I (PI I,Cf

•15.1:4=41;;;=•

PROJECT PNW-1204CULTURE OF MIXED CONIFERS

WEST SIDE CASCADES

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

Ph:uornenes de convergence chez les spirur-ides, en particulier clans lee sous-familles Ha-bronematinae Chitwood and Wehr, 1932 etSchistorophorinae L. Travassos, 1918; leurimportance por une classification naturelledes spirurides (Nematoda). Bull. Soc. Zool.Fr. 82: 88-102.

Inglis, W. G. 1965. The nematodes parasiticin the gizzard of birds: A study in morpho-

logical convergence. J. Helminthol. 39: 207-2x4.

Mawson, P. M. 1968. Habronematinae (Nema-toda: Spiruridae) from Australian birds. Par-asitology 58: 745-767.

Yeh, L. S. 1954. On two new species of thegenus Serticeps (Nematoda: Schistorophidae)from the gizzard of birds. J. Helminthol. 28:165-170.

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

R. L. RAUSCH AND V. R. RAZSCHArctic 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 shrews, Sores spp. (typehost: S. yaquinae Jackson) in Oregon, and is recorded also from the shrew-mole, Neiirotrichus gibbsii(Baird). C. maseri is the first capillariid to be reported from the urinary bladder of North Americaninsectivores_

In 1970-72, nematodes of the genus Capil-laria Zeder, 1800, were found in the urinarybladder of insectivores collected in Oregon byChris and Rita Maser, Oregon Coast EcologicalSurvey, Puget Sound Museum of Natural His-tory, who kindly sent the specimens to us forstudy. The species involved differs from anyrecorded from insectivores in Eurasia andNorth America and is described herein.

Materials and MethodsMost of the specimens had been fixed in

situ in 10% formalin solution. Fifty-six were-studied in detail, including some that werebroken in dissecting the anterior ends fromtissue of the urinary bladder. They werecleared by evaporation of a mixture of 2%glycerin in 70% ethanol, after which selectedspecimens were stained in light green SF dis-solved in glycerin. Two bladders with nema-todes in situ were embedded in paraffin, sec-tioned at 0.005 or 0.010 mm, and stained inhematoxylin—eosin, in Mallory's aniline bluecollagen stain, or by the periodic acid-Schiffreaction. For comparison, sections were simi-

larly prepared from the normal urinary bladderof a shrew collected in Alaska. The hosts wereidentified by Mr. Maser. The nomenclature ofshrews is in accordance with the recommenda-tions of Johnson and Ostenson (1959).

ResultsDescription

Numbers of specimens from which therespective measurements were derived areshown in parentheses. All measurements arein millimeters.

Capillaria maseri sp. n.(Figs. 1-8)

Slender nematodes; white in color when pre-served. Bacillary lines present. Mouth simple.Cuticle smooth.

MALE (23 specimens; 8 intact): Length(8) 7.3 to 20.5 (avg 17.1). Maximum width(22) 0.099 to 0.155 (avg 0.123), attained inposterior third of body: Width at cephalicextremity (8) 0.008 to 0.011 (avg 0.010);width at posterior extremity (23) 0.031. to

■

0

r).

6414

00

0.075 mm

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

0.036 (avg 0.032). Esophagus (8) 3.7 to 7.9long (avg 6.5); width at base (19) 0.049 to0.065 (avg 0.056). Length of muscular esoph-agus (4) 0.356 to 0.389 (avg 0.375). Ratioof length of esophagus to length of body (8)1:1.9 to 1:3.1 (ava 1;2.6). Twenty-six to 48esophageal cells observed. Distance from pos-terior end of esophagus to testis (19) 0.356to 1.178 (avg 0.645). Length of spicule (23)0.710 to 1.134 (avg 0.970). Spicule sheathaspinose, with minute folds. Caudal alae notpresent.

FEMALE (33 specimens; 10 intact): Length(10) 7.7 to 26.8 (avg 18.4). Maximum width(32) 0.073 to 0.168 (avg 0.145), attained inposterior third of body (containing uterus).Width at cephalic extremity (7) 0.008 to0.011 (avg 0.010). Width at posterior ex-tremity (33) 0.032 to 0.057 (avg 0.044).Width at level of vulva (30) 0.057 to 0.121(avg 0.092). Esophagus (10) 2.93 to 10.4(avg 7.5) long; width at base (28) 0.049 to0.070 (avg 0.062). Length of muscular esoph-agus (5) 0.378 to 0.470 (avg. 0.434). Thirty-nine to 41 esophageal cells observed. Ratioof length of esophagus to length of body (10)1:1.6 to 1:2.9 (avg 1:2.5). Ner •e ring (5)ca. 0.090-0.115 from anterior end. Distancefrom posterior end of esophagus to vulva (28)0.518 to 1.871 (avg 1.145). Distance fromcephalic extremity to vulva (9) 3.5 to 11.4(avg 9.0). Anus terminal. Egg (178) 0.060to 0.073 by 0.024 to 0.032 (avg 0.065 by0.028); shell smooth; polar plugs well defined.

TYPE itosr: Sorex yaquinae Jackson. Otherhosts S. vagrans Baird, S. bendirii (Merriam),S. trotvbridgii Baird, S. pacificus Coues, andNeiirotrichus gibbsii (Baird).

TYPE LOC.ALITY: Cascade Head Experi- -mental Forest, Lincoln County, Oregon.

HABITAT: Urinary bladder.TYPES: USNM Helm. Coll. No. 72256

holotype (male) and No. 72257 allotype; Nos.72258 and 72259 (paratypes).

DiscussionWith the exception of C. hepatica (Ban-

croft, 1893), which has been reported frommammals of diverse groups, species of Capi/-/aria from mammals are usually host-specific.That certain species have been found in hostsrepresenting two or more related families indi-cates that their occurrence may be influencedby ecologic, as well as phylogenetic, factors.Such is the case with C. maseri, occurring inboth shrews (Soricidae) and moles (Talpidae).Hol.vever, in any host infected, the respectivespecies appear to be organ-specific. In Eurasia,nematodes of the genus Capillaria have beenfound in the urinary bladder of moles andshrews, but none has been reported from thisorgan in North American insectivores. Thetwo species known from insectivores in NorthAmerica, C. rauschi Read, 1949, and C.blarinae Ogren, 1953, were described fromthe small intestine of Sorex cinereus Kerr andthe esophagus of Blarina brevicauda (Say),respectively. Wakelin (1968) suggested thatC. rauschi is possibly identical with C. kutoriRukhliadeva, 1946, described from the stomachof water shrews, Neomys fodiens (Pennant),from the Khopersk Reserve, European Russia.His hypothesis, if valid, would add anotherspecies of the genus Capillaria from mammalsto those few lcnown to be holarctic, of whichnone is from insectivores. It seems improbable,on zoogeographic grounds, that the two areconspecific; unfortunately, a conclusion cannotbe made from the descriptions of C. kutori(Rukhliadeva, 1946; Soltys, 1954). C. kuton'appears to be a common parasite of shrews ineastern Europe (Prokopfe, 1959; Arzamasovet al., 1969), and it was identified by Wakelin(1968) from the intestine of Sorex araneusin England.

ComparisonsA distinctive morphological characteristic of

C. rnaseri is the relatively great distance be-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 ofmale, slightly oblique. 3. Posterior end of niale. 4. Caudal extremity of male, with partially evertedspicule. 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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the mucosa. PAS stain, 0.010-mm section.

(avg 1.145 mm). C. maseri differs from thetwo species previously described from NorthAmerican shrews in other characters as wellas in habitat. From C. rauschi, it is distin-guished by the presence of bacillary lines,smooth cuticle, different position of the vulva,and smooth-shelled eggs of larger size; fromC. blarinae by the larger size, longer esopha-gus, longer spicule, absence of spines in spic-ule sheath, and, in females, by the presence ofan external vulvar process and a terminal analopening.

Several species of Capillaria occurring inthe urinary bladder of insectivores have beendescribed in Eurasia. C. capillaris (vonLinstow, 1882) and C. incrassata (Diesing,1551) have been recorded from both molesand shrews. Wakelin (1068) concluded thatC. capillaris could be synonymized with C.incrassata, a determination that was not ac-cepted by ProkopiZ:; and Mahnert (1970) or

Nadtochii and Rasskazova (1971). Both spe-cies have been found in western Eurasia aswell as in Kamchatka, in northeastern Siberia(Nadtochii and Rasskazova, 1971).

C. urinicola Soltys, 1952, described fromshrews in Poland, was later considered bySoltys (1954) to be identical with C. capillaris.However, comparisons by Furmaga (1959) ledto the conclusion that C. urinicola is distinct.

The similarity between C. sunci Chen, 1937,described from Sun= caeruleus (= S. murinusL.) (Soricidae) in China, and C. incrassatawas recognized by LOpez-Neyra (1947, p.140). Wakelin (1968, p. 392) stated thatthis species ". . . may well be considered asynonym of C. incrassata." •

C. reni Shaldybin, 1968, described fromwater shrews from the Mordovsk Reserve inEuropean Russia, might also be consideredhere, although it was found in the renal pelvisrather than in the urinary bladder.

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

C. maser( may be distinguished as followsfrom these other species inhabiting the uri-nary system of insectivores: from C. capillarisby its larger size, longer esophagus, vulvasituated farther posteriorly, and smooth-shelledegg; from C. incrassata by its larger size, moreslender anterior end, smooth cuticle, and longerspicule; from C. urinicola by its larger size,longer esophagus, presence of vulvar process,and a much larger egg; from C. rent, of simi-lar large size, by its longer spicule and, in thefemale, by its terminal anal opening andsmaller egg. The description of C. real doesnot permit comparison of other anatomicaldetails.

Parasite—host relationshipsC. maseri occurred in 43 (31%) of 138

shrews collected during November 1970—April1972. It was found also in one of seven shrew-moles, Neiirotricims but in none of 16Townsend's moles, Scapanus townsendii (Bach-man), or in 17 coast moles, S. orarius True.In 32 infected shrews, not segregated by spe-cies, numbers of nematodes ranged from 1 to29, with an average of 4. The ratio of malesto females in this series was 1:1.7. Five speci-mens were found in the bladder of the shrew-mole. Findings in mammals of the respectivespecies are summarized in Table 1. Mammalsof both sexes were infected.

The nematodes formed localized aggrega-tions on the floor of the urinary bladder in thevicinity of the neck, with the anterior portionof their bodies deeply embedded in themucosa. Males penetrated to the full lengthof the esophagus, females to the level of thevulva, permitting discharge of eggs directlyinto the lumen of the bladder. In sections ofincised, contracted bladders, in which the mu-cosa was thrown into deep, irregular folds, theanterior portions of the nematodes extendederratically through the thick epithelial layer,but not penetrating the connective tissue ofthe lamina propria, nor did they pass throughthe mucosa into the underlying muscularis(Fig. 9). No observations were made on re-lationships in the distended bladder, in whichthe respective layers of tissue would be muchflattened. Findings in tissue sections indicatethat C. maseri is not pathogenic in shrews. Theembedded portions of the nematodes were im-

Table 1. Occurrence of C. maseri in shrews and

Host No. examined No. infected

Soricidae 138 43Soren vogran., 16 10S. pacificw 27 5S. yaquinae 23 9S. bendini 13 3S. troirbridgii 59 16

Talpidae 40 1Neiirotrichus 7 1Scapanus ararius 17 —S. town.sentiii 16

mediately surrounded by normal-appearingepithelial cells; leukocyte infiltration, fibrosis,or other evidence of an inflammatory responsewere not observed. No eggs were found withinthe tissue of the host. The tissue reaction, ifany, evoked by other species of Capillaria in-habiting the urinary bladder of insectivoresapparently has not been described. A slidewith sections of the nematode in situ has beendeposited in the USNM Helm. Coll., No.72260. No early-stage infections involvingnewly established nematodes were detected,although a few immature specimens werefound. In the shrew-mole, one of the nema-todes was within the ureter; however, migra-tion from the bladder might have taken placeafter the death of the host.

Most of the insectivores were captured alongthe coast of Oregon in the vegetational zonecharacterized by Sitka spruce, a region of lowelevation, mild climate, and high annual pre-cipitation (Franklin and Dyrness, 1969). Ofthe five species of shrews involved, S. trow-bridgii, S. pac-ificus, and S. yaquinae inhabitforest and are nocturnal. S. bendirii is foundnear water, while S. vagrans inhabits meadowsas well as grassy areas along streams; both areactive at any time of day. During the driermonths (summer and autumn), all but S.cagrans were usually trapped within 25 in ofstreams. The shrew-mole has shrewlike habits;it usually occurs in forest and is active at anytime of day. The infected insectivores weretrapped in moist habitat in seral vegetation.C. maser( was found also in a single specimenof S. bendirii from the western foothills of theCascade Range (Lane County), at an elevationof about 1,000 m. (The foregoing data wereprovided by C. Maser.)

Seasonal differences in rates of infection in

112 • PROCEEDINGS OF THE HELMINTHOLOGICAL SOCIETY

shrews could not be discerned. Infected ani-mals were obtained during all months butSeptember and October. The lowest annualrates would be expected in autumn, when thepopulations of shrews consist predominantlyof young animals.

AcknowledgmentsWe express thanks to Mr. and Mrs. Chris

Maser, who provided the material studied aswell as a record of their field observations, andto Dr. B. Horning, Vet.-Bakteriologisches andParasitologisches Institut, Universitdt Bern,Switzerland, who provided the otherwise un-available description of C. rem.

Literature CitedArzamasov, I. T., I. V. Aferkusheva, 0. N.

Mikholap, and I. V. Chikilevskaia. 1969.Nasekomoiadny-e i ikh parazity na territoriiBelorussii. Nauka i Tekhnika, Minsk, 175 p.

Franklin, J. F., and C. T. Dyrness. 1969.Vegetation of Oregon and Washington. US-DA Forest Service Res. Paper PNW-80, Port-land, Oregon, 216 p.

Furmaga, S. 1959. Internal parasites of themole (Talpa europaea L.) in the Lublin en-vironment Acta Parasit. Polon. 7: 203-214.

Johnson, M. L., and B. T. Ostenson. 1959.Comments on the nomenclature of some

mammals of the Pacific Northwest. J. Mam-mal. 40: 571-577.

Lo'pez-Neyra, C. R. 1947. Los Capillarinae.Mem. Real Acad. de Ciencias Ser. CienciasNat., Madrid 12: 1-248.

Nadtochii, E. V., and T. T. Rasskazova.1971. Nematody zemleroek rtekotorykh ter-ritorii DaFnego Vostoka. In E. V. Gvozdev(ed.), Biologicheskie Problerny Severn, p. 93-99. Trudy Severo-vostochnogo KomplekskogoInst., Akad. Nauk SSSR, Magadan, 239 p.

Prokopie, J. 1959. Cizopasni Cervi nagichzoiravoi. Systematicko-faunisticka studie aekologicko-zoogeograficke zhodaoceni. desk.Parasit. 6: 87-134. , and V. Mahnert. 1970. Ober Helmin-

then der Kleinsauger (Insectivora, Rodentia)Tirols Wsterreichs). Ber. Nat.-Med. Ver.Innsbruck 58: 143-154.

Rukhliadeva, M. N. 1946. K izucheniiu nema-tod roda Capillaria Zeder, 18430, of kutory(Neomys fodiens Pall.). In V. P. Pod'iapol'-skaia (ed.), Germintologicheskii Sbornik, p.225-226. Akad. Nauk SSSR, Moscow-Lenin-grad, 302 p.

Soltys, A. 1954. Helminthofauna of Soricidae inthe Bialowieta National Park. Acta Parasit.Polon. 1: 353-402.

Wakelin, D. 1968. Nematodes of the genusCaadlaria Zeder, 1800 from the collection ofthe London School of Hygiene and TropicalMedicine. III. Capillariids from mammalianhosts. J. Helminthol. 42: 383-394.

A New Subfamily, Two New Genera, and Three NewSpecies 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 thenew genus Unisaccus and the new species, U. brisbanensis and U. spinosus; and the new genus andspecies 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) spentin the Parasitology Department, University ofQueensland, Brisbane, Australia, Brisbane Rivermullet were found to harbor in their small

1 Supported by NSF G6962.

intestines some trematodes that are uniqueamong haploporids in the possession of an un-divided cecum. They are described as twonew genera and three new species of a newsubfamily.