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Date Gb I (( 10 7 J. Plll'QsllOI"PaY'Osilol .. 87(1),87(1),2001.2001, p. 169-li2169-172 o@ AmericanAmerlcan Society of PurnsiioiogisisParasitologist. 2001

PROTOSPIRURA KAINDIENSIS N. SP. (SPIRURA: SPIRURIDAE) AND OTHER HELMINTHS FROM PSEUDOHYDROMYS (MURIDAE: HYDROMYINAE) FROM PAPUA NEW GUINEA

L. R,R. Smales School of Biological and Environmental Sciences.Sciences, Central Queensland University, Rockhampton, Queensland 4702, Australia

ABSTRACT: One cestode and 3 species ofnematodes are recorded from Pseudohydromys murinus and Pseudohydromys occiden· taUstalis (Muridae: Hydromyinae), from Papua New Guinea, for the first time. Heterakis fieldingijieldingi (Ascaridida: Heterakidea) has previously been known from Australia. Odilia sp,sp. resembles Odilia praeputialispraepulialis in the orientation of the synlophesyntophe and the number and size of ridges but differs in the length of spicule and lack of a praepuce,praepuce. Protospirura kaindiensis n. sp,sp. (Spirurida: Spiruridae) is readily distinguished from all other members of the genus by the number and arrangement of caudal papillae and the length of the spicules.

Although significant collections of New Guinea fauna have daundauo Province. The tissues of this latter animal were hard.hard, brittle, and difficult [0(0 dissect. been amassed by natural history museums over the past 140 yr, The 3 digestive tracts were examined with the aid of a dissecting and comprehensive surveys of the manunalianmammalian fauna began microscope, and allaU helminths found were removed for identification,identification. with the Archbold Expeditions of the 1920s to 1950s (Flannery, Nematodes were cleared in lactophenol prior to examination. Figures 1995), there have been no systematic attempts to document the were drawn with the aid of a drawing tube and measurements, in mil­ helminth parasites of the mammals. There are no published re­ limeters unless otherwise stated, taken using an ocular micrometer, Specimens have been deposited in the United States National Museum cords of gastrointestinal helminths from the rodents. Helminthological Collection.Collection, U.S,U.S. Dept.Dept, of Agriculture, Beltsville, The rodent fauna of New Guinea comprises only represen­ Maryland (USNM) and the BBM. tatives of the family Muridae. It isabundant and diverse, being one of the most species-rich, for the size of the landmass, in RESULTS the world (Flannery, 1995). The Hydromyini, 1 of 4 subgroups Cestode fragments were found in the small intestine of 1 P,P. of the Hydromyinae of Australia and New Guinea, are a mono­ murinus. 1\voTwo male Heterakis fieldingi Smales, 1996 (Ascari­ phyletic group, thought to have originated in montane New dida: Heterakidae) were found in the small intestine of P. mu­ Guinea.Guinea, with 9 genera, 8 endemic, including Pseudohydromys Tim~8.rinu.s, Measurements as follows: length, 4.7, 4.9; width 0.20, Rummler, 1934 (Flannery, 1995). Little is mown about the bi­ 0.27; esophagus length 512, 516 j.lJ11;j.lJll; esophagus bulb width 87.87, ology and habits of the eastern and western shrew-mouse, Pseu­ 86 !J.lll;/lorn; excretory pore 297, 313 fLJIlj.lJll from anterior end; tail 230, dohydromys murinus Rummler, 1934 and pseudohydromys oc­oc' 200 ~m/lorn long; spiCUles 250, 290 /lorn~m long; preanal sucker di­ cidentalis Tate, 1951. They are found in montane forest habitats ameter 40 IJ.rn.~m. This is a new host record. at altitudes between 2,100 and 3,600 m and may be predomi­ Seven nematodes from P. murinus (2 males, 5 females) Odi­ nantly insectivorous (Flannery, 1995). lia sp.sp, (Trichostrongyloidea: Heligmonellidae) were found in As part of an ongoing study on the helminth fauna of the the small intestine. Measurements as follows: rodents of New Guinea, all the available murids, stored as Males: Length 1.5, 1.8 width arat midbody 66, 63 /lorn;~m; ce­ whole specimens in 70% ethanol, in the BerniceBemice Bishop Mu­ phalic vesicle 34, 39 /lom~m long by 23, 22 ~m/lorn wide; esophagus seum, Honolulu, Hawaii (BBM) and the Australian Museum, 280, 300 f1IIllong;j.lJlllong; spicules 190, 180 fJomfJ-m long, Sydney were examined for gastrointestinal helminths. This ma­ma· Females: Length 1.8-2.2, width at midbody 59-74 /lorn;""m; ce­ terial included 6 specimens of Pseudohydromys. Examination phalic vesicle 34-50 ~mv-m long, 20-22 ""m/lorn wide; esophaguseso~hagus of these individuals revealed 3 species of nematode, 1 new to 260-320 long; vagiuavagina vera 30 ""m/lorn long; vulva to postenor end science is described in this paper, and new hosts records for the 84-96 fMU;j.lJll; tail 26-37 iJomjJ.m long; short, conical; eggs thin others are given below. shelled, elipsoidal 65 X 30 fJom.~m. This Odilia sp. differs from allaU There were no records of parasite material from Pseudohy­ the known species of OdilJaOdilia in spicule length and, or the numnum­w dromys in the national parasite collections of Australia, France, ber of ridges in the synlophe at the midbody region. Further the United Kingdom or the United States of America. study on additional specimens is needed to prepare a complete description. ...,., MATERIALS AND METHODS One larval nematode from P. occrdentaltsoccldentallS and 3 mtactIntact nem- males, female), pieces of fe~ales Complete digestive tracts of individuals of P. murinus and P.P, occi­oeci­ atodes (2 males, 1L female), pieces of 6 males and 4 females dentalis~entali!i were carefully removed from whole bodie~bodies that ~adhad been storedstore~ from P.p, murinus,murinus. Protospirura kaindienensis n.n, sp,sp. (Spiruro­(Sprruro­ Inin 70% ethanol. Of the 6 specimens of P. murinusmurmus Inin the museummuseu idea, Spirurida) were found in the stomach. Collection only 2 individuals, registration numbers 56592 and 56596, colte.ct~? had been preserved with digestive systems in situ. Both were colle,ete? DESCRIPTION by P. Woolley on 1] December 1982 and 6 December 1982 at Mt. Kamdl,Kalll I, elevation 2,2502250 m r217°21'S'S 146°41146<:041 'E in the Wau area of Moro~Morobe Pr?v­ protosplrura kaindienslskalndlensls n. sp. . '"" f P 'd t I'l's reoistratlOn (Figs. 1-10) !nee.Ince, Papua New Guinea. One specimen 0of P. • ocel enentaa IS,l, re'''stratlonlr.,. number 105241,105241 was collected by T. BukarnBukam on 8 ~u~ustAugust 197619?6 at Bok­Bok~ I'l' S iruroidea: Spiruridae.Spiruridac:, Large stout wonns,wonns. Cuticle thick, Ubet, elevation 2,850 m, 10 km east of Feramin 55°1313'8,S, 141141°14E,l4E. SanSan- Genera. P stn'ations Anterior extremity with highly developed with transverse .,.. . II dolabia raised above mouth openrng, Mouth opeOlngopemng dorsove~tra y pseu d nded by 2 larger lateral and 4 smallersmalIer submediansubmedIan el- elongateelongate,, surrousurreu b d' h I' '11 Received 6 April 2000; revised 9 August 2000; accepted 9 August . h J·thl'th 2 denticlesdenticles. Four large susubmeme Ian cep a lCIe papl ae,ae. Received 6 April 2000; revised 9 August 2000; accepted 9 August evatlOnS,c:vatlOnS, eac W • 2000.

169 170170 THETHE JOURNALJOURNALOFOF PARASITOLOGY,PARASITOLOGY, VOL.VOL, B7.B7, NO.1,NO, 1, FEBRUARYFEBRUARY 20012001

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2 subdorsal and 2 subventral papillae in cuticular depressions and 2 2 subdorsal and 2 subventral papillae in cuticular depressions and 2 DISCUSSION amphidsamphids formingforming outerouter circle.circle. Small labiallabial papillae not seen. Pharynx DISCUSSION thick-walled, laterally compressed. Esophagus divided into anterior thick-walled, laterally compressed. Esophagus divided into anterior Flannery (1995) suggested thatthat therethere have been multiplemultiple in­in­ muscularmuscular andand posteriorposterior glandularglandular portions. Nerve ring surrounds POSM vasions of the Australia-New Guinea region by murids, at least teriorterior ofofmuscularmuscular portionportion of esophagus,esophagus, excretory pore slightly poste. vasions of the Australia-New Guinea region by murids, at least riorrior toto nervenerve ring.ring. Deirids small,small, anterior toto nerve ring, Phasmids sub­ 1 being the arrival of thethe ancestral Hydromyinae inin thethe latelate terminal.terminal. Miocene, early Pliocene (about 5 million yr ago).ago). MovementMovement MaleMale (holotype(holotype andand paratype):paratype): Length 23, 20. width at midbody 0.48, of these now endemic rodents between Australia andand NewNew 0.48.0.48. AnteriorAnterior endend 0.16,0.16, 0.180.18 diameter. Pharynx 0.08, 0,07 long, Mus­ Guinea is thought to have been inin both directions andand cancan bebe cularcular portionportion ofofesophagusesophagus 0.30, 0,26 long;long; glandular portion of esoph­ agusagus 4.56,4.56, 4,764,76 long.long. Deirids 0.26, 0.22; nerve ring 0.40, 0,39; excretory related to the emergence of landland bridges across thethe TOITesTOITes StraitStrait porepore 0.47,0.47, 0.470.47 fromfrom anterioranterior extremity,extremity, Caudal alae thick. Posterior linking 2 land masses inin thethe Pleistocene era,era, Although Pseu­Pseu­ ventralventral surfacesurface ornamentedornamented with numerous longitudinallongitudinal striae preanally; dohydromys is 1 of the genera restrictedrestricted toto New Guinea (FIan.(FIan. irregularirregular andand transversetransverse patternspatterns postanally, Spicules markedly dissim­ nery, 1995), the helminth fauna itit harbors isis indicativeindicative ofofsuchsuch ilar,left spicule slender, 0.31, 0.33 long; right spicule stout, alate, 0.48, ilar,left spicule slender, 0.31, 0.33 long; right spicule stout, alate, 0.48, migrations. 0.450.45 long,long, GubernaculumGubernaculum 0.90 long.long. Nine pairs, 1 unpaired caudal pa­ pillae;pillae; 44 pairspairs preanal,preanal, II median on anterior anal lip, 5 pairs postanal Heterakis fieldingi has previously been recordedrecorded onlyonly fromfrom 11 papillae,papillae, secondsecond pairpair largest.largest. Tail conical 0.30, 0.33 long with blunt tip. of the 2 hydrornyine species thatthat occur inin both Australia andand FemaleFemale (holotype):(holotype): Length 40. width at midbody 0.73. Anterior end New Guinea, Hydromys chrysogaster Geoffroy, 18041804 andand onlyonly 0.250.25 diameter.diameter. PharynxPharynx 0.830.83long. Muscular portion of esophagus 0.24 in northern Australia. Odilia sp. were thoughtthought toto havehave evolvedevolved long;long; glandularglandular portionportion of esophagusesophagus 4.6 long.long. Deirids 0.31, nerve ring in isolation in Australia from nematodes resemblingresembling Nippo­Nippo­ 0.36,0.36, excretoryexcretory porepore 0.550.55 fromfrom anterior extremity. Vulva without or­ namentation,namentation, 99fromfrom anterioranteriorextremity.extremity. Vagina directed posteriorly. Tail strongylus Lane, 1923 a genus thatthat occurs conunonlyconunonly inin southsouth conicalconical 0.440.44 long,long, Eggs elliptical,elliptical, embryonated, 59.5-62.5 X 33-36 east Asia (Durette-Desset, 1985; Beveridge andand Durette-Desset, ~m,~m, 1992). The discovery of Odilia mallomyos Hasegawa andand Syaf­Syaf­ Larva:Larva: LengthLength 2.8,2.8, width atat midbody 0.16. Anterior end 0.07 di­ ruddin, 1994 in Mallamys Thomas, 1898 fromfrom IrianIrian JayaJaya andand ameter,ameter, EsophagusEsophagus 1.251.25 long,long, tailtail 0.15 long.long. now a second species of Odilia inin Pseudohydromys. bothboth hostshosts TaxonomicTaxonomic summarysummary endemic to New Guinea, give added weight toto thethe possibilitypossibility of Hydromyine host migration inin either direction.direction. ItIt throwsthrows SpecimensSpecimens deposited:deposited: Holotype USNM Helm. Coli. no. 89857, and open, however. the question of where thethe genus OdiliaOdilia maymay allotypeallotype 89858,89858, paratypesparatypes (1(1 male, pieces of 6 males, 4 females) BBM, Honolulu.Honolulu. Hawaii,Hawaii, reg.reg. no.no. 56592. have evolved. TypeType host:host: PseudohydromysPseudohydromys murinus. The finding of a new species of a spiruridspirurid P. kaindiensiskaindiensis may,may, TypeType locality:locality: Mt. Kaindi, 07°21 's's 146°41 'E,'E, Wau. Morobe Province. by contrast, be indicative of thethe relativerelative isolationisolationof muridmurid hostshosts SiteSite inin host:host: Stomach.Stomach. since their arrival in the New Guinea region.region. Protospirura sp.sp. Date of collection: December Date of collection: II December 1982. has not previously been reported fromfrom Australian rodents,rodents, butbut Etymology:Etymology: SpeciesSpecies namename isis takentaken fromfrom thethe type locality, the ecologically equivalent, superficially similarsimilar spiruridspirurid Mas­Mas­ RemarksRemarks tophorus muris Gmelin, 1780 has (Smales,(Smales, 1997).1997). FurtherFurther sur­sur­ veys are needed to determine whether Mastophorus isis restrictedrestricted AlthoughAlthough onlyonly 33 intactintact wonns were examined. the pieces of male to Australia and more recently arrived Rattus sp.sp. hosthost (Smales,(Smales, andand femalefemale wormsworms dissecteddissected fromfrom thethe host stomachs included both an M teriorterior andand posteriorposterior endsends thatthat were also examined. These measurements 1997) and Protospirura restricted toto New Guinea andand hydro­hydro­ werewere consistentconsistent withwith thosethose of thethe hototype, allotype, and paratype, given myine hosts. here.here. HasegawaHasegawa (1990)(1990) summarizedsummarized thethe status of Protospirura Seurat, ACKNOWLEDGMENTS 1914,1914, recognizingrecognizing 99 species.species. Pmtospirura kaindiensis n. sp" the 10th species,species, isis aa typicaltypical exampleexample of thethe group. My thanks to T. Flannery of thethe Australian .Museum.Museum a~da~d C.C. Kishi­Kishi­ ProtospiruraProtospirura kaindiensiskaindiensis cancan be readilyreadily differentiated from all other nami of the Bernice Bishop Museum forfor allowmgallowmg me toto dissectdissect hosts.hosts. membersmembers ofof thethe genusgenus inin havinghaving 5 pairs of postanal papillae (Protos­ This project was supported by an Australian Research Council Grant.Grant. pirurapirura okinavensisokinavensis Hasegawa, 1990 has 4 pairs and all the others more thanthan 55 pairs)pairs) andand thethe dimensionsdimensions of thethe spicules. LITERATURE CITED ~ther~ther distinguishingdistinguishing characterscharacters are as follows:follows: Protospiru:a anopIa BEVERIDGE, I., AND M.-C. DURRETIE-DESSET. 1992. Anew,species of KreiS,KreiS, 1938,1938, ProtospiruraProtospirura armeniana Alojan, 1951, Protosplrura cha,­ BEVERIDGE, I., AND M.-C. DURRETIE-DESSET. 1992. Anew,species of trichostrongyloid nematode, Odilia bainae from a native rodent, baudibaudi Vuylsteke,Vuylsteke, 1964,1964, Protospirura numidica Seurat, 1914 Protospr­ trichostrongyloid nematode, Odilia bainae from a native rodent, Rattus juscipes (Waterhouse). Transactions of the Royal Society of rurarura peromysciperomysci BaberoBabero andand Mathias, 1967, Protospirura suslica Schult, Rattus juscipes (Waterhouse). Transactions of the Royal Society of South Australia 116: 123-128. 19161916 andand P.P. okinavensisokinavensis havehave 4 denticles, not 2, on each element of South Australia 116: 123-128. DURRETI-DESSET, M.-C. 1985. Trichostrongyloid nematodes and t~e!rt~e!r thethe pseudolabia,pseudolabia, ProtospiruraProtospirura muricola Gedoelst, 1916 with 2 denticles DURRETI-DESSET, M.-C. 1985. Trichostrongyloid nematodes and vertebrate hosts: Reconstruction of the phylogeny of a parasItiC on each element of the pseudolabia has spicules of near equal length vertebrate hosts: Reconstruction of the phylogeny of a parasItiC on each element of the pseudolabia has spicules of near equal length group, Advances in Parasitology 24: 239-306. (u~out(u~out 0.42). Protospirura pseudomuris Yokohata and Abe, 1989, also group, Advances in Parasitology 24: 239-306. 0.42). Protospirura pseudomuris Yokohata and Abe, 1989, also FLANNERY, T. 1995. Mammals of New Guinea. Australian Museum, Wlth 2 denticles on each element of the pseudolabia (see Hasegawa. FLANNERY, T. 1995. Mammals of New Guinea. Australian Museum, Wlth 2 denticles on each element of the pseudolabia (see Hasegawa. Reed Books, Chatswood, Australia. 568 p. . .. 1990),1990), hashas thethe slenderslender rightright spiculespicule (0,68)(0,68) longerlonger than.th~br~ader left Reed Books, Chatswood, Australia. 568 p. . .. M AND D. M. SPRATI. 1995, Two newnew speciesspecies ofofOdiliaOdilia (0.33) (see Yokohata and Abe, 1989). Protospirura kamd,ensls has the GIBB ONS,•L ., )')' dd ' ' h h (0.33) (see Yokohata and Abe, 1989). Protospirura kamd,ensls has the (Nematoda: Heligmonellidae) fromfrom Austra IanIan roro ents,ents, WitWit com-com- broaderbroader rightright spiculespicule (0.46)(0.46) longerlonger thanthan thethe slender right (0.32),

...... f m Papua New Guinea. t.t. Anterior end.end. rightright laterallateral view.view. 2.2. FIGUREsFIGUREs 1-10.1-10. Protospirura kaindiensis n. sp. from pseudohydro~y: mu~mu~o:oventral view. 5. Male posterior end,end, ventralventral view,view, 6.6. Vagina,Vagina, ~ephalie~ephalie end, right lateral view. 3. En face view, 4. Cephalic end, optica sec~onFemale tail left lateral view. 10. Male posterior end, right lateral end, right lateral view. 3. En face view, 4. Cephalic end,gUber~a~u~~optica tail left lateral view. 10. Male posterior end, right lateral f1.ghtf1.ght laterallateral view.view. 7.7. Male tail,tail, ventral view. 8. Spicules and gUber~a~u~~ ,; 200 j.1m. '' VIew.VIew. ScaleScale BarsBars 1,1, 1010 = tootoo Il-ffi;Il-ffi; 2 = 25 f.Lm;f.Lm; 3. 4, 7, 8 = 50 J-l.m, •, 172 lHE JOURNAL OF PARASITOLOGY. VOL 87. NO 1. FEBRU~RY 2DO'

ments on O. bainae BcYeridgeBCYeridge & Durrette-DessetL 1991. S),stem­ t\Iuridae) of Irian Jaya., Indon~sia. Juurnal uf Ihe Hdminthotogic:l1 atic Parasitology 31: 67-75. Sodel}' of \\b.shingt..ln 61: 208-214. HAsEGAWA., H. 1990. Protospintra o/:ina\"ensis sp. n. (Nematoda: Spi­ S:Iot-u.ES. L. R. 1997. A review of Ihe helminth P;rr',J.sit~s of Australian ruridae) from Mus caroli on Okinawa Island. Japan. Journal of the rodents. Australian Journal of Zlll)logy 45: 50S-5:!J. Helminthological Society of 'Vashington 57: 153--156. YOKOH.UA. Y.• A..'\D H. ABE. 19S9. Two n~w ~pirurid nematodes in J:1p­ ---. AND SYAFRUDDIN. 1994. Odilia mallomyos sp. D. (Nematoda: anese moles. Mogt.'Tll spp. Jap:.mcsC' Journal of Par.l~i(()logy38: 92­ Heligmonellidae) from Mallomys l'O/hsdliidi meylandi (Rodentia:. 99.