A NEW GENUS AND SPECIES OF LUNGWORM (NEMATA: METASTRONGYLOIDEA) FROM AKODON

MOLLIS THOMAS, 1894 (RODENTIA: CRICETIDAE) IN PERU

Author(s): Maria Elizabeth Morales, John E. Ubelakert and Scott L. Gardner
Source: The Journal of Parasitology, Vol. 98, No. 3 (JUNE 2012), pp. 612-616
Published by: Allen Press on behalf of American Society of Parasitologists
Stable URL: http://www.jstor.org/stable/41511247
Accessed: 13-01-2016 17:24 UTC

REFERENCES
Linked references are available on JSTOR for this article:
http://www.jstor.org/stable/41511247?seq=1&cid=pdf-reference#references_tab_contents

You may need to log in to JSTOR to access the linked references.

Your use of the JSTOR archive indicates your acceptance of the Terms & Conditions of Use, available at http://www.jstor.org/page/
info/about/policies/terms.jsp

JSTOR is a not-for-profit service that helps scholars, researchers, and students discover, use, and build upon a wide range of content
in a trusted digital archive. We use information technology and tools to increase productivity and facilitate new forms of scholarship.
For more information about JSTOR, please contact support@jstor.org.

Allen Press and American Society of Parasitologists are collaborating with JSTOR to digitize, preserve and extend access to
The Journal of Parasitology.

http://www.jstor.org

This content downloaded from 155.247.166.234 on Wed, 13 Jan 2016 17:24:37 UTC
All use subject to JSTOR Terms and Conditions
 J.Parasito!., pp.612-616
2012,
98(3),
©American ofParasitologists
Society 2012

A NEW GENUS AND SPECIES OF LUNGWORM (NEMATA: M ET ASTRONG YLOI DE A) FROM

AKODON MOLLIS THOMAS, 1894 (RODENTI A: CRICETIDAE) IN PERU

MariaElizabethMorales*,JohnE. Ubelakert,and Scott L. Gardneri

ofProtozoología,
Department e Invertebrados
Helmintología Afines,Museode Historia Universidad
Natural, Nacional de San Marcos
Mayor
(MUSM),
Apartado Lima,Peru,e-mail:
14-0434, helminto_musm@yahoo.com
abstract: Akodonema luzsarmientae
n.g.,n.sp.(Nemata: is described
Metastrongyloidea) fromthepulmonary andheart
arteries
fromseveral of"soft
individuals mouse,"
grass Akodon mollis
(Rodentia:
Cricetidae), intheregion
collected Peru.Thenew
ofAncash,
genusandspecies bya reduction
isdistinguished rayto2 smallwidely
ofthedorsal separated
papillae.

Duringa studyof the helminth parasitesof the "softgrass Nematodes recovered werefixedandstored in 70%ethanol andlater
mollis habitat cleared with lactophenol forstudy.
mouse," Akodon Thomas,1894,from mountain shrub All nematodes usedin thisstudyweredeposited in theColección
onthewestern flankoftheAndesofPeru,anundescribed speciesof Helmintológica y de Invertebrados Relacionados delMuseode Historia
an angiostrongylid typeofnematode representing a newgenuswas Natural, MUSM,andintheParasite Collection oftheHarold W.Manter
foundinthepulmonary arteries andheart.Thenewspeciesshows Laboratory of Parasitology, University of Nebraska-Lincoln, Lincoln,
somesimilarity to speciesof Gallegostrongylus Nebraska. Measurements were madewith theaidofanocular micrometer
(Mas-Coma,1977) and measuring software (SigmaScanPro,Systat Software,Inc.,
digital
and Thaistrongylus (Ohbayashi, Kamiya,andBhaibulaya, 1979)in Chicago, Allmeasurements
Illinois). aregiven inmicrometers except where
thereduction, loss,orancestral absence,ofthedorsalray. noted,with therange followed bythemeaninparentheses. Drawings were
SpeciesofAkodon Meyen,1833,havea strictly Neotropical dis- madewith theaidofdrawing tubeattached toa CarlZeisslight microscope.
tribution,witha northern-most species,Akodon afflnisAllen,1912,
occurring inthewestern partoftheAndesinColombia(Eisenberg, DESCRIPTIONS
1989),andmanyotherspeciesdistributed overtheSouthAmerican
Akodonema n. gen.
continent to Tierradel Fuego,wherethecloselyrelatedAbothrix
hershkovitzi can be found(Patterson et al., 1984).Morethan40 Diagnosis: Elongate, delicate nematodes withbodyof bothsexes
filiform
andtapering tosmaller diameter atanterior end.Cuticular sheath
speciesofAkodon havebeendescribed (WilsonandReeder,2005), absent, cuticle with numerous fine, longitudinal striations.
Cephalicregion
andbecauseofcryptic species inthegenus, there areprobably many with 3,minute lips,eachbearing 2 small papillae. Esophagus claviformin
morethatarecurrently unrecognized. outline, excretory porewellanterior to esophageal-intestinal junction.
Akodon mollisis a smallrodent (meantotallength= 169.8mm) uterine Intestine filledwithbloodfromhost.Bothintestine andmilky-white
tubules spirally arranged within thepseudocoelom.
thatinhabits themontane regionandPacificlowlandsofnorthern Male:Tailcurved ventrally. Bursasmall, incomplete, 2 lateral
sidesnot
Perunorthward through thecentral highlands ofEcuador(Patton joinedanteriorly or posteriorly. Ventral andlateral raysofbursalong,
and Smith,1992)(Fig. 1). As is typicalof speciesincludedin narrow. Ventral rays fused formost oflength, lateralraysequal,arisingfrom
Akodon , their diet appears to consist of
mostly insects, seeds,and common trunk, divided to about 1/2of length. Dorsal ray but
absent, 2 small
anddelicate External dorsalrayspresent, elongate and
othervegetablematerial.There are no previousreportsof narrow. papillae present.
andwelldeveloped. Gubernaculum
Spicules thin,elongate present.
parasites fromA. mollis. Female: Oviparous andprodelphic withsimple ovejector.Vulvaopens
atposterior endjustanterior toanus,tailshort. Eggsellipticalinshape,
showing earlydevelopment ofjuveniles. Eggsdeposited intobloodstream,
MATERIALAND METHODS andlarvae hatch inlungs ofhost.
Between February 1988andOctober 1989,125specimens ofA. mollis
werecaptured usingSherman™ live traps,fromthe District of Taxonomie summary
Huayllacayán (lat10°14'56" S,long.77°26'00"W), Province ofBolognesi, Thegeneric nameis an arbitrary combination taken from
Department ofAncash, inthehigh-altitude punahabitat Etymology:
ofthecentral "Akodo " inreference tothehostrodent (Akodon) andtheword"nema,"
Andes ofPeru(2,100-3,256 m)(Fig.2). which means "thread" inGreek. Gender feminine.
Specimens ofAkodon inthisstudy werecaptured during a surveyofthe
reservoirs
potential ofLeishmania spp.bya research group of theInstitute
ofTropical Medicine "Alexander VonHumboldt" ofCayetano Heredia Akodonema luzsarmientae n. sp.
University in Lima,Peru,from19 different localities After
(caseríos). (Figs.1-7)
individualmiceweresampled forLeishmania spp.,they werekilled,and
thecarcasses frozen at -20 С and deposited in theDepartment of General: Bodyof bothsexesfiliform andtapering at anterior end.
oftheMuseode Historia Natural, Universidad Nacional Cuticle lightly transversely striated.Headsmoothly rounded. Mouth with
Mammalogy
Mayor deSanMarcos (MUSM),inLima,Peru.Subsequently, individual3 tinylips.Esophagus clubshaped. Excretory porecloseto nerve ring.
rodents werethawedand examined forparasites Nerve
usinga binocular Male ring in anterior third of esophagus.
in theLaboratory ofHelminthology at MUSM. (holotype and9 specimens): Bodylength 7.8to10.6mm;width at
dissectingmicroscope baseofesophagus 74to87,maximum width 110to162.Esophagus 150to
172longby35to39inmaximum width. Nerve ring 25to28from anterior
Received 22 January 2010;revised 24 November 2011;accepted 5 end.Excretory pore19to 22 from anterior end.Anterior endoftestes
December 2011. variable in position, notreaching baseofesophagus. Spiculesslender,
*Present address:Department of NaturalSciences, New Mexico conspicuously striated, equalor subequal, 230to 246longby5 wide;
Highlands University, LasVegas, NewMexico 87701. cephalic endsofspicules pointed, edgesirregular; distalendssmoothly
tDepartment of Biological Sciences,Southern Methodist University,rounded. Gubernaculum present, poorly cuticularized, 24 to 29 long.
Dallas,Texas75275. Bursasymmetrical andwelldeveloped forming twolateral lobes.Dorsal
{ Harold W.Manter Laboratory ofParasitology, University ofNebraska rayreduced to2 small papillae widely separated. Externodorsalrayslong,
StateMuseum, Lincoln, Nebraska 68588-0514. digitiform, arising separately at base.Lateral rayswithcommon trunk,
DOI: 10.1645/JP-GE-2440.1 posterolateral, mediolateral, and externolateral arising fromcommon

612

This content downloaded from 155.247.166.234 on Wed, 13 Jan 2016 17:24:37 UTC
All use subject to JSTOR Terms and Conditions
 ETAL.-ANEW
MORALES ANDSPECIES
GENUS OFLUNGWORM 613
METASTRONGYLOIDEA)
(NEMATA:

Figure1. Dorsalviewof"soft Akodon

mouse,"
grass mollis
Thomas,
1894(Cricetidae: Scalebar= 5 cm.
Akodontini).
Sigmodontinae,

trunk,allaboutequalinsizeandlength. Ventralrayssubequal, fused in

lower%,ventral trunkwidely separatefromlateraltrunk(Figs.5,6).
Female and8 other
(allotype paratype specimens;9 total
females included
intheparatype series):
Bodylength 17.5to26.9(21.6)mmwidth atbaseof
esophagus 100to165(135),maximum width240to320(274).Esophagus 132
to475longby114to117.Nerve 45to172from
ring anteriorend.Excretory
pore57to170from anterior
end.Anteriorendofovariesvariableinposition,
butnotextending anterior
tobaseofesophagus.Uterinetubules whiteinlife,
woundaround
spirally blood-filled
intestine a "barber-pole"
presenting
appearance common infamily.Vulvaopening neartipoftail166to 174.
Anus46to54from tipoftail.Phasmidial
pores notobserved. Tailbluntly
rounded. Eggsthinshelled,75 to 93 longand112to 196wide,laidin
pulmonary arteryanddeveloping andhatchinginlungtissue (Figs.1^4).
Taxonomie summary
Type host:Akodon mollis
Thomas, 1894,"softgrassmouse" (Cricetidae:
Sigmodontinae).
Symbiotypes: Akodon mollis Thomas, 1894,adultmalesandfemales,
Catalog numbers: 3608,3611,3623,3624,3628,3629,3642,3643,3645,
3649,3650,5760,5778,5782,5786,5787,5893,all deposited in the
Mammals Collection attheMuseodeHistoria Natural,MUSM.
Typelocality: Districtof Huayllacayán (10°14'56"S,77°26'00"W),
Province ofBolognesi, Department ofAncash, Peru.
Siteofinfection: Pulmonary arteryandbranches, heart,andlungs.
Prevalence Rodents
of infection: examined, 125of which27 were
infected (21.6%).
Intensity: Oneto9 nematodes. Totalnematodes recovered,78(34from
heart and44from lungs).
Typespecimens: Holotype male:ColecciónHelmintológica y de Figure2. Map showing the studyarea in oval (District of
Invertebrados Relacionados, Museode HistoriaNatural,MUSMm Huayllacayán, Province ofBolognesi, Department ofAncash, Peru)and
No. 1699.Allotype female:Colección HelmintológicaydeInvertebrados ofA.mollis
thedistribution Thomas, 1894,inPeru(shaded).
Relacionados, MUSM,No. 1700.Paratypes: ColecciónHelmintológica y
deInvertebrados Relacionados MUSM,Nos.1701,1837,1838,1839, and
Harold W.Manter ofParasitologyCollection thebasisof fragmented maleand female specimens. Thedescription
Laboratory (HWML48041, thatthegenus hasa strongly reduced bursa, theventral
HWML48042). suggests including
Thespecific namehonors Dr. Luz Sarmiento, Emeritus and lateral
rays,and the dorsal ray is reduced to andshort
papilla-like
Etymology: arcuate Ubelaker (1986)argued thatthegenus isverysimilar
to
Professor oftheUniversidad Nacional Mayor deSanMarcos, Lima,Peru, spicules.
andfounder oftheHelminthological Collectionat theNatural andshouldbe placedin synonymy.
Gallegostrongylus Thedifferences
History between thegenera Gallegostrongylus andThaistrongylus aremainly in
Museum, MUSM,in recognition of bothhercontribution to the
how the bursaof thelatterisreduced and ifthe of
lobes thedorsal
knowledge ofthePeruvian helminthology andherinspirational teaching are strongly
forthestudy ofprotozoan andother animal ray papilla-like or consistofveryshortrays.On theother hand,
parasites. Akodonema hasventral andlateral raysstrongly developed,similar
to
either orParastrongylus
Angiostrongylus (seeUbelaker, 1986forreview;
Remarks seealsoRoblesetal.,2008),andthedorsal rayisclearly reducedtowidely
Allmembers ofthisgroup ofworms dwellinthetissues oftheir hosts, separated papillae.The spiculesare also of the Angiostrongylus-
often inthebloodstream (Spratt, 1961).Thenewgenus, Akodonema, is Parastrongylus
type,being longandthin.
characterized byhaving well-developed ventral andlateralrays and with
thedorsal rayreduced toa pairofsub-ventral Thespicules
papillae. are DISCUSSION
elongate. The morphology of the bursaallowscomparisons with
Aelurostrongylus Cameron, 1927, MarsupostrongylusMackerras and
Sandars, 1953, Webster, 1978,
Within theAngiostrongylidae as described byAnderson (1978),2
Andersonstrongylus ThaistrongylusOhbaya- of are evident.A first ischaracterized
shi,KamiyaandBhaibulaya, 1959,and Gallegostrongylus Mas-Coma, groups genera group by having
1977, allofwhich havea reduced dorsalray,orthedorsalrayis deeply intact dorsalrays.Thisgroupshould remain intheAngiostrongylidae.
divided orabsent and/or papillaearepresentinitsplace.Thenewgenus is
most closelyassociated with thelatter2 genera.According toMas-Coma
(1977),in Gallegostrongylus , thedorsalrayis deeply divided witheach REDESCRIPTION
branch oftherayarising separately anddistant from eachother. The 1978
ventral andlateral Angiostrongylidae Anderson,
raysarewelldeveloped, andthespicules areshort,
delicate, and arcuate.As described by Ohbayashi et al. (1979), Diagnosis: With the charactersof the Bursa
family. highly developedor
Thaistrongylus isrepresented bya singlespecies,Thaistrongylusharinasu-reduced, complete,buttypical raysalways clearly Dorsalrayis
defined.
tai, collectedfrom a single
host(Rattus berdmoreiBlyth)anddescribed on present butsometimes divided.
Ventral andlateral rayswelldeveloped.

This content downloaded from 155.247.166.234 on Wed, 13 Jan 2016 17:24:37 UTC
All use subject to JSTOR Terms and Conditions
614 THEJOURNAL
OFPARASITOLOGY,
VOL.98,NO.3,JUNE
2012

Figures3-9. Akodonema luzsarmientaen.g.n. sp.,(3) anterior

endofgravid female from A. mollis
Thomas, 1894,ventral
view.(4) EggofA.
luzsarmientae
n.g.n.sp.(5)Magnified
posterior extremity ofA.luzsarmientae
n.g.n.sp.,showinganusandglands with
associated rectum.
(6)Posterior
ofgravid
extremity female ofA. luzsarmientaen.g.n. sp.showing thevulva, andeggsintheuterus.
vagina (7) Posterior ofa maleofA.
extremity
luzsarmientae
n.g.n.sp.,lateral
view.(8) Posterior
extremity ofa maleofA. luzsarmientaen.g.n.sp.,ventral viewofspicules
view.(9) Ventral ofA.
n.g.n.sp.Scalebars= 20(xm
luzsarmientae (Figs.3,5); 100ц.т(Figs.4,6);40jim(Figs.7-9).
Remarks Romanov, 1952,StefanskostrongylusDrozdz,1970,andTribo strongylus
Anderson, 1963.A keyforthesegenera isprovidedby Anderson (1978).
inthisfamily
Included areAngiostrongylus
Kaminski,
1905,
Didelphos- A second groupofgenera splitintoseveral generally
species a
displays
Prestwood,
trongylus 1976,Filostrongylus
Mackerras,1955,Gurtlia dorsalraymodified from a single intact to a pairofdivided
structure
1933,Heterostrongylus
Wolffhugel, Travassos,
1925,Madangiostrongylus
papillaeor papillae-like
structures.Thisgroupofnematodes willneed
ChabaudandBrygo, 1960,Parastrongylus 1928,Pulmostrongylus
Baylis, revision, after
preferably anadequate analysis
phylogenetic is conducted
Rodentocaulus
Hsu,1935, OrlovandKutass,
Schulz, 1939, ofthegroup.
Sobolevingylus

This content downloaded from 155.247.166.234 on Wed, 13 Jan 2016 17:24:37 UTC
All use subject to JSTOR Terms and Conditions
 ETAL.-ANEW
MORALES ANDSPECIES
GENUS OFLUNGWORM 615
METASTRONGYLOIDEA)
(NEMATA:

Figure10. Akodonema n.g.n.sp.intheheart

luzsarmientae ofitshost.
Arrowshowsa female
wormin theheartofA. mollis Thomas, 1894, Figure11. Akodonema n.g.n.sp.in thelungsofA.
luzsarmientae
original
magnification
(25X). mollis
Thomas,
1894,original (20X).
magnification

Generaincluded in thisgroupincludeAkodonema n. gen.,Aeluros-hasbeenshownwithA. morerai inpopulations ofA. azaraefrom

Cameron,
trongylus 1927,Andersostrongylus 1978,Antechinos-
Webster, Argentina (Robles et al., 2008). However, further studiesare
trongylusSpratt,1981,MadafilaroidesChabaudand Brygoo, 1960, needed to demonstrate if these infections are fatal; some
Gallego lusMas-Coma,
strongy Mackerras
1977,Marsupostrongylus and
andBhaibulaya,experimental withAngiostrongylus
infections schmidti werenot
Sandars,1953,andThaistrongylusOhbayashi,
Kamiya,
1979. fatalto larger hostssuch as cotton ratsand white rats,butthey
Thisgroupcanbedistinguished fromtheAngiostrongylidae
mainlyby were fatalto rodents with smallerbody size(Kinsella, 1987).The
thecharacters
ofthebursathatshowa gradual as discussed
reduction, by present studyconstitutes a thirdreportofan angiostrongylid-type
Webster(1978).Thisgroupshows tonematodes
relationships to
belonging of a smallmammalin Peruand extendsthehostand
theFilarioidea
inwhich thecaudalregion
ofthemales remnantsparasite
possesses
ofventralandlateral geographical distributionofthisnematode to thePeruvian Andes
rays.
in SouthAmerica.
DISCUSSION ACKNOWLEDGMENTS
To date,only2 speciesof angiostrongylid-types ofnematodes We thankDr. Alejandro Llanosand Abraham Cáceresforthe
have beenreported frommammalsin Peru.Theyare Angios- opportunity to examine therodents at theMuseode Historia Natural,
MUSM.Specialthanks arealsodueto Dr. BenNelson, NewMexico
trongylus costaricensisMorera and Céspedes, 1971, in the LasVegas, NewMexico, forcontinued inhis
Highlands
University, support
mesenteric vesselsof "pichicobarba blanca,"Saguinusmystax laboratory byproviding working spaceto thefirstauthor. Thanks to
(thetamarin), from Iquitos(Loreto),andParastrongylus sp.inthe Professor R. Severino,oftheLaboratorio deZoologiadeInvertebrados of
pulmonary vein of "the paintedbig-eared mouse,"Auliscomys theFacultadde Ciencias of
Biológicas the Nacional
Universidad Mayor de
SanMarcos(MUSM),Lima,Peru,forhiscontinued encouragement and
pictus,fromCasapalca (Lima) (Sarmientoet al., 1999).Until valuable helpwith laboratoryresources. thanks
Special toElenaVivar of
recently,just2 previously reportsof angiostrongylidsin rodents the Department of Mammalogy of MUSM forcorroborating the
fromSouth Americawere known:Angiostrongylus lenziiin identificationof therodents and providinginformation of thehost.
Akodonmontensis Thomas,1913,fromRio de Janeiro,Brazil Furthermore, thanks toDr.Victor Pacheco forhishelpwiththeAkodon's
erai in Akodon picture andreferences.Thanks areduetoLeonardo Romero oftheFaculty
(Souza et al., 2009), and Angiostrongylus mor of BiologicalScience of MUSM forhistechnical assistance withthe
azarae(Fischer,1829)fromBuenosAires,Argentina (Robleset photographs andDr.DianaSilvafrom MUSMforhiscomments onthe
al., 2008);bothnematodes werefoundin thepulmonary arteries manuscript.Support wasalsoprovided bytheJ.Teague SelfEndowment
oftheirrodenthosts. Fundforvisiting scholarsin theHaroldW. Manter Laboratory of
Akodonis one of themostdiversegeneraof rodentsin Peru, Parasitology, University ofNebraska-Lincoln,Lincoln,Nebraska.
with13 speciesrecognized to date and manyothersyetto be
described LITERATURECITED
(Pachecoet al., 2009).The helminthfaunaofA. mollis
from Peru is apparentlydiverse,and the findingof A. Anderson, R. C. 1978. Keyto genera ofthesuperfamily Metastrongy-
luzsarmientae n. could be the fact of the loidea.
No.5.In Keystothenematode ofvertebrates,
parasites R.C.
n.g. sp. corroborating Anderson, A. G. Chabaud,and S. Wilmot Commwealth
correlationbetweenspeciesrichness and (eds).
positivecoevolutionary AgriculturalBureaux. Farnham Royal,U.K.,p. 1^0.
altitude(M. Morales,pers.obs.); however, further studiesare Eisenberg, J. F. 1989.Mammals of theNeotropics. The northern
necessary to testthishypothesis. Neotropics.Vol.I. Panama, Colombia,Venezuela,Guyana, Suriname,
It is possiblethatA. luzsarmientaemaybe a causeofmortality FrenchGuiana.University of ChicagoPress,Chicago, Illinois,
inpopulations 449p.
ofA. mollisintheAndes,becausethisparasite may Kinsella,J.M. 1987.Studies on thelifecycleandhostspecificity of
provokeinflammation of thelungand heartvessels,interfering Parastrongylus schmidti(Nematoda: Proceedings
Angiostrongylidae).
withthenormalfunction of thesevitalorgans(Figs.10-11),as oftheHelminthological SocietyofWashington 54:245-248.

This content downloaded from 155.247.166.234 on Wed, 13 Jan 2016 17:24:37 UTC
All use subject to JSTOR Terms and Conditions
616 THEJOURNAL
OFPARASITOLOGY, 2012
VOL.98,NO.3,JUNE

Mas-Coma, S. 1977.Gallegostrongylus n.gen.,

ibicensis n.sp.(Nematoda: Akodon azarae(Rodentia: in Argentina.
Cricetidae) Journal of
Metastrongylidae),parasite
pulmonaire de Musmusculus Linnaeus, Parasitoloev94:515-519.
1758(Rodentia: Muridae)a Ibiza(Baleares).
Annales L.,M.Tantaleán,
deParasitolo-Sarmiento, andA.Huiza.1999. Nemátodos parásitos
gicHumaine etComparée 52:637-642. del hombre y de los animales en el Perú.RevistaPeruanade
Ohbayashi,M.,M. Kamiya, andM. Bhaibulaya. 1979.Studiesonthe Parasitología14:9-65.
parasitefaunaofThailand 1. Twonewmetastrongylid S. A.R. С. Thiengo,
nematodes,Souza,J.,R. O. Simões, W.S. Lima,E. M.Mota,R.
Angiostrongylussiamensis
sp.n. andThaistrongylusharinasutai
gen. Rodriguez-Silva, andA. Maldonado,
R. M. Lanfredi, Jr.2009.A
et sp. n. (Metastrongyloidea; fromwildrats.
Angiostrongylidae) newMetastrongylidae species(Nematoda: A
Metastrongylidae):
Japanese JournalofVeterinaryResearch 27:5-10. lungworm fromAkodon montensis(Rodentia:Sigmodontinae)in
Pacheco,V. R.,R. Cadenillas, E. Salas,С. Tello,andH. Zeballos. Brazil.
Journal ofParasitology95:1507-1511.
2009.Diversidad y endemismo de losmamíferos delPerú.Revista Spratt,D. M. 1961.Antechinostrongylus gen.et sp.
disgubernaculus
Peruana deBiología 16:5-32. (Nematoda: from
Angiostrongylidae) themarsupialmouse Antechinus
Patterson,В.D.,M.H. Gallardo,andК. E. Freas.1984.Systematic of swainsonii
(Marsupialia: Journal
dasyuridae). ofParasitology
67:90-93
miceof thesubgenus Akodon (Rodentia: in southernUbelaker,J. E. 1986.Systematics
Cricetidae) of speciesreferred
to thegenus
SouthAmerica. Fieldiana,
Zoology, newser.23:1-16. Journal
Angiostrongylus. ofParasitology
72:237-244.
Patton,J.L., and M. F. Smith.1992.Evolution andsystematicsof Webster, W. A. 1978.Andersostrongylus gen.et sp. n.
captivensis
akodontine rodents(Muridae:Sigmodontinae)ofPeru, withempha- (Metastrongyloidea: fromthelungsof thestriped
Filaroididae)
sisonthegenus Akodon. Memorias delMuseodeHistoria Natural, shunk,Mephitis mephitis.Journal
ofParasitology64:459-462.
Universidad NacionalMayor deSanMarcos, Lima21:83-103. Wilson,D. E., and D. M. Reeder.2005.Mammalspeciesof the
Robles,M. Del R., G. T. Navone,and M. Kinsella.2008.A new world.A taxonomie and geographic reference.
JohnsHopkins
Angiostrongylid(Nematode) speciesfrom thepulmonary arteries
of UniversityPress,Baltimore,Maryland.2,142p.

This content downloaded from 155.247.166.234 on Wed, 13 Jan 2016 17:24:37 UTC
All use subject to JSTOR Terms and Conditions