A New Subfamily, Two New Genera, and Three New Species of Haploporid Trematodes1

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1 112 PROCEEDINGS OF THE HELMINTHOLOGICAL SOCIETY shrews could not be discerned. Infected animals were obtained during all months but September and October. The lowest annual rates would be expected in autumn, when the populations of shrews consist predominantly of young animals. Acknowledgments We express thanks to Mr. and Mrs. Chris Maser, who provided the material studied as well as a record of their field observations, and to Dr. B. Horning, Vet.-Bakteriologisches und Parasitologisches Institut, Universitat Bern, Switzerland, who provided the otherwise unavailable description of C. rent. Literature Cited Arzamasov, I. T., I. V. Merkusheva, O. N. Mikholap, and I. V. Chikilevskaia Nasekomoiadnye i ikh parazity na territorii Belorussii. Nauka i Tekhnika, Minsk, 175 p. Franklin, J. F., and C. T. Dyrness Vegetation of Oregon and Washington. US- DA Forest Service Res. Paper PNW-80, Portland, Oregon, 216 p. Furmaga, S Internal parasites of the mole (Talpa europaea L.) in the Lublin environment. Acta Parasit. Polon. 7: Johnson, M. L., and B. T. Ostenson Comments on the nomenclature of some mammals of the Pacific Northwest. J. Mammal. 40: Lopez-Neyra, C. R Los Capillarinae. Mem. Real Acad. de Ciencias, Ser. Ciencias Nat., Madrid 12: Nadtochii, E. V., and T. T. Rasskazova Nematody zemleroek nekotorykh territorii Dal'nego Vostoka. In E. V. Gvozdev (ed.), Biologicheskie Problemy Severa, p Trudy Severo-vostochnogo Komplekskogo Inst., Akad. Nauk SSSR, Magadan, 239 p. Prokopic, J Cizopasni cervi nasich hmyzozravcu. Systematicko-faunisticka studie a ekologicko-zoogeograficke zhodnoceni. Cesk. Parasit. 6: , and V. Mahnert Uber Helminthen der Kleinsauger (Insectivora, Rodentia) Tirols (Osterreichs). Ber. Nat.-Med. Ver. Innsbruck 58: Rukhliadeva, M. N K izucheniiu nematod roda Capillaria Zeder, 1800, ot kutory (Neomys fodiens Pall.). In V. P. Pod'iapol'- skaia (ed.), Gel'mintologicheskii Sbornik, p Akad. Nauk SSSR, Moscow-Leningrad, 302 p. Soltys, A Helminthofauna of Soricidae in the Bialowieza National Park. Acta Parasit. Polon. 1: Wakelin, D Nematodes of the genus Capillaria Zeder, 1800 from the collection of the London School of Hygiene and Tropical Medicine. III. Capillariids from mammalian hosts. J. Helminthol. 42: A New Subfamily, Two New Genera, and Three New Species of Haploporid Trematodes1 W. E. MARTIN Department of Biological Sciences, University of Southern California, Los Angeles, California ABSTRACT: A new subfamily, Unisaccinae, of the family Haploporidae is established to include the new genus Unisaccus and the new species, U. brisbanensis and U. spinosus; and the new genus and species Unisaccoides vitellosus found in the intestines of mullet in the Brisbane River, Queensland, Australia. These worms are unique among haploporids in having a single cecum. During sabbatical leave ( ) spent in the Parasitology Department, University of Queensland, Brisbane, Australia, Brisbane River mullet were found to harbor in their small 1 Supported by NSF G6962. intestines some trematodes that are unique among haploporids in the possession of an undivided cecum. They are described as two new genera and three new species of a new subfamily.

2 OF WASHINGTON, VOLUME 40, NUMBER 1, JANUARY The worms were fixed without pressure by injecting them into hot 5% formalin. They were stained with Mayer's iron alum or Mayer's paracarmine. Living specimens were also studied. All measurements are in microns unless designated otherwise; averages are in parentheses. Unisaccinae n. subfamily DIAGNOSIS: Haploporidae. Body small, tegument spinose. Eyespot remnants present. Oral sucker subterminal, acetabulum in anterior half of body. Cecum undivided. Hermaphroditic duct with or without spirally arranged pads bearing spines. Internal and external seminal vesicles present, the former enclosed along with prostate cells and prostatic bulb in hermaphroditic sac. Testis single in posterior half of body. Vitellaria follicular. Ovary immediately anterior to testis. Laurer's canal and seminal receptacle uterinum present. Embryonated egg with eyespotted miracidium (as far as known). Excretory bladder Y-shaped with sphincter near exit. Type genus Unisaccus. Unisaccus gen. n. DIAGNOSIS: Unisaccinae. Body fusiform, oral sucker subterminal, acetabulum in anterior half of body. Prepharynx, pharynx, and esophagus well developed. Cecal sac lobed or nonlobed. Vitelline follicles few, lateral between levels of testis and acetabulum. Ovary small, simple, near anterior margin of testis. Laurer's canal and receptaculum seminis uterinum present. Uterus extends to near posterior end of body. Eggs relatively large and numerous, each mature one with eyespotted miracidium containing the outline of a redia. Testis in posterior half of body. Hermaphroditic duct with spined pads. Genital pore midventral between acetabulum and pharynx. Type species Unisaccus brisbancnsis. Unisaccus brisbanensis sp. n. (Figs. 1-3, 6) DESCRIPTION (measurements based on 24 mounted specimens): With characters of the genus. Tegument spined to near posterior end of body. Body length (780), width (320). Oral sucker (75) long, (90) wide; acetabulum (82) in diam; prepharynx and esophagus subequal, (230) long; pharynx (35) long, (55) wide; cecum may bear shallow pouches at corners. Hermaphroditic sac (129) long, (150) wide; hermaphroditic duct with spirally arranged pads about 20 long, each bearing two spines about 7 long and supported by a delicate sclerotized basal lattice about 7 long. Internal seminal vesicle tubular; external vesicle saccular and filled with sperm in mature specimens. Sperm about 142 long. Testis oval to spherical, (122) long, (81) wide; ovary median, oval to spherical, (16) long, (17) wide: Laurer's canal opens dorsally posterior to ovary, with proximal bulb containing cilia which beat toward exit. Vitelline follicles 4 to 6. Uterus from near posterior end of body to acetabular level. Eggs operculate, with thin yellow shells (58) long, (27) wide. Arms of excretory bladder longer than stem, bladder often containing a small concretion. Holotype deposited as No. 715 in the Hancock Parasitology Collection, University of Southern California. HOST: Mugil cephalus L. SITE: Small intestine. LOCALITY: Brisbane River, Brisbane, Queensland, Australia. Unisaccus spinosus sp. n. (Figs. 4, 5, 8) DESCRIPTION (measurements based on 55 mounted specimens): With characters of genus. Tegument spined, body (700) long, (274) wide. Mouth subterminal; oral sucker (159) long, (192) wide; acetabulum spined, (131) in diameter. Prepharynx (248) long; pharynx (91) long, (174) wide; esophagus about as long as pharynx; cecum with shallow pockets at corners. Testis nearly spherical to oblong, (171) long, (168) wide. External seminal vesicle elongate and saccular. Hermaphroditic duct lining bears spiraling rows of pads, each pad with two or rarely three spines on a reticular sclerotized base. Ovary nearly spherical, (61) in diam. Laurer's canal as in preceding species. Eggs with thin,

3 114 PROCEEDINGS OF THE HELMINTHOLOGICAL SOCIETY Figures Ventral view of Unisaccus brisbanemis. A, acetabulum; C, cccum; E, excretory bladder; ES, eyespot remnants; EV, external seminal vesicle; GP, genital pore; H, hermaphroditic sac; O, ovary; P, pharynx; S, oral sucker; T, testis; V, vitellaria. 2. Hermaphroditic sac enclosing internal seminal vesicle, prostate bulb, and spined hermaphroditic duct. 3. Optical section of cecum.

4 OF WASHINGTON, VOLUME 40, NUMBER 1, JANUARY llmrn Figures Ventral view of Unisaccus spinosus. Symbols same as Figure Lateral view of 17. spinosus. 6. Spine pad, spines, and sclerotized support of U. brisbanensis. Spine pads of U. spinosus similar but smaller and more numerous.

5 116 PROCEEDINGS OF THE HELMINTHOLOGICAL SOCIETY Figures Hermaphroditic sac of V. spinosus enclosing spined hermaphroditic duct (HD), internal seminal vesicle (IV), and prostate bulb (PB). 8. Miracidium of V. spinosus with enclosed redia. 9. Vnisaccoides vitellosus, ventral view.

6 OF WASHINGTON, VOLUME 40, NUMBER 1, JANUARY yellow, operculated shells, (102) long and (53) wide; embryonated egg with ocellate miracidium containing a well-developed redia about 37 long and 25 wide, with pharynx,1 about 10 long and 12 wide. Hermaphroditic sac (260) long, (210) wide. Genital pore immediately anterior to acetabulum. Excretory bladder with fairly long stem, arms reaching to level of vitellaria. Holotype deposited as No. 717 in the Hancock Parasitology Collection. HOST: Lisa argentea (Quoy and Gaimard). SITE: Small intestine. LOCALITY: Brisbane River, Brisbane, Queensland, Australia. Unisaccoides gen. n. DIAGNOSIS: As in Unisaccus except vitelline follicles numerous, mainly lateral, from hermaphroditic sac to near posterior end of body where they may become confluent; uterus from testis to hermaphroditic sac. Type species Unisaccoides vitellosus. Unisaccoides vitellosus sp. n. (Fig. 9) DESCRIPTION (measurements based on four specimens, only one ovigerous): Body (650) long, (217) wide. Oral sucker (67) long, (77) wide. Prepharynx (99) long; pharynx (60) long, (85) wide; esophagus about 60 long; cecum (115) long, about 112 wide. Acetabular diameter (66). Hermaphroditic sac about 90 long, 50 wide. Ovary (47) in diam; testis (120) long, (76) wide. Eggs with thin, yellow operculated shells, (62) long and about 42 wide; none showed miracidia but perhaps would if older. Stem of excretory bladder shorter than arms which reach to near the acetabular level. Holotype deposited as No. 716 in the Hancock Parasitology Collection. HOSTS: Mugil cephalus and Lisa argentea. SITE: Small intestine. LOCALITY: Brisbane River, Brisbane, Queensland, Australia. Discussion Attempts to determine the life cycles were made by exposing laboratory-reared snails, Posticobia brazieri (Smith), to the miracidia of Unisaccus brisbanensis and U. spinosus but none became infected. Posticobia brazieri was chosen because it serves as intermediate host for some other haploporids (Martin, 1973). The description of the digestive tract in the family Haploporidae must be emended to include those with an undivided cecum. In young adults of the species described in this paper the cecum lacks the lobes of older specimens. Evidently the cercaria has a single cecum. Also the cercaria must bear eyespots since all adults showed remnants of these structures. The single saccular cecum might indicate a fairly close relationship to the Haplosplanchnidae in which the cecum is tubular and usually longer. Certain species of Haploporidae and Haplosplanchnidae develop in the same species of mullet. While alive, specimens of the trematodes described above were seen to evert the hermaphroditic duct through the genital pore in a manner suggesting its functioning as a male copulatory organ. When not everted, the hermaphroditic duct may serve as a vagina. After the specimen shown in Figure 9 was drawn, it was accidentally crushed while removing immersion oil. Additional fish were examined and another specimen was found that contained eggs. This was deposited as the type specimen although it does not show some structures as clearly as the original specimen because of the eggs. Unfortunately the eggs collapsed during fixation so their measurements are as near approximations to actual size as could be obtained. Acknowledgments I am greatly indebted to Professor J. F. A. Sprent, Head of the Parasitology Department, University of Queensland, for the use of laboratory facilities and encouragement in many ways; to Dr. John Pearson, Reader in Parasitology, for furnishing laboratory-reared snails and help in many other ways; to Miss Marilyn Gadisckie for assistance in making slides; and to Mr. Jim Davie for help in collecting fish. Literature Cited Martin, W. E. Life cycle of Saccocoelioides pearsoni n. sp. and the description of Lecithobotrys sprenti n. sp. (Trematoda: Haploporidae). Trans. Am. Microsc. Soc. 92:

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