I (PI I ,Cf PROJECT PNW-1204 CULTURE OF MIXED CONIFERS •15.1:4=41;;;=• WEST SIDE CASCADES

- OF WASHINGTON, VOLUME 40, NUMBER 1, JANUARY 1973 • 107

Ph:uornenes de convergence chez les spirur- logical convergence. J. Helminthol. 39: 207- ides, en particulier clans lee sous-familles Ha- 2x4. bronematinae Chitwood and Wehr, 1932 et Mawson, P. M. 1968. Habronematinae (Nema- Schistorophorinae L. Travassos, 1918; leur toda: Spiruridae) from Australian birds. Par- importance por une classification naturelle asitology 58: 745-767. des spirurides (Nematoda). Bull. Soc. Zool. Yeh, L. S. 1954. On two new species of the Fr. 82: 88-102. Serticeps (Nematoda: Schistorophidae) Inglis, W. G. 1965. The nematodes parasitic from the gizzard of birds. J. Helminthol. 28: in the gizzard of birds: A study in morpho- 165-170.

Capillaria maseri sp. n. (Nematoda) from Insectivores (Soricidae and ) in Oregon

R. L. RAUSCH AND V. R. RAZSCH Arctic Health Research Cznter, Bureau of Community Environmental Management, Health Services and Mental Health Administration, Department of Health, Education, and Welfare, Fairhanks, Alaska

13 sr nacr : Capillaria maseri so. n. is described from the urinary bladder of , Sores spp. (type host: S. yaquinae Jackson) in Oregon, and is recorded also from the -mole, Neiirotrichus gibbsii (Baird). C. maseri is the first capillariid to be reported from the urinary bladder of North American insectivores_

In 1970-72, nematodes of the genus Capil- larly prepared from the normal urinary bladder laria Zeder, 1800, were found in the urinary of a shrew collected in Alaska. The hosts were bladder of insectivores collected in Oregon by identified by Mr. Maser. The nomenclature of Chris and Rita Maser, Oregon Coast Ecological shrews is in accordance with the recommenda- Survey, Puget Sound Museum of Natural His- tions of Johnson and Ostenson (1959). tory, who kindly sent the specimens to us for study. The species involved differs from any Results recorded from insectivores in Eurasia and Description North America and is described herein. Numbers of specimens from which the Materials and Methods respective measurements were derived are shown in parentheses. All measurements are Most of the specimens had been fixed in in millimeters. situ in 10% formalin solution. Fifty-six were- studied in detail, including some that were Capillaria maseri sp. n. broken in dissecting the anterior ends from (Figs. 1-8) tissue of the urinary bladder. They were cleared by evaporation of a mixture of 2% Slender nematodes; white in color when pre- glycerin in 70% ethanol, after which selected served. Bacillary lines present. Mouth simple. specimens were stained in light green SF dis- Cuticle smooth. solved in glycerin. Two bladders with nema- MALE (23 specimens; 8 intact): Length todes in situ were embedded in paraffin, sec- (8) 7.3 to 20.5 (avg 17.1). Maximum width tioned at 0.005 or 0.010 mm, and stained in (22) 0.099 to 0.155 (avg 0.123), attained in hematoxylin—eosin, in Mallorys aniline blue posterior third of body: Width at cephalic collagen stain, or by the periodic acid-Schiff extremity (8) 0.008 to 0.011 (avg 0.010); reaction. For comparison, sections were simi- width at posterior extremity (23) 0.031. to

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0

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6414

0 0

0.075 mm OF WASHINGTON, VOLUME 40, NUMBER 1, JANUARY 1973 109

0.036 (avg 0.032). Esophagus (8) 3.7 to 7.9 Discussion long (avg 6.5); width at base (19) 0.049 to With the exception of C. hepatica (Ban- 0.065 (avg 0.056). Length of muscular esoph- croft, 1893), which has been reported from agus (4) 0.356 to 0.389 (avg 0.375). Ratio of diverse groups, species of Capi/- of length of esophagus to length of body (8) /aria from mammals are usually host-specific. 1:1.9 to 1:3.1 (ava 1;2.6). Twenty-six to 48 That certain species have been found in hosts esophageal cells observed. Distance from pos- representing two or more related families indi- terior end of esophagus to testis (19) 0.356 cates that their occurrence may be influenced to 1.178 (avg 0.645). Length of spicule (23) by ecologic, as well as phylogenetic, factors. 0.710 to 1.134 (avg 0.970). Spicule sheath Such is the case with C. maseri, occurring in aspinose, with minute folds. Caudal alae not both shrews (Soricidae) and moles (Talpidae). present. Hol.vever, in any host infected, the respective FEMALE (33 specimens; 10 intact): Length species appear to be organ-specific. In Eurasia, (10) 7.7 to 26.8 (avg 18.4). Maximum width nematodes of the genus Capillaria have been (32) 0.073 to 0.168 (avg 0.145), attained in found in the urinary bladder of moles and posterior third of body (containing uterus). shrews, but none has been reported from this Width at cephalic extremity (7) 0.008 to organ in North American insectivores. The 0.011 (avg 0.010). Width at posterior ex- two species known from insectivores in North tremity (33) 0.032 to 0.057 (avg 0.044). America, C. rauschi Read, 1949, and C. Width at level of vulva (30) 0.057 to 0.121 blarinae Ogren, 1953, were described from (avg 0.092). Esophagus (10) 2.93 to 10.4 the small intestine of cinereus Kerr and (avg 7.5) long; width at base (28) 0.049 to the esophagus of Blarina brevicauda (Say), 0.070 (avg 0.062). Length of muscular esoph- respectively. Wakelin (1968) suggested that agus (5) 0.378 to 0.470 (avg. 0.434). Thirty- C. rauschi is possibly identical with C. kutori nine to 41 esophageal cells observed. Ratio Rukhliadeva, 1946, described from the stomach of length of esophagus to length of body (10) of water shrews, Neomys fodiens (Pennant), 1:1.6 to 1:2.9 (avg 1:2.5). Ner •e ring (5) from the Khopersk Reserve, European Russia. ca. 0.090-0.115 from anterior end. Distance His hypothesis, if valid, would add another from posterior end of esophagus to vulva (28) species of the genus Capillaria from mammals 0.518 to 1.871 (avg 1.145). Distance from to those few lcnown to be holarctic, of which cephalic extremity to vulva (9) 3.5 to 11.4 none is from insectivores. It seems improbable, (avg 9.0). Anus terminal. Egg (178) 0.060 on zoogeographic grounds, that the two are to 0.073 by 0.024 to 0.032 (avg 0.065 by conspecific; unfortunately, a conclusion cannot 0.028); shell smooth; polar plugs well defined. be made from the descriptions of C. kutori TYPE itosr: Sorex yaquinae Jackson. Other (Rukhliadeva, 1946; Soltys, 1954). C. kuton hosts S. vagrans Baird, S. bendirii (Merriam), appears to be a common parasite of shrews in S. trotvbridgii Baird, S. pacificus Coues, and eastern Europe (Prokopfe, 1959; Arzamasov Neiirotrichus gibbsii et al., 1969), and it was identified by Wakelin (Baird). (1968) from the intestine of Sorex araneus TYPE LOC.ALITY: Cascade Head Experi- - in England. mental Forest, Lincoln County, Oregon. HABITAT: Urinary bladder. Comparisons TYPES: USNM Helm. Coll. No. 72256 A distinctive morphological characteristic of holotype (male) and No. 72257 allotype; Nos. C. rnaseri is the relatively great distance be- 72258 and 72259 (paratypes). tween the end of the esophagus and the vulva

Figures 1-8. Capillaria maseri sp. n. I. Anterior end of female. 2. En face view of cephalic end of male, slightly oblique. 3. Posterior end of niale. 4. Caudal extremity of male, with partially everted spicule. 5. Terminal portion of esophagus, female. 6. Details of vulva and associated structures. 7. Caudal extremity of female. 8. Structure of egg.

110 • PROCEEDINGS OF THE HELMINTHOLOGICAL SOCIETY

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(avg 1.145 mm). C. maseri differs from the Nadtochii and Rasskazova (1971). Both spe- two species previously described from North cies have been found in western Eurasia as American shrews in other characters as well well as in Kamchatka, in northeastern Siberia as in habitat. From C. rauschi, it is distin- (Nadtochii and Rasskazova, 1971). guished by the presence of bacillary lines, C. urinicola Soltys, 1952, described from smooth cuticle, different position of the vulva, shrews in Poland, was later considered by and smooth-shelled eggs of larger size; from Soltys (1954) to be identical with C. capillaris. C. blarinae by the larger size, longer esopha- However, comparisons by Furmaga (1959) led gus, longer spicule, absence of spines in spic- to the conclusion that C. urinicola is distinct. ule sheath, and, in females, by the presence of The similarity between C. sunci Chen, 1937, an external vulvar process and a terminal anal described from Sun= caeruleus (= S. murinus opening. L.) (Soricidae) in China, and C. incrassata Several species of Capillaria occurring in was recognized by LOpez-Neyra (1947, p. the urinary bladder of insectivores have been 140). Wakelin (1968, p. 392) stated that described in Eurasia. C. capillaris (von this species ". . . may well be considered a Linstow, 1882) and C. incrassata (Diesing, synonym of C. incrassata." • 1551) have been recorded from both moles C. reni Shaldybin, 1968, described from and shrews. Wakelin (1068) concluded that water shrews from the Mordovsk Reserve in C. capillaris could be synonymized with C. European Russia, might also be considered incrassata, a determination that was not ac- here, although it was found in the renal pelvis cepted by ProkopiZ:; and Mahnert (1970) or rather than in the urinary bladder. OF WASHINGTON, VOLUME 40, NUMBER 1, JANUARY 1973 •

C. maser( may be distinguished as follows Table 1. Occurrence of C. maseri in shrews and from these other species inhabiting the uri- nary system of insectivores: from C. capillaris by its larger size, longer esophagus, vulva Host No. examined No. infected situated farther posteriorly, and smooth-shelled Soricidae 138 43 Soren vogran., 16 10 egg; from C. incrassata by its larger size, more S. pacificw 27 5 S. yaquinae 23 9 slender anterior end, smooth cuticle, and longer S. bendini 13 3 spicule; from C. urinicola by its larger size, S. troirbridgii 59 16 longer esophagus, presence of vulvar process, Talpidae 40 1 Neiirotrichus 7 1 and a much larger egg; from C. rent, of simi- ararius 17 — lar large size, by its longer spicule and, in the S. town.sentiii 16 female, by its terminal anal opening and smaller egg. The description of C. real does not permit comparison of other anatomical mediately surrounded by normal-appearing details. epithelial cells; leukocyte infiltration, fibrosis, or other evidence of an inflammatory response Parasite—host relationships were not observed. No eggs were found within the tissue of the host. The tissue reaction, if C. maseri occurred in 43 (31%) of 138 any, evoked by other species of Capillaria in- shrews collected during November 1970—April habiting the urinary bladder of insectivores 1972. It was found also in one of seven shrew- apparently has not been described. A slide moles, Neiirotricims but in none of 16 Townsends moles, Scapanus townsendii (Bach- with sections of the nematode in situ has been man), orarius True. deposited in the USNM Helm. Coll., No. or in 17 coast moles, S. 72260. No early-stage infections involving In 32 infected shrews, not segregated by spe- newly established nematodes were detected, cies, numbers of nematodes ranged from 1 to although a few immature specimens were 29, with an average of 4. The ratio of males found. In the shrew-mole, one of the nema- to females in this series was 1:1.7. Five speci- todes was within the ureter; however, migra- mens were found in the bladder of the shrew- tion from the bladder might have taken place mole. Findings in mammals of the respective after the death of the host. species are summarized in Table 1. Mammals Most of the insectivores were captured along of both sexes were infected. the coast of Oregon in the vegetational zone The nematodes formed localized aggrega- characterized by Sitka spruce, a region of low tions on the floor of the urinary bladder in the elevation, mild climate, and high annual pre- vicinity of the neck, with the anterior portion cipitation (Franklin and Dyrness, 1969). Of of their bodies deeply embedded in the the five species of shrews involved, S. trow- mucosa. Males penetrated to the full length bridgii, S. pac-ificus, and S. yaquinae of the esophagus, females to the level of the inhabit forest and are nocturnal. S. bendirii is found vulva, permitting discharge of eggs directly near water, while S. vagrans inhabits meadows into the lumen of the bladder. In sections of as well as grassy areas along streams; both are incised, contracted bladders, in which the mu- cosa was thrown into deep, irregular folds, the active at any time of day. During the drier months (summer and autumn), all but S. anterior portions of the nematodes extended cagrans erratically through the thick epithelial layer, were usually trapped within 25 in of streams. The shrew-mole has shrewlike habits; but not penetrating the connective tissue of it usually occurs in forest and is active at any the lamina propria, nor did they pass through time of day. The infected insectivores were the mucosa into the underlying muscularis trapped in moist habitat in seral vegetation. (Fig. 9). No observations were made on re- C. maser( was found also in a single specimen lationships in the distended bladder, in which of S. bendirii from the western foothills of the the respective layers of tissue would be much Cascade Range (Lane County), at an elevation flattened. Findings in tissue sections indicate of about 1,000 m. (The foregoing data were that C. maseri is not pathogenic in shrews. The provided by C. Maser.) embedded portions of the nematodes were im- Seasonal differences in rates of infection in 112 • PROCEEDINGS OF THE HELMINTHOLOGICAL SOCIETY

shrews could not be discerned. Infected ani- mammals of the Pacific Northwest. J. Mam- mals were obtained during all months but mal. 40: 571-577. September and October. The lowest annual Lopez-Neyra, C. R. 1947. Los Capillarinae. Mem. Real Acad. de Ciencias Ser. Ciencias rates would be expected in autumn, when the Nat., Madrid 12: 1-248. populations of shrews consist predominantly Nadtochii, E. V., and T. T. Rasskazova. of young . 1971. Nematody zemleroek rtekotorykh ter- ritorii DaFnego Vostoka. In E. V. Gvozdev Acknowledgments (ed.), Biologicheskie Problerny Severn, p. 93- We express thanks to Mr. and Mrs. Chris 99. Trudy Severo-vostochnogo Komplekskogo Inst., Akad. Nauk SSSR, Magadan, 239 p. Maser, who provided the material studied as Prokopie, J. 1959. Cizopasni Cervi nagich well as a record of their field observations, and zoiravoi. Systematicko-faunisticka studie a to Dr. B. Horning, Vet.-Bakteriologisches and ekologicko-zoogeograficke zhodaoceni. desk. Parasitologisches Institut, Universitdt Bern, Parasit. 6: 87-134. Switzerland, who provided the otherwise un- , and V. Mahnert. 1970. Ober Helmin- available description of C. rem. then der Kleinsauger (Insectivora, Rodentia) Tirols Wsterreichs). Ber. Nat.-Med. Ver. Literature Cited Innsbruck 58: 143-154. Rukhliadeva, M. N. 1946. K izucheniiu nema- Arzamasov, I. T., I. V. Aferkusheva, 0. N. tod roda Capillaria Zeder, 18430, of kutory Mikholap, and I. V. Chikilevskaia. 1969. (Neomys fodiens Pall.). In V. P. Podiapol- Nasekomoiadny-e i ikh parazity na territorii skaia (ed.), Germintologicheskii Sbornik, p. Belorussii. Nauka i Tekhnika, Minsk, 175 p. 225-226. Akad. Nauk SSSR, Moscow-Lenin- Franklin, J. F., and C. T. Dyrness. 1969. grad, 302 p. Vegetation of Oregon and Washington. US- Soltys, A. 1954. Helminthofauna of Soricidae in DA Forest Service Res. Paper PNW-80, Port- the Bialowieta National Park. Acta Parasit. land, Oregon, 216 p. Polon. 1: 353-402. Furmaga, S. 1959. Internal parasites of the Wakelin, D. 1968. Nematodes of the genus mole ( europaea L.) in the Lublin en- Caadlaria Zeder, 1800 from the collection of vironment Acta Parasit. Polon. 7: 203-214. the London School of Hygiene and Tropical Johnson, M. L., and B. T. Ostenson. 1959. Medicine. III. Capillariids from mammalian Comments on the nomenclature of some hosts. J. Helminthol. 42: 383-394.

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

W. E. MARTIN Department of Biological Sciences, University of Southern California, Los Angeles, California 90007

Aris-raAcr: 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 (1970-71) spent intestines some trematodes that are unique in the Parasitology Department, University of among haploporids in the possession of an un- Queensland, Brisbane, Australia, Brisbane River divided cecum. They are described as two mullet were found to harbor in their small new genera and three new species of a new 1 Supported by NSF G6962. subfamily.