COLLABüRATüRS

FLIES LICE E. PAUL CATTS KE CHUNG KIM University of Delaware Pennsylvania State University Newark, Delaware University Park, Pennsylvania

LICE PROTOZOANS KARY C. EMERSON NORMAN D. LEVINE Arlington, Virginia University of Illinois Urbana, Illinois PENTASTOMIDS ALEX FAIN Institute de Médecine Tropicale Prince Léopold ROBERT E. LEWIS Antwerp, Belgium Iowa State University Ames, Iowa NEMATODES, ACANTHOCEPHALANS, LE.ECHES LICE DONALD HEYNEMAN ROGER D. PRICE University of California University of Minnesota San Francisco, California St. Paul, Minnesota

PARASITES OF FISHES GLENN L. HOFFMAN BUGS Eastern Fish Disease Laboratory RAy:yrOND E. RYCK:YL\N Leetown, "Vest Virginia Loma Linda University Loma Linda, California TICKS HARRY HOOGSTRAAL TREMATODES, CESTODES U.S. Naval Medical Research Unit MARIETTA VOGE No. 3 University of California Cairo, U.A.R. Los Angeles, California

PARASITES OF AMPHIBIANS AND REPTILES MITES HAROLD M. KAPL\N CONRAD YUNKER Southern Illinois University Rocky Mountain Laboratory Carbondale, Illinois rr-,~.. > Hamilton, Montana t',,·,;

IiL>t R~'{,",,::<,;, r 1

Chapter 16

PENT ASTOMIDS

by Alex Fain

P ENTASTOMIDS wmtitute a highly Natural infections have been reported aberrant group of (Fain 1961; in sorne endothermal species occasionally Heymons 1935). They have a wormlike used in the laboratory. Experimental in­ and generally annulated appearance. The fections have also been produced, The body is white. legless, and either cylindrical pentastomids that may be encountered in or flat. endothermal laboratory species are listed These arthropods are typically heter­ in Table 16.1. The most important species oxenous parasites. In the most evolved are described below. species, the adults live in the respiratory Linguatula sel'rata tract of carnivorous animaIs, usually , and the larvae develop in the tis­ (Tongue Worm) sues of various animaIs, usually . , a relatively benign The intermediate host becomes in­ parasite, is found throughout the world, fected by drinking water or eating food but its exact incidence is unknown. It is contaminated by fecal material or by commonest in Europe, especially eastern mucus from the respiratory tract of an ani­ Europe (Heymons 1942). and has been re· maI harboring adult pentastomids; the de­ ported from the United States (Stiles 1895), finitive host becomes infected by eating South America (Gelormini and Roveda animaIs or viscera containing nymphs. 1938). South Africa (OrtIepp 1934), Asia The development in the intennediate (Faust 1927; Heymons 1935), the Philip­ host gener:llly takes several months. The pine Islands (Tubangui :lnd :\lasiluilgan development:ll stages include a prim:lry 19:16), ,-\ustralia (Pullar 1936), and ~ ew or migr:lting bna; a secondary or resting Ze:lland (Gurr 1953). larva which molts sever:ll times; and a ter­ .-\dults accur in the nasal passages of tiary larva or nymph which encysts in the the dog and other canids, and rarely in tissues of the host, generally in the peri­ damestic farm species :lncl man (Heymons toneal cavity. The terti:lry larva usuaUy 1942). The nymph sometimes occurs in the remains encysted until ingested by the wild ~orway rat, black rat, guinea pig, definitive hast. but sometimes. for reasons rabbit, cat, titi monkey, gelada baboon, not dearly understood. it escapes from its and man (Bochefontaine 1876; Heymons cystic envelope and migrates through the 1942; Kuntz, Myers. and Vice 1967; Strong. tissues of the intermediate host, causing Shattuck, and Wheeler 1926). lt is com­ acute peritonitis and possibly death (Chal­ mon in domestic farm species. A report mers 1899). of canine nymphal linguaculosis in Japan , 493 ~ >, l z 1 494 PARASITES OF ENDOTHERMAL LABORATORY

is of doubtful validity (Yamashita and Ohbayashi 1954). The nymph was proh­ abl)' ATmillifer moniliformis. The incidence of L. sen-ata in labora­ tory specimens is unknown. It is not likely ~ ,...~ to be encountered except in dogs obtained "'" ~. from pounds or in rodents obtained from their natural habitat. The mouse, guinea pig, and rock squirrel (Otospermophilus beecheyi) have been infected experimen­ . tally (Heymons 194~; Hobmaier and Hob· maier 1940; Koch 190i). /' 1"'..0l.. MORPHOLOGY , The adults have a transparent, tongue­ shaped body with approximately 90 annuli (Fig. 16.1) (Sambon 1922). The anterior end has two pairs of simpIe retractile (,1' hooks. The female is 80 to 130 mm long \~ and 10 mm wide, and reddish orange eggs ...o!!!!!I!D ,') "­ are visible along the median line of the ~ '\'.., \.. ~ .....'" - 5mm FIG. 16.2. Linguatula serrata nymphs. (Cour. tesy of A. Fain, Institut de Médecine Tropi· cale Prince Léopold.)

body; the male is 20 mm long and 3 to 4 mm wide. The nymph, sometimes called Pentastomum denticulatum, is 4 to 6 mm long and 1 mm wide (Fig. 16.2). It has spinous body rings and two pairs of binate hooks (Fig. 16.3). The egg is oval, about iO to 90 p. in diameter, and is individually enclosed in a thin bladderlike envelope con­ taining a clear fluid. It has a thick chitin­ ous shell containing an embryo with rudi­ mentary mouthparts and four short legs, each bearing two clawlike hooks. On the back of the em brvo is the so-called dorsal organ or facette. ­

LIFF. CYCLE Eggs are expelled from the definitive host in the nasal mucus or are swallowed and passed in the feces (Hobmaier and Hobmaier 1940). When ingested by an in­ termediate host, thev hatch in the intes­ tine. The resultant hrvae migrate to the internaI organs, usually the mesenteric FIG. 16. J. Linguatula serrata. (Le!t) Male. lymph nodes, and after about 6 months (Righi) Female. (Courtesy of A. Fain, Institut and nine molts, they develop into infective de Médecine Tropicale Prince Léopold.) nymphs. These nymphs remain viable in PENTASTOMIDS 495

ined for this parasite. Infected dogs can be treated by spraying the nasal passages with an aerosol containing ascaridol (the active ingredient in chenopodium oil) (Enigk and Düwel 1957), or the parasites can be removed surgically (Olt and Strôse 1914). There is no treatment for nymphal infections.

PUBLIC HEALTH CONSIDERATIONS This parasite is not an important pub­ lic health problem (Fain 1960). The nymph and, very rarely, the adults have been re­ FIG. 16.3. Linguatula senala nymph. Note ported in man. spi nous body rings and the two pairs of binate hooks. (Courtesy of A. Fain, Institut de Méde· Porocephalus cine Tropicale Prince Léopold.) Porocephalus crotali occurs in North and South America, P. clavatlls occurs in the intermediate hast for over 2 vears. The South America, and P. subulifer is con· definitive hast becomes infected' by ingest­ fined ta tropical Africa (Fain 1961, 1966; ing viscera containing the infective stage, Heymans 1935). Porocephalus nymphs are but the method by which nymphs get ta occasionally found in the viscera of sorne the nasal cavities is unknown. This may endothermal laboratary species. They are occur while contaminated food is being relatively benign parasites. Adults usually masticated or possib!y later during emesis. live in the lung of large snakes. Adults survive about 2 years in the defini· The P. crotali nymph has been found tive hast. Thev feed on nasal mucus and in deer mice and the cotton rat in the secretions and ~ccasionally on blood (Hey· Vnited States (Layne 1967; Self and Mc­ mans 1942). Murry 1948), and there is a doubtful rec­ ord in a marmoset (Sagllinus) (Heymons PATHOLOCIC EFFECTS 1935). The nymph of P. clavatus has been Vsually there are no signs of infec­ reported from the common marmoset (Hey­ tion, but a severe catarrhal or suppurative mons 1935), from laboratory tam:lfins rhinitis and epistaxis sometimes occur (Saguin1ls nigricollis) in the United States (Enigk and Düwel 1957; Heymans 1942). (Cosgrove, Nelson, and Gengozian 1968; Restlessness, sneezing, and difficult breath­ Nelson, Cosgrove, and Gengozian 1966), ing are occasionally seen. The sense of and, erroneously, from African primates smell is ohen reduced or abolished. and man (Fiennes 1967). The nymph of The nymph generally does not pro­ P. subulifer has been found in a guenon duce signs and is an inciden tal necropsy (Heymons 1935) and in a ~a1ago (Fain finding, appearing as a small fibrous or 1961), and nymphs of an unidentified spe­ calcified tuberde in the viscera. A massive cies of Porocephallis have been recO\'ered infection has camed peritonitis in an ex­ from a squirre1 monkey (A. Fain, unpub­ perimentally infected guinea pig (Koch lished data). 1907). The rat, mouse, and hamster have been experimentally infected with eggs of DJAC;-.lOSIS P. crotali (Esslinger 1962a, b), ancl the Diagnosis is based on clinical signs and mouse, rat, and guinea pig have been ex­ the presence of L. serrata eggs in the feces perimentally infected with eggs of P. cla­ or nasal mucus. vatus (da Fonseca 1939). Porocephalus cla

496 PARASITES OF ENDOTHERMAL LABORATORY ANIMALS

1966; Heymons 1935; Penn 1942; Self and McMurry 1948).

PATHOLOCIC EFFECTS In deer mice and the cotton rat, the nymph locates in the viscera, mesentery, and abdominal and thoracic walls (Layne 1967). It produces no signs or serious pa· thology. In primates, the nymph encysts in many tissues (Fig. 16.5. 16.6), including the liver. lungs. peritoneum. and meninges, but produces little or no injury (A. Fain. unpublished data; Nelson, Cosgrove, and Gengozian 1966). Inflammatory reaction is minimal (Fig. 16.7) unless the nymph dies; then a foreign body reaction and graduaI resorption occur (Fig. 16.8).

FIc. 16.4. Porocephalus nymph. (Courtesy of DIAGNOSIS A. Fain, Institut de Médecine Tropicale Prince Léopold.) Since This parasite usually does not produce signs or lesions, diagnosis is made by finding the nymph at necrops)'. Cosgrove 1968). The incidence of P. crotali and P. subulifer in laboratory specimens is CONTROL unknown. They are not likely to be en­ No special procedures other than rou­ countered except in sorne primates and tine sanitation are necessan. The Poro­ wild rodents obtained from endemic areas. cephalus nymph can occur only in labora· tory species permined to ingest food con­ I\fORPHOLOCY taminated with feces of infected snakes. The Porocephalus nymph has a cylin· There is no treatment. drical, smoothly annulated body which is often club shaped (Fig. 16.4) (Heymons PUBLIC HEALTH CONSIDERATIONS 1935). Nymphs of P. crotali and P. da· POl'Ocephalus is of no known public vatus are about 8 to 14 mm long; the health importance. AU reports of the nymph of P. subulifer is 7 to 15 mm long nymph in man are of doubtful \'alidity and 1.2 to 1.5 mm wide. Each nymph has (Fain 1960). approximately 30 to 45 annuli. Two un· equal pa~rs of hooks are located at the Arrnillifer ar711illatlls anterior end around the mouth. The inner This relatively benign parasite occurs pair is simple; each of the outer hooks naturally only in tropical A[rica, where it has an accessory spine. The adults are is common. The nymphs develop in var· similar, only larger. ious endothermal species and are fre­ quently found in the rhesus monkey, other LlFE CYCLE macaques, galagos, guenons, mangabeys, The life cvcle resembles that of Lin­ baboons, the chimpanzee, dog, and man guatula sen'ata' except that the adults oc· (Fain 1960, 1961, 1966; Fiennes 1967; Hey. cur in snakes instead of in the dog and mons 1935). There is a report of the oc­ other canids. Adults of P. crotali are corn· currence of the nymphs in a New World mon in rattlesnakes (Crotalus) and also monkey, a capuchin, from a European zoo occur in the cottonmouth water moccasin (Desportes and Roth 1943), but This infec­ (Aneistrodon piscivorus); adults of P. cla­ tion was probably acquired in the zoo. vatus occur in boas (Boa, Epicrates and Adults of A"millifer armillatus live in the Eunectes); and those of P. stlbulifer occur lung of large snak·es. only in file snakes (Mehelya) (Fain 1961, The incidence of A. arrnillatus in lab­ FIG. 16.5. Porocephalus clavatus nymphs (arrows) in the liver and lungs of a tamarin (Saguinus nigricollis). (From Nelson, Cosgrove, and Gengozian 1966. Courtesy of American Association for Laboratory Science.)

FIG. 16.6. Porocephalus nymph on the liver FIG. 16.7. Histologie section of encysted Poro­ of a squirrel monkey. (Courtesy of A. Fain, In­ cephalus clavatus nymph in the liver of a tama­ Stitut de Médecine Tropicale Prince Léopold.) rin. (A) Liver tissue. (B) Parasite. Note ab­ sence of inflammation. (From Nelson, Cos­ grove, and Gengozian 1966. Courtesy of Ameri· can Association for Laboratory Animal Science.) i "'""------­ .J 498 PARASITES OF ENDOTHERMAL LABOR\\TORY ANIMALS

FIG~ 16.9. Armillife,' annillatus nymphs. (From Fain 1961. Courtes)' of Musée royal de l'Afrique Centrale.)

1961). Often they are located beneath the capsule or are embedded in the superficial layers of the liver. They usually cause lit­ tle or no reaction in the host even at high levels of infection, but there is one report of peritonitis and death caused by Armilli­ fcr nymphs in 7 of 24 laboratory manga­ beys (Whitney and Kruckenberg 1967). FIG. 16.8. Histologie section of the liver of a DIAGNOSIS tamarin containing degenerating Porocephalus Diagnosis is based on finding nymphs clavatus nymph. Note inflammatory reaction. at necropsy. (From Nelson, Cosgro\'e. and Gengozian 1966. Counes)' of American Association for Labora· tory Animal Science.) '-' --:9 oratory specimens is unknown. It is nOl .,~. '~.~,: ~~ likely to be encountered except in primates :-<;~~,.~ ,~':.., obtained from tropical Africa. 0':;" ~.... 0 . ·r~._ '_ "i . '. r., .' ~.,',. MORPHOLOGY #.~.',':';~:.t'~.~~' The nymphs have a cylindrical, an­ • nulated body about 13 to 23 mm long ...... ' ~_.~,j. \, (Fig. 16.9) (Fain 1961). The annuli are :~.9~ ... ,.: .. Çf~Q.ilI thick and projecting, and their number .}~~ .,~,'.' ;~U varies from 15 to 19 in the male nymph . and from 18 to 22 in the female nymph. ;~.:. ~ ~ .~~ .e~~~~èl l~ The hooks are simple. Adults closely re· •. semble the nymphs but are larger. ":' "~"""--A ....~.-..Î...... '.' ~.;::.:. .~...... -.;: :. """-.--: ., -. nJ> LIFE CYCLE ..:>...... , 'ë). ~;.:;:'.:. ....}~.',.~~f' '~.',2\ .., •... "".·..*_':V::J....~~.... The adults live in the lung of large _L ro_ . .'._. ". .:sI";; """, \~~_•• ? . v)' ·C:.':,,,, .•c. ... African snakes (Python, Bitis) (Broden and ~,,-~,'- - ,;::~-- -~-~; "'4" ~ Rodhain 1907, 1908-1909, 1910). The in· ~ ~ (,,:' •... ;-: .. , ..... termediate host becomes infected by swal­ .'..:...• ~.~. -::; lowing food contaminated by feces 'CW" or saliva. FIG. 16.10. nymphs en· PATHOLOGIC EFFECTS cysted in the omentum of a green monkey. Encysted nymphs are commonly found (From Fain 1961. Counesy of Musée royal de in the peritoneal cavity (Fig. 16.10) (Fain l'Afrique Centrale.) r PENTASTOMIDS 499 CONTROL PUBLIC HEAI.TH CONSIDERATIONS In sorne parts of tropical Africa man Other than routine sanitation, no spe­ is frequently infected (Bouckaert and Fain cial control procedures are necessary. In· 1959; Cannon 1942), but the parasite in fection can only occur if laboratory spe­ endothermal laboratory species is of no cies are permitted ta ingest eggs passed public health importance. Man can only in the feces or saliva of infected snakes. be infected by ingesting eggs passed in There is no treatment. snake feces or saliva. TAnI.f: HU. Prnta~nomld~ afff"rliuR: rndothrrll1al lahoralory aoi01al~

Public GcoRnphic F.ndolhermal Location MClhod of Indd("rlce l'af hologic IlraIth It.. r:uilc Di~(Tihution Ho.l in HO~I InFrction ln nature In I:Iboral;;-ry­ Hrcel. Importance Refcrrncc

1_;"Kllatu/a Worldwidc J)rfinitivc: Adllh: J)rriniti\'c: ~·;OInnl(JII in Untenown: lJ~;ual1y 11011<"; Rare in RochefonlaÎl1e Jerrata· dog,olher Il;)''al C3\'Îty ingc"lioll uf

• J)hrll!ll"c(\ in trxt.

"IJIICJ" ,\p' '1 TAilLE Ili.1 (amlin'U'll)

Public Geographie t:IU..ltll hc.:rmal I..oL~liun l\Iethod al Incidence Ilathologic lIeallh I)arasilc Disuihulioll IJu.!It ill nust JnfCClioli -In n;llurc 1n laborawry Elfect. Importance Reference -~-_.._.­ --­ Gig/iul.l1a Madagascar JIIlcnucdhuc: ~Ic.st.'nl('ry Uuknown; pre­ Unkno\'f'1l Uukno,... n IkniM,u q:~ts Noue Chabaud ~lId b'lIInpti ICUlur ;'1.)(: .umahly by in mcscntcry Chu,!uet 1954 (CI,,:i,o­ Îllgc:u;on of galt:lIs cggs p3sscll lJy tIItNtill~J dcfini.ivc laos(

Ar//lillif<1 'l'rOllilal IlltcruH.:dii1(C; h:..-iloucal JuKcslÏoli of eggs Common Unknown USIll.l)ly henil(u Commun in J)c:sportes allJ unnillutus· Africa dug, c.:avity [,aucd by cySl!I in man in Roll. 1945 mOllkcy. dcfinilivc host peritoneal endemic Fain 1960, 1961 mlu:r C3VÏly; rarcly arcas; Fienne. 1967

rl '//Iil/ifcr As;a. IUICllllcdialC: Viscera. lugcslion oC cggs Unknowil Unknowli l1nkuowli R.ue in hin 1966 JUcmili/urmiJ Auslr<.llia vadous pcritolleal pa..ed by man Worm.1967 mammuls. t:avity dcfinilive host Va,na,hita and pm~sjhly (pythons) Ohbaya.hi 1954 dog, cal. cynornolgus moule)', man

• UiM:us)cd in teXI.

------::--­,...- ...... ~.=.=._=====;:==;;;:;;;;;;;;;;;;; 502 PARASITES OF ENDOTHERMAL LABORATORY ANIMALS REFERENCES other hosts in China. Am. J. Trop. Med. 7:311-25. Bochefontaine, M. 1876. Pentastome denti­ Fiennes, R. 1967. Zoonoses oE primates: The culé provenant du poumon d'un cobaye. epidemiology and ecology oE simian dis­ Compt. Rend. Mem. Soc. Biol. Paris 28:261. eases in relation to man. WeidenEeld and Bouckaert, Lo, and A. Fain. 1959. Een geval Nicolson, London. 190 pp. van nymphale porocephalose met dodelijk Fonseca, F. da. 1939. Observaçoes sç>bre 0 cielo verloop. Ann. Soc. Belge Med. Trop. 39: evolutivo de Porocephalus clavatus, espe­ 793-98. cialmente sobre 0 seu orquidotropismo em Broden, A., and J. Rodhain. 1907. Contribu­ cobaias. Mem. InsL Butaman (Sao Paulo) tion à l'étude de Porocephalus monili­ 12:185-90. formis. Ann. Trop. Med. Parasitol. 1907: Gelonnini, N., and R. J. Roveda. 1938. Lin­ 493-504. guatuZa seTTata. Inst. Parasitol. Enferm. --. 1908-1909. Contribution à l'étude de Parasit. Uni\'. Buenos Aires 1:3-12. porocephalus moniliformis. Ann. Trop. Gurr, L. 1953. A note on the occurrence oE Med. ParasitoI. 1908-1909: 303-13. Linguatula serrata (Frohlich, 1789) in the --. 1910. Contribution à l'étude de Poro­ wild rabbit, Oryctolagus CUlljeulus, in New cephalus moniliformis. Ann. Trop. Med. Zealand. New Zealand J. Sei. Technol. 35: ParasitaI. 1910:167-76. 49-50. Cannon, D. A. 1942. Linguatulid Heymons, R. 1935. , pp. 1-268. of man. Ann. Trop. Merl. Parasitol. 36: In H. G. Bronn's h.lassen und Ordnungen 160-66. des Tierreichs. Vol. 5, Sect. 4, Bk. 1. Aka­ Chabaud, A. G., and Marie-Therese Choquet.. demische VerlagsgesellschaEt M.B.H., Leip­ 1954. Nymphes du Pentastome Gigliolella zig. (n. gen.) brumpti (Giglioli 1922) chez un Lemurien. Riv. Parassitologia 15:331-36. 1942. Der Kasenwunn des Hundes (Linguatula !errata Froelich), seine \Vine Chalmers, A. J. 1899. A case of Pentastoma constrictum. Zentr. Bakteriol. Parasitenk. und Beziechungen zur europaischen Tier­ 26:518. welt, seine Herkunft und praktische be­ Cosgrove, G. E., B. Nelson, and No Gengozian. deutung auf Grund unserer bisherigen 1968. Helminth parasites ot the tamarin, Kennmisse. Z. Parasitenk. 12:607-38. Saguinus tuscicollis. Lab. Animal Care Hobmaier, A., and M. Hobmaier. 1940. On the 18:654-56. liEe-cycle of Linguatula rhinaria. Am. J. Desportes, C., and P. Roth. 1943. Helminthes Trop. Med. 20:199-210. récoltés an cours d'autopsies pratiquées sur Koch, M. 1907. Zur h.ennmis des Parasitismus différents mammifères morts a la ménagerie der Pentastomen. VerhandI. Deut. Ges. du Museum de Paris. Bull. Museum Hist. Pathol. 17:265-79. Nat. Paris 15: 108-14. Kuntz, R. E., Betty J. Myers, and T. E. Vice. Enigk, K., and D. DüweI. 1957. Feststellung 1967. Intestinal protozoans and parasites oE und Behandlung des Linguatula-Befalles the gelada baboon (Theropithecus gelada beim Hund. Deut. Tieraerzù. Wocilschr. Rüppel, 1835). Proc. Helminthol. Soc. 64:401-3. Wash. D.C. 34:65-66. Esslinger, J. H. 1962a. Development of PorD­ Layne, J. N. 1967. Incidence of Porocephalus cephalus crotali (Humboldt, 1808) (Pen­ crotali (Pentastomida) in Florida mammals. tastomida) in experimental hosts. J. Para­ Bull. Wildlife Dis. Assoc. 3: 105-9. sitaI. 48: 452-56. Nelson, B., G. E. Cosgrove, and N. Gengozian. --. 1962b. Hepatie lesions in rats experi­ 1966. Diseases of an imported primate memally infeeted with Porocephalus cTOtali Tamarinus nigricollis. Lab. Animal Care (Pentastomida). J. ParasitaI. 48:631-38. 16:255-75. Fain, A. 1960. La pentastomose chez l'homme. Olt, A. and A. Strôse. 1914. Die Wildkrank­ Bull. Acad. Roy. Med. Belg., Ser. VI, 25: heiten und ihre Bekampfung. J. Neumann, 516-32. Neudamm. 649 pp. --. 1961. Les pentastamidés de l'Afrique Ortlepp, R. J. 1934. Note on the occurrence Centrale. Ann. Musee Roy. Afrique Cen­ oE the tongue-worm, Linguatula seTrata, in trale, Ser. 8, Sei. Zool. 92: 1-115. a dog in South AErica. J. S. AErican Vet. 1966. Pentastomida of snakes-their Med. Assoc. 5:113-14. parasitological role in man and animaIs. Penn, G. H., Jr. 1942. The life history of Po­ Mem. Inst. Butaman (Sao Paulo) 33:167­ TOcephalus cTOtali, a parasite of the Louisi­ 74. ana muskrat. J. Parasitol. 28:277-83. Faust, E. C. 1927. Linguatulids from man and Pullar, E. M. 1936. A note on the occurrence of PENTASTOMIDS 5°3 Linguatula seT1'ata (Frohlich, 1789) in Aus­ of the Hamilton Rice Expedition to the naUa. Australian Veto J. 12:61-64. Amazon. Harvard Univ. Press, Cambridge, Sambon, L. W. 1922. A synopsis of the farnily Mass. Linguatulidae. J. Trop. Med. Hyg. 25: Tubangui, M. A., and Victoria A. Masiluiigan. 188-206, 391-428. 1936. Notes on Philippine linguatulids Self, J. T., and G. E. Cosgrove. 1968. Penta­ (Arthropoda: Pemastomida). Philippine stome larvae in laboratory primates. J. J. Sei. 60:399-403. ParasitoI. 54:969. Whitney, R. A., Jr., and S. M. Kruckenberg. Self, J. T., and F. B. McMurry. 1948. Poro­ 1967. Pentastomid infection associated cephalus crotali Humboldt (Pentastomida) with peritonitis in mangabey monk.eys. J. in Oklahoma. J. Parasitol. 34:21-23. Am. Veto Med. Assoc. 151:907-8. Sti1es, C. W. 1895. On the recent occurrence Worms, M. J. 1967. Parasites in newly im­ of Linguatula rhinaria, Railliet, 1886, and ported animaIs. J. Inst. Animal Tech. 18: Taenia echinococcus, V. Siebold, 1853, in 39-47. the United States of North America. Trans. Yamashita, J., and M. Ohbayashi. 1954. On a Fint Pan-Am. Med. Congr. Wash., D.C. tongue worm, Linguatula SeT1'ata Frolich, (Part 2): Il63-65. 1789, showing the sarne parasitism as in the Strong, R. P., G. C. Shattuck, and R. E. Whee­ intermediate hast within a dog body (in 1er. 1926. Other~ parasitic infections of Japanese). Mem. Fac. Agr. Hokkaido Univ. animaIs, pp. Il8-47. In ~Jedical Report 2:146-48.