in reducing the chaos which exists in the genus Tcenia can be corrected very easily. We have the satisfaction of knowing that our work has been adopted by a number of specialists in helmiuthology and hardly feel called upon to apologize for the attempts we have made, even if not approved of by Veterinarian Meyner. A second error Dr. Meyner has fallen into is that he has forgotten that every classification must undergo an evolution (with epigenesis !). He would evidently have us include all four-suckered tapeworms in the heterogeneous and collective genus Tcenia, "until at least the typical chief-forms can be looked upon as established," although he failed to give an explanation of what he meant by "the typical chief-forms." If he refers to the type species of genera, he has lost sight of the fact that the type species of a genus is designated by any given author, and can not be changed after once being so designated. The type species is, therefore, optional, subject of course to certain principles, in the case of the author who designates it and obligatory in the case of other authors. Possibly Meyner refers to the most highly specialized species in the different groups. If so, however, his advice to wait until we study up such forms and decide which are the most highly specialized is wanting in reason. It is certainly far better to create a new genus for a form when we can not logically unite it with the known genera, than it is to place it with forms with which it does not agree in any important character. NO. 1105. PROCEEDINGS OF THE NATIONAL MUSEUM. 209 Of course our efforts at classification are experimental we all admit that; but from the very nature of things all efforts at classification in practically unknown groups are and must be experimental. The classi- fications must be changed time and again as new facts are discovered. Nor have our experiments (or, as Meyner puts it, "derartige Experi- inente") counted upon immediate general recognition (allgemeine Aner- kenuuug) ; it was not with that end in view that we published them. We do not expect to see our proposed classifications adopted by zoolo- gists at large until they have stood the test of other specialists in helminthology. We have not heard as yet, however, of any marked disapproval of the genera proposed from workers who were acquainted with the forms and who were competent to pass judgment on the case. When such authors propose a better classification, they can certainly count on IHauchard, Eailliet, Hassall, and myself as four helmintholo- gists who are ready to follow them. At present, however, I maintain that the classification originally proposed by Blanchard and since that time considerably expanded by Eailliet, Hassall, and myself is a far more natural and satisfactory classification oi the forms treated than any other classification ever proposed for the same forms. I am fully convinced, after a study of several thousand specimens, that the main features of the proposed division will stand, although the details of the system may undergo some changes. Helminthologists, as a class, are ultra-conservative in every line except species-making and yet as long as the Eudolphi-Diesing school exerts such a powerful influence in wield- ing the yardstick instead of the microscope, perhaps this generic con- servatism should be looked upon as a blessing. A third error of Meyner's is that he does not understand the views which he has attempted to criticise, or the relative rank of the groups proposed, and he ascribes to authors propositions which they never made. Thus he states (page 6) : Diese Anoplocephalinen theilt er (R. Blanchard) dann init Riicksicht auf die Anord- nung der Geschlecktsorgane in 3 Unterfamilien [!] ein und zwar (1) Genre Moniezla * * * ; (2) Genre Auoplocephala * * * ; (3) Genre Bertia Meyner thus makes the terms subfamily and genus synonymous rather a novel idea in systematic zoology; he accredits (page 8) Blauch- ard and Eailliet with a family "Anoplocephalen," although he states a few lines before that Eailliet accepted "Anoplocephaline" as a sub- family. Upon the same page he speaks of Bertia as a genus and Cteno- tcenia and Andrya as "Arten." It does not seem to me at all strange that our efforts should "fail totally in their object" with a worker who confounds such terms as species, genus, subfamily, and family. A fourth error into which Meyner has fallen in the passage quoted is the assumption that we have taken only a few species into consid- eration in making our classifications. True, we have not felt called upon to give a list of all the species of cestodes with which we have acquaintance, either through personal study or through the publications Proc. N. M. vol. xix 14 210 TAPEWORMS OF HARES AND RABBITS STILE '. of our colleagues, but it would be going a little far to assume that we did not have other forms in mind at the time we revised a por- tion of the species known to us. Consistency being such a virtue, however, if Meyner considers the species mentioned in the former papers by the authors he criticises as representing the sum total of the species known to them, he will probably not object to applying the same criterion to himself. As he has published original work on only two species, however, it might be difficult for him to justify himself in attempting upon the basis of these two species to overthrow the con- clusions which have been based upon a study of a much larger number of forms by four different specialists, all of whom agree in the general arrangement and differ with each other only in matters of detail. Meyner's remarks represent an excellent example of what is so com- mon in science to-day, namely, a tendency to jump at conclusions and to generalize in a too dogmatic manner when one is not thoroughly at home on the subject he has under discussion. The above remarks are, I think, sufficient to show that Meyner's amusing criticisms would better have been submitted to more careful thought before they were published. They can not have much weight with helminthologists, and should not have much weight with zoolo- gists in other specialities. A number of genera not discussed in this paper have been proposed by authors for various forms in the family Tasniidse. Some of these genera must be rejected. In regard to f some of the others, judgment must be reserved for the present. I hope, however, to publish before long a summary of all the genera proposed. Regarding the newly pro- posed genera for avian cestodes, see Stiles, 1 1896. Exchange of cotypes. There seems to be a popular impression among workers that it is a perfectly easy matter for a scientist to read a description of a species or genus and judge of the validity of the pro- posed form without examining specimens. This impression is certainly true in some cases, but must always be taken cum grano salis. I do not hesitate to assert that not one-half of the species and genera of parasitic worms ever described can be rightly judged from their descriptions, nor is this always attributable either to the description or to the ability of the worker attempting to pass judgment upon the case. It is rather attributable to the undeniable fact that a person obtains an entirely different impression from a study of the objects from what he obtains from reading a description, be it ever so detailed and complete. Many a species or genus has been accepted or rejected by an author who would have decided differently if he could have examined cotypes of the forms he was discussing. In view of these facts, which I am confident the specialists in hel- minthology will admit, I wish to appeal again to helminthologists to 1 Report upon the present knowledge of the tapeworms of poultry, Bull. 12, Bureau Animal Industry, Washington, D. C. NO.] 105. PROCEEDINGS OF THE RATIONAL MUSEUM. 211 deposit cotypes with the chief museums of the world. It has been our policy to do this as far as possible, and as soon as the international postal laws 1 are amended so as to allow the acceptance of these specimens in the international mails, the system will be extended. As a matter of justice, I believe the type specimen belongs in the national museum of the country in which the specimen was collected. The first duplicate (paratype) belongs by right to the Berlin Museum, as that museum now possesses more types of parasitic worms than prob- ably any other collection in the world. I would set forth the claims of the United States National Museum for the second duplicate on the grounds that the European workers all have easy access to the speci- mens at Berlin, while the frequent sending of a rare and valuable specimen from Europe to this continent is attended with more or less danger of loss. Japan certainly has a claim to the third duplicate. It is indeed time that helminthologists give the question of the pres- ervation of types their serious consideration. Many of the older col- lections can not be traced 5 Davaine's collection is evidently destroyed, and many more private collections will follow it if more attention is not given to this subject. CONCLUSIONS. The chief results of this paper may be briefly summarized as follows: 1. The time has come when helminthologists must donate their type specimen of every species to museums where they can be properly cared for and where they may be open to the inspection of other specialists. I maintain that the original type rightfully belongs to the national museum of the country in which the specimen was collected ; that the second specimen (first duplicate first paratype) rightfully belongs to the Berlin Museum ; that the United States National Museum has a valid claim for the third specimen (second duplicate), and that Japan has a valid claim for the fourth specimen. 2. In order to insure the preservation of types and cotypes, I invite the specialists in helminthology to unite upon some regular plan for the deposit and exchange of such specimens. 3. Permanent mount in balsam appears to me to be a much better method of preserving a type than to retain it as an alcohol specimen. 4. The known adult leporine cestodes belong to the five genera: Anoplocephala, Andrya, Bertia, Cittotcenia, and Davainea. 5. Of these genera Anoplocepliala, Cittotoenia, and Davainea are con- sidered as -perfectly valid genera; Andrya and Bertia are left subjiidice, although it is my impression that they will finally be recognized as valid, as soon as sufficient material can be properly studied. 'Owing to the absurd international postal regulation (16. 3. b.) excluding zoological specimens from the international mails, some of the paratypes mentioned as " distrib- uted" in this paper have not yet been sent to the authors named; they will be forwarded as soon as an opportunity presents itself. 212 TAPEWORMS OF HARES AND RABBITS STILES. VOL.XIX. 6. The microscope must supplant the yardstick and internal anatomy must take the place of external form in judging the validity of cestode and trematode genera and species. 7. The principle of homoplasy must be recognized by helmintholo. gists as well as by other zoologists, and any classification which leaves this important and well-recognized principle out of account can be taken only as a preliminary (although often necessary) study (p. 204). 8. The median field of the Tseniidie is the seat of the most active lat- eral growth, and the same rule will probably be found to apply to other families of Cestoda. No particular longitudinal zone of the median field can, however, be named as the zone of most active growth in all Ta3niida3 (p. 205). 9. The armed young cestode which I mentioned in Note 31 [ (1895) is not the young of an anoplocephaline tapeworm, as Curtice, Braun, Kailliet, Neumann, and I had inferred, but represents the young stage of the single-pored cestode referred to in my paper in 1895. 2 It evi- dently belongs to Davainea salmoni. 10. The double-pored cestode with occasional single pores, described in my paper in 1895, 3 is Cittotcenia variabilis angusta. 11. I have also found some very young stages of an unarmed cestode in the intestine of the cottontail rabbit, which probably belong to Cittotcenia variabilis. This young stage corresponds to what we may expect to find as the larval form of Moniezia expansa of cattle and sheep, and I doubt whether it will be possible to distinguish it from the young of that species. This renders the question of the origin of the tape, worms of cattle and sheep more complicated than it was formerly sup- posed to be, and demands the strictest experimental proof on the part of any author who suspects that he has solved the mystery of the life history of the cestodes of cattle and sheep. 12. The head of a cestode increases in size after the parasite reaches its final host, as is shown by a comparison of the younger speci- mens. 13. None of the adult leporine tapeworms thus far described in Europe have as yet been found in America. The American forms which have been published as " Tccnia pectinata" must be distributed over several species typical to this continent. 14. The following table includes all of the genera at present recog- nized in the subfamilies Ta3niin, Mesocestoidime, Anoplocephalinse, and Dipylidiinse of the family Ta3niidae. A number of other genera have been proposed, but some of them must fall as synonyms, while judg- ment upon others must be reserved. Several of the genera in this key 1 Notes sur les Parasites 31: Une phase prdcoce du Teuias ,nia\v\ the subfamily Dipylidiinse, see my Report upon the Present Knowledge of the Tapeworms of Poultry, Bull. 12, Bureau of Animal Industry, 1896. * 1863, Hesocestoides, VAILLANT, L'Institut, No. 1524. Type, Mesocestoides ambiguus, Vaillant, 1863. 1885, Pti/chophysa, HAMANN, Zeitschr. f. wiss. Zool., XLII, p. 740. Type, Tcenia lineata, Goeze, 1782. 3 1758, Twnia, LINNAEUS, Systema naturae, 10 ed., p. 646, 819. Type by elimination Tawia solium, Linmvus, 1758. 1800, Ali/selminthus, ZEDEH, Erster Nachtrag zur Natnrg. d. Eingeweidewiirmer, p. 221. Tamia, Linnaeus, renamed, hence type Tania solium, Linmeus, 1758. 1803, Halt/sis, ZEDER, Anleitung zur Naturg. d. Eingeweidewiirmer, p. 298. Alysel- minthus, Zeder, renamed, hence type Tcenia solium, Linmeus, 1758. 1850, Arhyncliotwnia, DIESING, Systema helminthum, I, p. 497. Subgenus containing type of genus, hence type Ta>nia solium, Linnrous, 1758. 1858, Acanthotrias, WEINLAND, Human Cestoides, p. 51. Type, Cysticercus acantho- trias, Weinland, lSoS = Ciislicercuscellnlosw, Rudolphi^:larva of Twnia solium. 1858, Tamiarhyncltns, WEINLAND, Human Cestoides, p. 46. Type, Tcenia mediocanellata, Kiichenmeister, 1852 = Tcenia saginata, Goeze, 1782. 1858, Echinococcifer, WEINLAND, Human Cestoides, p. 52. Type, Tania echinococcus, Siebold, 1853. 1863, Cystotwnia, LEUCKART, Die menschlichen Parasiten, I, p. 223. As subgenus including type of genus, hence type Tcenia solium, Linnaeus, 1758. To this list of synonyms must be added the subgenera Cysticercus, Cocnurus, and Echinococcus. They take the date and authority of the author who first used them in a subgeneric sense and not the date and authority of the author who first pro- posed them, since according to the International Code, section 48, b, the generic and specific names of larval cestodes (Cysticercus, Cocnurus, Echinococcus, Finna, Hydatigena, Hydatis. etc.) are not entitled to the law of priority. The genus Tainia is frequently divided into three subgenera : Cysticercus ( = Cys- totcenia\ Ccenurus, and Echinococcus. Tasnia must be substituted for the first sub- genus (type T. aolium) and Echinococcifer must probably be substituted for the third subgenus. 214 TAPEWORMS OF HAKES AND RABBITS STILES. Egg with thin transparent shells; frequently in egg capsules; in some cases scattered through the segments; head nearly always armed with hooklets on rostelluni; larval stage a cysticercoid ; adults in birds and mammals. Dipylidiince (p. 193), 27 Anoplocephalince. 1 5. Ova with well developed pyriform body; genital canals pass dorsally of longi- tudinal canals and main nerve trunk (not yet satisfactorily demonstrated for Andrya) 6 Ova without pyriform body; genital canals pass dorsally of ventral canal and nerve, but ventrally of dorsal canal (not yet demonstrated for Stilesia centripunctata) . 10 6. Uterus a transverse tube, generally continuous, in some of the double-pored forms divided (double), one tube for each ovary, and generally provided with proximal and distal pouches; dorsal canal lies dorsal to dorso-lateral of ventral canals ; pedunculated prostatic gland absent 7 Uterus more like a network in its early stages; afterwards the boundaries of the meshes nearly or quite disappear and the uterus (or uteri) appears like a sac. . 9 1 The genera of Anoplocephalince may also be determined by the following key, which although simpler than the one given above, is much more artificial : Anoplocephalina. A. Genital pores single B Genital pores double F B. Genital pores unilateral or irregularly alternate with decided tendency to unilaterality . C Genital pores regularly alternate or nearly so; pedunculate prostatic glands absent D C. Pores unilateral; testicles unilateral or nearly so in median field opposite pores; uterus transverse with proximal and distal pouches; pedunculate prostatic gland absent Anoplocephala, 1 1 Pores irregularly alternate with decided tendency to unilaterality; uterus forms a network in its younger stage, later the meshes become nearly or quite obliterated; testicles extend across the median field to or beyond the ovary; distinct pedunculated round or elongate prostatic gland present. .. Andrya, 26 D. Egg without pyriform body E. Egg with well-developed pyriform body; uterus transverse with thin- walled proximal and distal pouches; genital canals pass dorsally of dorsal and ventral canals and main longitudinal nerve trunk Bertia, 14 E. Uterus single and transverse, without pouches, or double, a cornucopia-like egg pouch being present near each ventral canal, the median transverse connecting canal not visible; genital canals pass dorsally of nerve and ventral canal, but ventrally of dor- sal canal (not shown for S. centripunctata) *. Stilesia. Uterus transverse but undulate, with thick-walled asconspore or cornucopia-like egg pouches; genital canals pass dorsally of nerve and ventral canal, but ventrally of dorsal canal Thysanosorna. F. Egg without pyriform body; uterus transverse and undulate, with thick-walled ascon- spore or cornucopia-like egg pouches; genital canals pass dorsally of ventral canals and nerves, but ventrally of dorsaj canal Thusanosoma. Egg with well- developed pyriform body ; genital canals dorsally of ventral an I dorsal canals and nerves G G. Uterus a transverse tube, single or double, with thin-walled egg pouches ; dorsal canal dorsal to dorso-lateral of ventral canal ; vagina ventral of cirrus pouch on both sides of the segment Cittotcenia, 19 Uterus reticulate, double; dorsal canal dorsal to dorso-median of ventral canal; vagina ventral of right cirrus pouch, dorsal of left cirrus pouch Moniezia. NO. 1105. PROCEEDINGS OF THE NATIONAL MUSEUM. 215 7. Genital pores single 8 Genital pores double; vagina ventral of cirrus pouch on both sides of the seg- ment Cittotcenia (p. 170), 19 8. Pores unilateral; testicles unilateral, or nearly so, in median field. Anoplocephala (p. 150), 11 Pores regularly or irregularly alternate; testicles extend ncrosg the median field (not shown as yet for B. studeri and B. satyri) Bertia (p. 160), 14 9. Genital pores irregularly alternate with marked tendency to unilaterality ; pedunculated round or elongate prostatic gland near ventral canal on pore side of segment; dorsal canal lies dorsal to dorso (? lateral) of ventral canal. Andrya (p. 154), 26 Genital pores double; pedunculated prostatic gland absent; vagina ventral of right cirrus and dorsal of left cirrus Moniezia. 1 10. Uterus single, transverse, but undulate, with asconspore or cornucopia-like egg pouches; genital pores single or double; testicles form a band in median field or are divided into two lateral groups, one in each lateral field Thysanosoma.* Uterus single and transverse, or double, being represented by a cornucopia-UTce pouch on each side of the segment; genital pores irregularly alternate; testicles in two lat- era I groups and absent from median line Stilesia.* Anoplocephala. 11. Head with posterior lobes; body may attain 8 to 25 mm. long by 3 to 15 mm. wide; head 2 to 3 mm. broad. Host: Horse (Equus caballus) A.perfoliata. Head without posterior lobes 12 12. Head 4 to 6 mm. broad; strobila 9 to 80 cm. long by 5 to 20 mm. broad. Host: Horse (Equus caballus) and Mule (Equus asinus X caballus) A.plicata. 4 Head less than 1 mm. broad 13 13. Strobila 10 mrn. long by 1.5 to 2.25 mm. broad; contains 10 to 28 segments; distal segments may become nearly as long as broad; genital glands confined to proximal third of worm ; testicles 15 to 30 111 each segment ; cirrus pouch 0.48 mm. long. Host: European rabbit (Lepus cuniculus) and Mountain hare (L. variabilis) A. wimerosa. Strobila attains 6 to 30 mm. long by 4 to 6mm. broad; 35 to 53 segments present, the distal 4 to 8 completely filled with ova; testicles 60 to 100 in each segment; cirrus pouch attains 0.8 mm. in length. Host: Horse (Equus caballus). A.mamillana (p. 153)' Strobila 10 to 16 cm. long by 6 to 8 mm. broad; 60 to 80 testicles present. Host: Arctomys, sp A. transversaria (p. 154). 1 1891, Moniezia, R. BLANCHARD, Me"m. Soc. zool. France, IV, p. 187. Type, Twnia expansa, Rudolphi, 1810. For the species of this genus see Stiles & Hassall, 1893, A Revision of the Adult Cestodes of Cattle, Sheep, and Allied Animals, Bull. 4, Bureau of Animal Industry, U. S. Dept. Agriculture. >2 1835, Thysanosoma, DIESING, Med. Jahrb. d. ost. Staat. n. Folge, VII, pp. 105-111, pi. in. Type, Thysanosoma actinioides, Diesing, 1835. For the species of this genus see Stiles in Stiles & Hassall, 1893, Bull. 4, Bureau of Animal Industry. 3 1893, Stilesia, RAILLIET, Traite de Zool. me'd.'et agric., I, pp. 277-278. Type, Tcenia globipunctata, Rivolta, 1874. For the species of this genus see Stiles in Stiles & Hassall, 1893, Bull. 4, Bureau of Animal Industry. 4 Doubtful whether this specific term should stand, as it is antedated by equina, 1781, and magna, 1789. 216 TAPE WORMS OF HAEES AND EA BEITS- STILES. Inertia. 14. Hosts: Rodents; 1 cirrus pouch distinctly muscular, prominent, with vesicula seminalis; receptaculum seminis round and prominent; pores very regularly alternate ; calcareous corpuscles present or absent 15 Hosts: Primates and lemurs; 1 cirrus pouch? ; calcareous corpuscles present 16 15. Genital pores in posterior half of lateral margin; cirrus pouch about 0.5 mm. long; posterior segments show a tendency to Itecome narrow and thick; strobila attains 33 mm. in length by 6 mm. in breadth, and contains about 90 segments, calcareous cor- puscles not observed. Host: Yellow-haired porcupine (Erethizoii epixanthus) and Canada porcupine (E. dorsatus), North America IS. americana (p. 165) Genital pores in about the middle of the lateral margin ; cirrus pouch about 0.4 mm. long ; posterior segments do not become narrow ; strobila attains 23 to 47 mm. in length by 5 to 6.5 mm. in breadth, and contains 60 to 95 segments; calcareous corpuscles present. Host: Rabbit (Lepus, sp. ?), North America. B. americanaleporis (p. 167). 16. Cirrus pouch not so prominent as vagina, and with but weak musculature; receptacu- lum seminis round to elongate and not very prominent; pores irregularly alternate; ovary confined to pore side of median field; uterus transverse with proximal and distal pouches 17 Cirrus pouch very muscular; receptaculum seminis elongate; pores irregularly alter- nate; ovary extends to aporose portion of median field; uterus transverse with prox- imal and distal pouches; strobila 24 to 220mm. long by 5 to 11 mm. broad, and con- tains SO to 400 segments. Host: Fining lemur (Galeopithecus volans), India. B. plattica (p. 164). Cirrus pouch? vesicula seminalis?; uterus composed of 30 to 35 polyhedral packages. 18 17. Strobila attains 140 mm. or more in length by 8 to 10 mm. in breadth. Host: Black howler ( Alouatta caraya), Paraguay B. mucronafa (p. 162). Strobila attains 84 mm. or more in length by 6.5 mm in breadth. Host: Bonnet mon- key (Macacus siuicus), India B. conferta (p. 163). 18. Strobila attains 130 mm. in length by 15 mm. in breadth by 2.5 mm. in thickness; geni- tal pore alternates very regularly; ova 53 to 60 //. Host: Chimpanzee (Aiithropo- pithecus troglodytes) B. studeri (p. 161). Strobila attains 245 mm. or more in length by 10 mm. in breadth by 2 mm. in thick- ness; pores irregularly alternate; cirrus pouch claviform; ova 35 to 38 /< by 30 to 32ju. Host: Oran-utau . (Simia satyrus) B. satyri (p. 161). Cittotainia. 19. Cirrus pouch muscular, generally pyriform, with distinct outline, and swollen proximally by the vesicula serainalis Marmota>-Group, 20 Cirrus pouch tubular, resembling the nozzle of a hose, of equal or nearly equal diameter throughout its entire length, and not swollen proximally by any prominent vesicula seminalis Pectinata-Group, 23 20. Testicles arranged in a band extending across the median field 21 Testicles comparatively few in number, absent from median line, arranged in two triangular groups, one corresponding to each ovary; cirrus pouch small, 0.16 mm. long, scarcely reaching the longitudinal nerve; pores in distal part of lateral margin; female glands near ventral canal; dorsal canal dorsal or dorso-lateral of ventral canal; strobila attains 80 cm. in length by 10 mm. in breadth. Host: Common wild and tame rabbits (Lepus cuniculus and L. cnui- culus domesticus), Europe C. ctenoides (p. 179). 21. Testicles not confined to space between the ovaries 22 Testicles confined to distal portion of segment between oraries; cirrus pouch about 0.5 mm. long by 0.17 mm. broad, crosses the canals ; female glands some distance from 1 Characters satisfactory for an analytical key are still lacking. NO. 1105. PROCEEDINGS OF THE NA TIONAL MUSE UM. 217 lateral canals ; transverse uterus single with proximal and distal pouches; dorsal canal 'between, ventral canal and nerve; stroblla may attain 112 mm. long by 5 to IS mm. broad. Host : Marmot ( Arctomys inarmota), Europe. C. marmotm ( p. 172). 22. Cirrus pouch -very large and prominent, may attain 1.12 mm. long by 0.32 mm. broad; pores at or near distal corner of lateral margin; female glands not far removed from dorsal canal; several accessory ventral canals; strobila may attain 80 cm. long by 15mm. broad. Host: Wild rabbit (Lepus cuniculus), Europe C. denticulata (p. 174). Cirrus pouch small, 0.24 mm. long; female glands considerably removed from dorsal canal; dorsal canal lateral or dorso-lateral of ventral canal ; reccptaculum seminis very large; segments imbricate; strobila attains 40 mm. long by 5.5 mm. broad. Host: Prairie gopher (Geornys bursarius), North America.. C. prwcoquis (p. 181). 23. Cirrus pouch about 1 mm. long, extends some distance median of longitudinal canals ; testicles arranged in a band in distal portion of median field, extend- ing beyond ovaries from canal to canal; ovary some distance from longitudi- nal canals; uterus single, with well-developed proximal and distal pouches; strobila attains 40 cm. in length by 8 to 10 mm. in breadth. Hosts: Common hare (Lepus timidus) and mountain hare (L. variabilis) Europe. C.pectinata (p. 184). Cirrus pouch not over 0.5 mm. long, extends scarcely median of lateral canals'; uterus single or double in the same strobila. Hosts : Lepus, North America. 24 24. Testicles in two groups, one for each ovary, extending laterally of ovaries but absent from median field ; strobila attains 57 mm. or more in length by 10 mm. in breadth. Host: Cottontail rabbit (L. sylraticus) C. perplexa (p. 189). Testicles in a band confined to distal portion of median field between the ovaries; strobila attains 100 to 180 mm. in length by 10 mm. in breadth. Host: Cottontail rabbit (L. sylvaticus) and marsh hare (L. palustris). C. variabilis (p. 190), 25. 25. Strobila nearly always more than 3 mm. broad ; posterior flap of segments nearly straight; segments slightly imbricate: genital pore in about the middle of the lateral margin. Host: Cottontail rabbit (L. sylvaticus). C. variabilis (p. 192). Strobila nearly always more than 3 mm. broad; posterior flap of segments lobed; segments imbricate; genital pores generally in distal half of lateral margin. Host : Marsh hare (L. palustris) C. variabilis imbricata (p. 193). Strobila rarely over 2 mm. broad; posterior flap straight; segments not imbri- cate; genital pores generally in distal portion of lateral margin, generally double, occasionally single. Host: Cottontail rabbit (L. sylvaticus'). C. variabilis angusta (p. 193). Andrya (European). 26. Genital pores near posterior corner of segment ; prostatic gland elongate ; head about 1 mm. in diameter; testicles comparatively few in number, confined chiefly to aporose side of median field; cirrus pouch 0.32 to 0.34 mm. long; strobila attains 60 to 80 cm. in length by 5 mm. in breadth. Host: Common hare ( Lepus timidus), Europe - A . rhopalocephala (p. 155). Genital pores in about the middle, or in the distal half of the lateral margin; prostatic gland round; head about 0.5 mm. in diameter; testicles about 50 in number, scattered through entire median field; cirrus pouch 0.4 to 0.48 mm. long; strobila attains 100 cm. in length by 8 mm. in breadth. Host: Wild rabbit (Lepus cuniculus), Europe - ^- cuniculi (p. 158). DipylidiincB. 27. Suckers unarmed - ** Suckers armed with minute booklets (with four exceptions known only in birds) 3 28. Genital pores double ^ 29 Genital pores single, irregularly alternate or unilateral 31 218 TAPE WORMS OF HARES AND RAB BITS STILES. VOL. xix. 29. Two submedian ovaries in each segment 30 One median ovary in each segment Amabilia. 1 30. Several rows of hooks upon rostellum Dipylidium. 2 A single row of hooks upon rosteUum Cotugnia.* 31. Dorsal root of hooks much longer than ventral root or prong ; ventral root very short; hooks 8 to 12 (rarely to 26} in number; known only in birds Drepanidotwnia.* Dorsal root of hooks about the same length as ventral root and prong 32 32. Dorsal root shorter than prong or ventral root ; hooks generally less than 20 in number (range from 10 to 26} ; genital pores unilateral or irregularly alternate ; known only in birds Dicranolamia. 5 Dorsal root longer than prong or ventral root ; if rosteUum is armed, there are 24 to 30 hooks present ; genital pores unilateral (on left of segments) ; three testicles nor- mally present in each segment ; retractile rostellum armed with minute hooklets or rudimentary and unarmed; eggs with three envelopes ; parasitic in man, chiroptera, insectivora, rodents, and insectivorous birds ; larval stage in insects or myriapoda. - Hymenolepis.* 33. Hooks on suckers arranged in circular rows on border; hooks on rostellum resem- ble a hammer in form and are arranged in a double row; with four exceptions known only in birds Davainea (p. 194), 35. '1893, Amabilia, DIAMARE, Boll. Soc. Nat. Napoli, 1. ser., VII (1-2), p. 13. August 28, 1893. Type, Tcenia lamelligera, Owen, 1835. ? 1896, Diploposthe, JACOBI, Zool. Auzeiger, XIX, No. 505, June 15, 1896, pp. 268-269. Type, Tcenia Icvvis, Diesiug. See Stiles, 1896, Bull. 12, Bureau of Animal Industry, p. 31. From the diagnosis given by Jacobi I fail to see how Diploposthe diifers from Amabilia. 1863, Dipylidium, R. LEUCKART, Die menschlicheu Parasiten, I, p. 400. Type, Tcenia elfyrtica, Batsch, 1786 = Tcvnia canina, Linnaeus, 1758. 1858, " Alyselminthus, ZEDER, 1800 pp.," of Weinland with Tcvnia cucumerina, Bloch, 1782 Tamia canina, Liumeus, 1758, as type. 1884, Micro tcenia, SEDGWICK, as subgenus in CLAUS & SEDGWICK, Elementary Text-book of Zoology, I, p. 336. Type, Tamia cucumerina, Bloch, 1782 = Tcenia canina, Linnaeus, 1758. For species and bibliography see Diamari's monograph (1893) II Gen ere Dipylidium, Lkt., Atti R. Accad. Scienze fisische e matematiche, 2 ser., VI, No. 7, 31 pp., pis. I-III. 3 1873, Cotugnia, DIAMARI, Boll. Soc. Nat. Napoli, 1. ser., VII (1-2), p. 11. August 28, 1893. Type, Tfenia digonopora, Pasquale, 1890. See Stiles, 1896, Bull. 12, Bureau of Animal Industry, p. 29. 4 1892, Drepanidotcenia, RAILLIET, Bull. Soc. zool. France, XVII, p. 116. Type, Tcmia lanceolata, Bloch, 1782. ? 1896, Choanotcenia, RAILHET, Rec. me"d. ve"t., 8 ser., Ill (5), March 15, 1896, p. 159. Type, Tcenia infundibuliformis, Goeze, 1782. For species and bibliography of this genus see Stiles, 1896, Bull. 12, Bureau of Animal Industry, pp. 36-45, 60-61. 6 1892, Dicranotosnia, RAILLIET, Bull. Soc. zool. France, XVII, p. 116. Type, Tcenia coronula, Dujardin, 1845. For species and bibliography of this genus see Stiles, 1896, Bull. 12, Bureau of Animal Industry, pp. 32-36. 6 1858, Hymenolepis, WEINLAND, Human Cestoides, p. 49, 52. Type, Tcenia Jlavopunc- tata, Weinland, 1858= Toenia diminuta, Rudolphi, 1819. 1858, Lepidotrias, WEINLAND, as subgenus of Hymenolepis, Human Cestoides, p. 52. Type, Tceniamurina, Dujardin, 1845, proposed by Weinland, but this subgenus includes the type of the genus. For species and bibliography of the genus see R. Blanchard, 1891, Hist. zool. et med. des Teiriades du genre Hymenolepis, Weinlaud. Paris. 112 pp., 22 figs. NO. 1105. PROCEEDINGS OF THE NATIONAL MUSEUM. 219 Hooks on stickers arranged in several transverse rows ; hooks of infundibulum very small (4 /<) arranged in a single row ; known only in birds Ophryocotyle. 1 Hooks on suckers arranged in one median set and two lateral sets ; hooks on rostellum provided with long dorsal root and arranged in a single row ; known only in birds. Ecliinocotyle* Subfamily f 34. Anterior extremity of strobila expanded in form of a hammer Fimbriaria.* A n terior segments become calci/orm and function as pseudoscolex Idiogenes. 4 Davainea. 35. Found in birds ; see Stiles, 1896, 5 p. 45. Found in mammals r 36 36. Genital pores unilateral ; a single egg in each egg capsule 37 Genital pores generally alternate; strobila attains 85 mm. or more in length by 3 mm. in breadth; suckers not invaginated; eggs grouped 3 to 15 in each egg capsule. Host: Eastern Jackass hare (Lepus melanotis) and Cottontail rabbit (L. sylvaticus) I), salmoni (p. 198). 37. Suckers (always?) invaginated; strobila attains 105mm. long by 3 mm. broad. Host : Arizona cottontail (L. arizonm) D. retractilis (p. 195). Suckers not invaginated 38 38. Rostellum armed with double row of about 90 hooks, IS /.i long ; strolnln attains 250 to 300 mm. long, composed of 500 to 600 segments. Host : Man (Homo sapiens). D. madagascariensis (p. 194). Rostellum large, armed with (?) a single row of numerous minute hooklets ; suckers armed with 8 to 10 rows of hooks. Host: Common Indian Pangolin (Manis peu- tadactyla) D. contorta (p. 195). COMPENDIUM OF THE PARASITES ARRANGED ACCORDING TO THEIR HOSTS. In the following list are given the hosts of the parasites discussed in this paper, so far as they are known to me. I have personally exam- ined the species starred (*) for the hosts under which they are given. For the name of the collector in each case see text. The numbers given with the names of the hosts are those of von Linstow's Compendium and Nachtrag. The geographical distribution refers to the host. One '1870, Ophryocotyle, FRIIS, Videnskab. Meddel. fra den Naturhist. Foren. Kjoben- havn [aar 1869], 1870, pp. 121-124, pi. I. Type, Ophryocotyle proteus, Friis, 1870. R. Blaiichard gives the date as 1869, Braun gives it 1870. Original not at my dis- posal. For species see Stiles, 1896, Bull. 12, Bureau of Animal Industry, p. 56. 2 1891, Echinocotyle, R. BLANCHARD, Bull. Soc. zool. France, IV, p. 423. Type, E. rosseteri, R. Blan chard, 1891. 3 1802, Fimbriaria, FROLICH, Der Naturforscher, XXIX, pp. 13-14. Type, Tanta malleus, Goeze, 1782. 1850, Ithynchotwnia, DIESING (as section or subgenus of Tojnia}, Systema Helmiii- thum, I, p. 521. Type, Twnia malleus, Goeze, 1782. 1892, Epision, LINTON, Proc. U. S. Nat. Mus., XV, p. 100. Type, Epision plicatus, Linton, 1892. 4 1868, Idiogenes, KRABBE, Videns. Meddel. fra d. Naturhist. Foren. Kjobeuhavn (for Aaret 1867), 1868, pp. 122-126. Type, Idiogenes otidis, KRABBE, 1868. 6 Stiles, 1896, Report upon the Present Knowledge of the Tapeworms of Poultry, Bull. 12, Bureau of Animal Industry, U. S. Department of Agriculture. 220 TAPEWORMS OF HARES AND RABBITS STILES. VOL.XIX. or two parasites not discussed in this paper are included in the com- pendium for reasons obvious in each case. 1 have elsewhere (1896) made a plea for the adoption of a modern sys- tem of nomenclature for the hosts, and have endeavored to introduce here the correct names for most of the hosts cited. In the main, Flower and Lydekker's Mammals Living and Extinct (1891) has been followed; in the few cases that I have departed from the names given in that work I have done so upon the advice of Dr. T. S. Palmer. In order to pre- vent confusion by this change in host names., I have added the names used by von Linstow in his compendium or by other authors in their writings, cross-referencing the synonyms to the proper name. It is impossible to give a monographic list of the genus Lepus and to estab- lish the correct names of all the forms until that genus is revised, by a specialist in mammalogy. ALOUATTA CARAYA (Humboldt, 1811). Black Howler. (Stentor caraya, Humboldt; 20. Mycetes niger ; 31. Celms caraya, Fischer). (Southern Brazil, Paraguay, Bolivia.) * Bertia mucronata. Tcenia megastoma. ANTHROPOPITHECUS TROGLODYTES (Linnaeus, 1758). Chimpanzee. (2a. Simia troglodytes ; Troglodytes niger Geoff.) (Western and Central Equatorial Africa.) Bertia studeri. 92. ARCTOMYS MARMOTA (Linnaeus, 1758). Alpine Marmot. (Mountains of southern Europe Alps, Pyrenees, and Carpathians.) * CittotoRnia marmottv. 926. ARCTOMYS, sp. ? (Turkestan.) Anoplocephala transversaria. 31. CEBUS CARAYA, Fischer vide ALOUATTA CARAYA. 30. CEBUS CAPUCINUS (Linnaeus, 1758). Weeping Capnchin. (Paraguay to United States of Colombia.) "Taenia No. 2, Gottheil," p. 164. 248. EQUUS CABALLUS (Linnaeus, 1758). Horse. *AnoplocepJiala mamillana. Anoplocephala perfoliata. Anoplocephala plicata. Anoplocephala plicata pediculata. Anoplocephala plicata strangulata. 124. ERETHIZON DORSATUS (Linnaeus, 1758). Canada Porcupine. (Boreal region of eastern North America.) * Bertia americana. Tosnia laticephala, Leidy. ? " Tcenia pectinata" with unilateral pores=?, recorded by Cobbold, 1862. ERETHIZON EPIXANTHUS, Brant. Yellow-haired Porcupine. (Mountains of Western United States.) * Bertia americana. GALEOPITHECUS VOLANS (Linnaeus, 1758). Flying Lemur, Common Colngo or Cobego. (Sumatra, Borneo, Java, Malay Peninsula, Tenasserim, and Siain.) Bertia plastica. GEOMYS BURSARIUS (Shaw, 1800). Prairie Gopher. (Upper Mississippi Valley, southward to southern Illinois, Missouri, and eastern Kansas.) * Cittotcenia prcecoquis. PROCEEDINGS OF THE NATIONAL MUSEUM. 221 1. HOMO SAPIENS, Linmeus, 1758. Man. BothriocepUalus cordatns. * Bothrloceplialus latns. Davainea madagascariensis Dipylidium caninum. Hymcnolepis diminuta. Hymenolepis murina. Krabbea grandis* *Tcenia confasa. *Tcenia saginata. *Tamia solium. 12. INUUS CYNOMOLGUS vide MACACUS CYNOMOLGUS. LEPUS AMERICANUS, Erxleben, 1777. Northern Hare. (Wooded districts, New England to Minnesota, and south to Virginia, along the Alleghanies. ) ? " Toenia pectinata," reported by Curtice, ' 1892, p. 232. LEPUS ARIZONA, Allen, 1877. Arizona Cottontail. (Arizona, southern Nevada, and desert region of southern California.) * Davainea retractilis. 137. LEPUS CUNICULUS. Common European Wild Rabbit. Andrya cuniculi. See p. 155. * Anoplocephala wimerosa. *Cittotamia ctenoides. *Cittota'nia denticulata. 137a. LEPUS CUNICULUS DOMESTICUS. Common Domesticated Rabbit. *Cittot(vnia ctenoides. ? " Tcema pectinata," * reported by Curtice, 1892, p. 232. LEPUS MELANOTIS, Mearns, 1890. Eastern Jackass Hare. (Plains from Texas to Nebraska, west to Rocky Mountains.) * Davainea salmoni. LEPUS PALUSTRIS, Bachrnan, 1837. Marsh Hare. (Marshy lowlands of South Atlantic States, from North Carolina southward.) *Cittot(vnia variabilis imbricata. 138. LEPUS SYLVATICUS, Bachman, 1837. Cottontail. (Eastern United States.) *CUtot(Knia perplexa. *CitlotcKnia variabilis. *Cittotce / nia variabilis angusta. ^Davainea salmoni. LEPUS TEXIANUS, Waterhouse, 1848. Black-Tailed Jack-Rabbit. (Great Basin. ) "An undescribed Tamia," 2 reported by Curtice, 1892, p. 233. 140. LEPUS TIMIDUS, Linnajus, 1758. Common European Hare. (All parts of Europe except the north of Russia, the Scandinavian peninsula, and Ireland.) ? Andrya cuniculi. See p. 155. * Andrya rhopalocephala. * Cittotamia pectinata. 139. LEPUS VARiAiiiLis, Pallas, 1778. Mountain Hare. (Northern Eurasia.) * AnoplocepJiala ivimerosa. *Cittotcenia pectinata. l l am unable to trace this specimen, but most of Curtice's T. pectinata is C. variabilis. -Impossible to tell definitely what species Curtice referred to, but possibly the parasite is Davainea salmoni and the host L. melanotis. 222 TAPEWORMS OF HARES AND RABBITS STILES. LEPUS, SP. ? (Some North American species.) *Bertia americana leporis. ?LEPUS SP.? (?WASHINGTONI, S. F. Baird, 1855). Western Varying Hare. (Puget Sound.) * Cittotwnia (?) varidbilis. MACACUS CYNOMOLGUS (Linnaeus, 1758). Crab-eating Macaque. (12. Innus eynomoh/us.) (Malay Peninsiila and Philippine Archipelago.) " Tcenia No. 1, Gottheil." See p. 1S4. " Tcenia No. 2, Gottheil." See p. 164. MACACUS sixicus (Linnaeus, 1771). Bonnet Monkey. (Southern India.) Bertia conferta. MAXIS PEXTADACTYLA. Common Indian Pangolin. (India aud Ceylon.) Davainea contorta. 20. MYCETES NIGER, vide ALOUATTA CARAYA. 3. PlTHECUS SATYRUS, vide SlMIA SATYRUS. SIMIA SATYRUS, Linnaeus, 1758. Orang or Oran-Utan. (3. Pitfiecus satyrus.) (Sumatra and Borneo.) Bertia satyr i. 2a. SIMIA TROGLODYTES, vide ANTHROPOPITHECUS TROGLODYTES. TROGLODYTES NIGER, vide AXTHROPOPITIIECUS TROGLODYTES. BIBLIOGRAPHY By ALBERT HASSALL, M. R. C. V. S. [If the date of publication under the author's name is inclosed in parentheses, the citation has not been verified.] BAIRD, W. 1853. Catalogue of the species of Ento- zoa or Intestinal Worms contained in the Collection of the British Museum, London. 132 pp., Tab. I-II. DE BLAINVILLE. 1828. Art., Vers. Dictionnairedes Sci- ences naturelles, Strasbourg, Tom. LVII,pp.365-625,Pls.XXVII-XLVIII. BLANCHARD, E. 1848. Recherches sur 1'organisation des vers. Ann. des Sci. uat., 3 s st% V ^.^r-y-Vr'* >.?> r ANOPLOCEPHALINE TAPEWORM FROM THE COTTONTAIL RABBIT. FOR EXPLANATION OF PLATE SEE PAGE 231. U. S. NATIONAL MUSEUM PROCEEDINGS, VOL. XIX PL. XXI ARMED TAPEWORM FROM THE ARIZONA COTTONTAIL RABBIT. FOR EXPLANATION OF PLATE SEE PAGE 231. U. S. NATIONAL MUSEUM ti ve PROCEEDINGS, VOL. XIX PL. XXII ci ARMED TAPEWORMS FROM THE ARIZONA COTTONTAIL RABBIT, THE COMMON INDIAN PANGOLIN, AND THE COTTONTAIL RABBIT. FOR EXPLANATION OF PLATE SEE PAGE 231. U. S. NATIONAL MUSEUM PROCEEDINGS, VOL. XIX PL. XXII ARMED TAPEWORM FROM THE COTTONTAIL RABBIT AND THE EASTERN JACKASS HARE. FOR EXPLANATION OF PLATE SEE PAGE 232. U. S. NATIONAL MUSEUM PROCEEDINGS, VOL. XIX PL. XXIV ARMED TAPEWORM FROM THE EASTERN JACKASS HARE. FOR EXPLANATION OF PLATE SEE PAGE 232. U. S. NATIONAL MUSEUM PROCEEDINGS, VOL. XIX PL. XXV w 5 YOUNG STAGES OF AN ARMED TAPEWORM FROM THE COTTONTAIL RABBIT. FOR EXPLANASION OF PLATE SEE PAGE 232.