REVIEW ARTICLE /ARTÍCULO DE REVISIÓN
UPDATE OF CHECKLIST OF DIGENEAN PARASITES OF WILD BIRDS FROM
ARGENTINA, WITH COMMENTS ABOUT THE EXTENT OF THEIR INVENTORY
ACTUALIZACIÓN DEL LISTADO DE DIGENEOS PARÁSITOS DE AVES SILVESTRES
DE LA ARGENTINA, CON COMENTARIOS ACERCA DE SU INVENTARIO
1 1*
Fabiana Beatriz Drago & Lía Inés Lunaschi
1 División Zoología Invertebrados, Museo de La Plata, UNLP, Paseo del Bosque S/Nº, 1900 La Plata, Buenos Aires Province,
Argentina. *Financial support: UNLP (grant 11/N751) and CIC (grant 833/14)
Corresponding author at: División Zoología Invertebrados, Museo de La Plata, UNLP, Paseo del Bosque S/Nº, 1900 La
Plata, Buenos Aires Province, Argentina. E–mail address: fdrago@fcnym.unlp.edu.ar
Neotropical Helminthology, 2015, 9(2), jul-dec: 325-350.
ABSTRACT
Keywords: Argentina - checklist - Digenea - inventory - parasites - wild birds.
We present an update of the checklist of digenean parasites of birds from Argentina published by
Lunaschi et al. (2007), including all additional records published since 2007 up to present and
some records omitted in the preceding version. This new report includes information about 74
digenean taxa, distributed in 18 families and 45 genera, of which 68 species were reported
parasitizing wild birds, 6 species were obtained experimentally, and 6 taxa identified at the
generic level were recovered in wild birds and/or experimental host. Consequently, the actual
number of digeneans recorded from Argentinean wild birds amounts 112 species in 65 genera of
20 families. The extension of this inventory was evaluated together with the number of host-
parasite relations reported and the geographical distribution in the different ecoregions of the
country. The species accumulation curve and the accumulation curve of host-parasite relations
show that the inventory of digeneans of wild birds is far from being completed, even in the most
studied areas. The taxonomic status of some species listed in Lunaschi et al. (2007), such as
Lyperosomum oswaldoi, Parastrigea brasiliana, Lyperorchis inexpectabilis and Stephanoprora
podicipei, was reconsidered according to new studies.
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ISSN Versión impresa 2218-6425 ISSN Versión Electrónica 1995-1043
INTRODUCTION
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Neotropical Helminthology. Vol. 9, Nº2, jul-dec 2015
We present an update of Lunaschi et al. (2007),
including all additional records published
since 2007 up to present, some omitted reports
in the preceding version, and the taxonomic
relocation of some species listed in the former
work, according to new investigations.
Lunaschi et al. (2007) list information
published between 1909 and 2006, consisting
of 89 adult digenean species of birds from
Argentina (81 species parasitizing natural host
and 8 obtained experimentally) and 6
digeneans identified at generic level. In the last
8 years, the number of papers on digeneans
from birds had a significant increase. In this
update, we present information about 69 adult
digenean species from birds (65 parasitizing
wild birds and 4 obtained experimentally) and
7 digenea identified at generic level. The aim
of this paper was to estimate the extent of
inventory of digenean parasites of Argentinean
wild birds, as well as the number of host-
parasite relations using species accumulation
curves, and to evaluate the distribution of these
helminths in different regions of the country.
This update was prepared on the basis of data
published since 2007 to present from various
regions of Argentina. The list of digenean taxa
is presented in alphabetical order, within the
different taxonomic hierarchy. Each record
contains information about the species name,
authorship and year, as well as the synonymous
under which original records appeared,
host(s), site of infection and localities
(geographical coordinates when possible).
Information on the life cycle of species listed
herein or by Lunaschi et al. (2007) is also
given. Detail concerning the material
repository is listed, where this information was
included in the original publication: collection
acronym and code numbers. Reference to the
literature is arranged chronologically. Species
RESUMEN
Palabras clave: Digenea – parásitos - aves silvestres – listado – inventario - Argentina.
En este trabajo se presenta una actualización del listado de digeneos de aves de Argentina
publicado por Lunaschi et al. (2007), incluyendo todos los registros adicionales publicados desde
2007 hasta la actualidad y algunos reportes omitidos en la versión precedente. Estos nuevos
registros incluyen información de 74 taxa de digeneos distribuidos en 18 familias y 45 géneros,
de los cuales 68 especies fueron reportadas parasitando aves silvestres, 6 especies fueron
obtenidas experimentalmente y 6 taxa identificados a nivel genérico reportados en aves silvestres
y/o hospedadores experimentales. Por consiguiente, el número actual de digeneos reportados en
las aves silvestres de la Argentina alcanza 112 especies distribuidas en 65 géneros de 20 familias.
La extensión de este inventario fue evaluada junto con el número de relaciones hospedador-
parásito reportadas y la distribución geográfica en las distintas ecoregiones del país. Las curvas
de acumulación de especies y de relaciones hospedador-parásito muestran que el inventario de
digeneos de aves silvestres se encuentra lejos de ser completado, aún en las áreas más estudiadas.
La posición taxonómica de algunas especies listadas en Lunaschi et al. (2007), tales como
Lyperosomum oswaldoi, Parastrigea brasiliana Lyperorchis inexpectabilis y Stephanoprora
podicipei, fue reconsiderada de acuerdo a nuevos estudios.
Drago & Lunaschi
MATERIAL AND METHODS
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Update of checklist of Digenean parasites of wild birds from Argentina
referred to in theses, dissertations and
scientific meetings were not considered. The
taxonomic classification of digeneans is
presented according to Gibson et al. (2002),
Jones et al. (2005) and Bray et al. (2008). The
nomenclature of birds follows Remsen et al.
(2014). The map of ecoregions of Argentina is
taken from Brown & Pacheco (2006).
Codes for the helminthological collections are
as follow: collection at the Natural History
M u s e u m , L o n d o n , U K ( B M N H ) ;
Parasitological Collection of the Centro
Nacional Patagónico, Puerto Madryn,
Argentina (CNP-Par); Invertebrates collection
of the Museo Argentino de Ciencias Naturales
Bernardino Rivadavia, Buenos Aires,
Argentina (MACN-Pa); Helmintological
Collection of the Museo de La Plata (MLP-
He), Buenos Aires Province, Argentina;
Parasitological Collection of the Universidad
Nacional del Comahue, Bariloche, Argentina
(UNCo-Pa). Abbreviations for the life cycle
are as follow: Hi : first intermediate host and
1
Hi : second intermediate host. In order to
2
evaluate the extent of the inventory, only taxa
identified at specific level were counted.
A total of 74 reports of adult digeneans from
birds are listed here. The new data consist
mostly of species new to science, new host
records, new geographical records and data on
their life cycle. These reports include 68
species parasitizing wild birds, 6 species
obtained experimentally, and 6 taxa identified
at generic level recovered in wild birds and/or
experimental host. These species are
distributed in 18 families and 45 genera. There
was an increase of 32 species in the last 8 years.
A total of 29 bird species are listed as definitive
hosts of digeneans, two species of birds and
one of mammals as experimental hosts, and 27
hosts are listed as intermediate hosts (Table 1).
Superfamily Brachylaimoidea Joyeux &
Foley, 1930
Family Brachylaimidae Joyeux & Foley, 1930
Genus Brachylaima Dujardin, 1843
Brachylaima yupanquii Freitas, Kohn &
Ibáñez, 1967
Host: Cariama cristata (Linnaeus, 1766)
(Cariamidae).
Site of infection: small intestine.
Locality: La Marcela farm (26°17'35''S;
59°06'67''W), Pirané, Formosa Province.
Material deposited: MLP-He 6444.
Reference: Lunaschi & Drago (2012).
Genus Glaphyrostomum Braun, 1901
Glaphyrostomum propinquum Braun, 1901
Host: Gui ra gui ra (Gmelin, 1788)
(Cuculidae).
Site of infection: cloaca.
Locality: La Marcela farm (26°17'35”S;
59°06'67”W), Pirané, Formosa Province.
Material deposited: MLP-He 5575.
Reference: Lunaschi & Drago (2009a).
Family Leucochloridiidae Poche, 1907
Genus Pojmanskia Zamparo, Brooks &
Causey, 2003
Pojmanskia riosae Zamparo, Brooks &
Causey, 2003
Host: Taraba major (Vieillot, 1816)
(Thamnophilidae).
Site of infection: cloaca.
Locality: La Marcela farm (26°17'35''S;
59°08'38''W), Pirané, Formosa Province.
Material deposited: MLP-He 6731.
Reference: Lunaschi et al. (2014).
Superfamily Clinostomoidea Lühe, 1901
Family Clinostomidae Lühe, 1901
Genus Clinostomatopsis Dollfus, 1932
Clinostomatopsis sorbens (Braun, 1899)
Dollfus, 1932
Host: Tigrisoma lineatum (Boddaert, 1783)
(Ardeidae).
Site of infection: esophagus.
RESULTS
Locality: Bellaco stream (26º14'S; 58º07'W),
Pirané, Formosa Province.
Material deposited: MLP-He 5573.
Reference: Lunaschi & Drago (2009a).
Genus Clinostomum Leidy, 1856
Clinostomum marginatum (Rudolphi, 1819)
Braun 1899
Host: Tigrisoma lineatum (Boddaert, 1783)
(Ardeidae).
Site of infection: esophagus.
Locality: Bellaco stream (26º14'S; 58º07'W),
Pirané, Formosa Province.
Material deposited: MLP-He 5574.
Reference: Lunaschi & Drago (2009a).
Superfamily Cyclocoeloidea Stossich, 1902
Family Cyclocoelidae Stossich, 1902
Genus Spaniometra Kossack, 1911
Spaniometra variolaris (Fuhrmann, 1904)
Host: Rostrhamus sociabilis (Vieillot, 1817)
(Accipitridae).
Site of infection: air-sacs.
Locality: La Marcela farm (26°17'35”S;
59°06'38”W), Pirané, Formosa Province.
Material deposited: MLP-He 6718, MLP-He
6719.
Reference: Drago et al. (2014).
Family Eucotylidae Cohn, 1904
Genus Tanaisia Skrjabin, 1924
Tanaisia dubia Freitas, 1951
Hosts: Himantopus melanurus Vieillot, 1817
(Recurvirostridae), Pitangus sulphuratus
(Linnaeus, 1766) (Tyrannidae).
Site of infection: ureters.
Locality: La Marcela farm, (26°17'35”S;
59°08'38”W), Pirané, Formosa Province.
Material deposited: MLP-He 6771, MLP-He
6672.
Reference: Lunaschi et al. (in press).
Superfamily Diplostomoidea Poirier, 1886
Family Diplostomidae Poirier, 1886
Genus Austrodiplostomum Szidat & Nani,
1951
Austrodiplostomum mordax Szidat & Nani,
1951
Host: Phalacrocorax brasilianus (Gmelin,
1789) (Phalacrocoracidae).
Site of infection: intestine.
Locality: Lacombe lagoon (35º49'S; 57º49'W),
Lezama, Buenos Aires Province.
Material deposited: MLP-He 6411.
Life cycle: Hi : metacercaria e of
2
Austrodiplostomum cf. mordax, in brain, optic
nerve and spinal chord in Odontesthes
bonariensis (Valenciennes, 1853), from
Lacombe and Salada Grande lagoons, Buenos
Aires Province.
References: Drago et al. (2011), Drago (2012).
Austrodiplostomum ostrowskiae Dronen, 2009
Host: Phalacrocorax brasilianus (Gmelin,
1789) (Phalacrocoracidae).
Site of infection: large intestine.
Locality: La Marcela farm (26°17'S;
59°06'W), Pirané, Formosa Province.
Material deposited: MLP-He 6412.
Reference: Drago et al. (2011).
Genus Dolichorchis Dubois, 1961
Dolichorchis lacombeensis Lunaschi &
Drago, 2006
Hosts: Ardea cocoi Linnaeus, 1766; Ardea
alba Linnaeus, 1758 (Ardeidae).
Site of infection: intestine.
Locality: Lacombe lagoon (35° 49'S; 57°
49'W), Lezama, Buenos Aires.
Material deposited: MLP-He 5972; MLP-He
5973.
Reference: Drago & Lunaschi (2011a).
Genus Hysteromorpha Lutz, 1931
Hysteromorpha triloba (Rudolphi, 1819) Lutz,
1931
Host: Phalacrocorax brasilianus (Gmelin,
1789) (Phalacrocoracidae).
Site of infection: intestine.
Localities: Lacombe lagoon (35º49'S;
57º49'W), Lezama, Buenos Aires Province; La
Marcela farm (26°17'S; 59°06'W), Pirané and
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Clorinda (25°17'S; 57°43'W), Formosa
Province.
Material deposited: MLP-He 6414.
Reference: Drago et al. (2011).
Genus Neodiplostomum Railliet, 1919
Neodiplostomum travassosi Dubois, 1937
Host: Buteogallus meridionalis (Latham,
1790) (Accipitridae).
Site of infection: intestine.
Locality: La Marcela farm (26°17'35”S;
59°06'38”W), Pirané, Formosa Province.
Material deposited: MLP-He 6714.
Reference: Drago et al. (2014).
Genus Posthodiplostomum Dubois, 1936
Posthodiplostomum macrocotyle Dubois,
1937
Host: Busarellus nigricollis (Latham, 1790)
(Accipitridae).
Site of infection: small intestine
Locality: La Marcela farm (26°17'35”S;
59°06'38”W), Pirané, Formosa Province.
Material deposited: MLP-He 6716, MLP-He
6717.
Reference: Drago et al. (2014).
Posthodiplostomum nanum Dubois, 1937
Host: Ardea alba Linnaeus, 1758 (Ardeidae)
Site of infection: intestine.
Localities: Lacombe lagoon (35º49'S;
57º49'W), Lezama, Buenos Aires Province and
Clorinda (25°17'S, 57°43'W), Formosa
Province.
Material deposited: MLP-He 5974, MLP-He
5975.
Host: Tigrisoma lineatum (Boddaert, 1783)
(Ardeidae).
Site of infection: intestine.
Locality: Bellaco stream (26º14'S; 58º07'W),
Formosa Province.
Material deposited: MLP-He 5977.
Reference: Drago & Lunaschi (2011a).
Posthodiplostomum sp.
Host: Ardea cocoi Linnaeus, 1766 (Ardeidae).
Site of infection: intestine.
Locality: Lacombe lagoon (35º49'S;
57º49'W), Lezama, Buenos Aires Province.
Material deposited: MLP-He 6013.
Reference: Drago & Lunaschi (2011a).
Posthodiplostomum sp.
Natural definitive host: unknown.
Experimental hosts: Gallus gallus (Linnaeus,
1758) (Phasianidae), Mus musculus Linnaeus,
1758 (Muridae).
Life cycle: Hi : sporocysts and cercariae in
1
Anisancylus obliquus (Broderip & Sowerby,
1832). Hi : metacercariae in Galaxias
2
maculatus (Jenyns, 1842) from Patagua Lake,
Neuquén Province.
Material deposited: experimental adults from
chicks MACN-Pa 558/11-12, UNCo-Pa
239/1-6; from mice MACN-Pa 558/ 8-10,
UNCo-Pa 240/1-13; metacercariae MACN-Pa
558/5-7, MACN-Pa 558/2-4, UNCo-Pa 241/1-
12, UNCo-Pa 242/1-3; sporocyst MACN-Pa
558/1, UNCo-Pa 243/1-3.
Reference: Ritossa et al. (2013).
Genus Sphincterodiplostomum Dubois, 1936
Sphincterodiplostomum musculosum Dubois,
1936
Host: Tigrisoma lineatum (Boddaert, 1783)
(Ardeidae).
Site of infection: intestine.
Locality: Bellaco stream (26º14'S; 58º07'W),
Pirané, Formosa Province.
Material deposited: MLP-He 5970.
Reference: Drago & Lunaschi (2011a).
Genus Tylodelphys Diesing, 1850
Tylodelphys adulta Lunaschi & Drago 2004
Host: Phalacrocorax brasilianus (Gmelin,
1789) (Phalacrocoracidae).
Site of infection: small intestine.
Locality: Clorinda (25°17'S; 57°43'W),
Formosa Province.
Material deposited: MLP-He 6413.
Reference: Drago et al. (2011).
Tylodelphys brevis Drago & Lunaschi, 2008
Host: Mycteria americana Linnaeus, 1758
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Update of checklist of Digenean parasites of wild birds from Argentina
(Ciconiidae).
Site of infection: Small intestine.
Locality: Clorinda (25°17'S; 57°43'W),
Formosa Province.
Material deposited: Holotype, MLP-He 5741;
paratypes MLP-He 5742.
Reference: Drago & Lunaschi (2008).
Host: Busarellus nigricollis (Latham, 1790)
(Accipitridae).
Site of infection: intestine.
Locality: La Marcela farm (26°17'35”S;
59°06'38”W), Pirané, Formosa Province.
Material deposited: MLP-He 6715.
Reference: Drago et al. (2014).
Tylodelphys elongata (Lutz, 1928) Dubois,
1937
Host: Tigrisoma lineatum (Boddaert, 1783)
(Ardeidae).
Site of infection: intestine.
Locality: Bellaco stream (26º14'S; 58º07'W),
Pirané, Formosa Province.
Material deposited: MLP-He 5971.
Reference: Drago & Lunaschi (2011a).
Family Strigeidae Railliet, 1919
Genus Apharyngostrigea Ciurea, 1927
Apharyngostrigea ardearum (Lutz, 1928)
Dubois, 1968
[Syn. Apharyngostrigea brasiliana Szidat,
1929 of Labriola & Suriano (1998)]
Site of infection: intestine.
Host: Ardea alba Linnaeus, 1758 (Ardeidae).
Localities: Lacombe lagoon (35º49'S,
57º49'W), Lezama, Buenos Aires Province and
Clorinda (25°17'S; 57°43'W), Formosa
Province.
Material deposited: MLP-He 5965, MLP-He
5966.
Host: Ardea cocoi Linnaeus, 1766 (Ardeidae).
Locality: Lacombe lagoon (35º49'S, 57º49'W),
Buenos Aires Province.
Material deposited: MLP-He 5967.
Hosts: Ardea alba Linnaeus, 1758, Bubulcus
ibis (Linnaeus, 1758) (Ardeidae).
Locality: De Monte Lagoon, Buenos Aires
Province.
Material deposited: MLP-He 5968, MLP-He
5969.
Comments: In Argentina, this species was
cited by Labriola & Suriano (1998) as
Apharyngostrigea brasiliana (Szidat, 1928)
Szidat, 1929 parasitizing A. alba and B. ibis
from De Monte lagoon. Later, was listed as
Parastrigea brasiliana (Szidat, 1928) Dubois,
1964 in Lunaschi et al. (2007). Finally, these
specimens were re-examined and transferred
to A. ardearum (see Drago & Lunaschi,
2011a).
Reference: Drago & Lunaschi (2011a).
Genus Australapatemon Sudarikov, 1959
Australapatemon canadensis Dubois &
Rausch, 1950
Host: Cygnus melancoryphus (Molina, 1782)
(Anatidae).
Site of infection: small intestine.
Localities: Lacombe lagoon (35º51'42''S,
57º53'39''W), Lezama, Buenos Aires and
Pellegrini Lake, Cinco Saltos (38º42'15''S,
67º59'47''W), Río Negro Province.
Material deposited: MLP-He 6065; MLP-He
616-617/C.
Reference: Drago & Lunaschi (2010).
Australapatemon magnacetabulum Dubois,
1988
Natural definitive host: unknown in Argentina.
Experimental definitive hosts: Gallus gallus
(Linnaeus, 1758) (Phasianidae), Anas
platyrhynchos Linnaeus, 1758.
Site of infection: anterior intestine.
Life cycle: Hi : sporocysts and cercaria in
1
gonad and hepatopancreas of Biomphalaria
tenagophila (d'Orbigny, 1835). Hi :
2
metacercariae in parenchyma of Helobdella
adiastola Ringuelet, 1972, Helobdella
triserialis (Blanchard, 1849), Haementeria
e i c h h o rn i a e Ringue let, 197 8, a n d
Haementeria sp. from Tres Palmeras, Valley of
Lerma, Salta Province.
Material deposited: MACN-Pa 521/1-9 from
A. platyrhynchos; MACN-Pa 521/10-19 from
G. g. domesticus; MACN-Pa 521/20
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331
sporocysts from B. tenagophila; MACN-Pa
521/21-24 sagittal sections of adults from G.
domesticus; MACN-Pa 521/25 histological
sections of leeches containing metacercariae.
Reference: Davies & Ostrowski de Núñez
(2012).
Genus Cardiocephaloides Sudarikov, 1959
Cardiocephaloides physalis (Lutz, 1926)
Host: Spheniscus magellanicus (Foster, 1781)
(Spheniscidae).
Site of infection: intestine.
Locality: coast of Península Valdés (42°04'-
42°53'S; 63°38'-64°30'W), Chubut Province.
Material deposited: MLP-He 6085.
Reference: Díaz et al. (2010).
Genus Strigea Abildgaard, 1790
Strigea elliptica (Brandes, 1888) Szidat, 1928
Host: Buteogallus meridionalis (Latham,
1790) (Accipitridae).
Site of infection: small intestine.
Locality: Bellaco wetland (26°17′S,
59°06′W), Formosa Province.
Material deposited: MLP-He 5885.
Reference: Lunaschi & Drago (2009b).
Strigea falconis brasiliana Szidat, 1929
Hosts: Phalacrocorax brasilianus (Gmelin,
1789) (Phalacrocoracidae), Milvago
chimachima (Vieillot, 1816), Caracara
plancus (Miller, 1777) (Falconidae).
Site of infection: small intestine.
Locality: La Marcela farm (26°17'S;
59°06'W), Pirané, Formosa Province.
Material deposited: MLP-He 6415, MLP-He
6712, MLP-He 6713
Reference: Drago et al. (2011, 2014).
Strigea inflecta Lunaschi & Drago, 2012
Host: Cariama cristata (Linnaeus, 1766)
(Cariamidae).
Site of infection: small intestine.
Locality: La Marcela farm (26°17'35''S;
59°06'67''W), Pirané, Formosa Province.
Material deposited: Holotype MLP-He 6445;
paratypes MLP-He 6446.
Reference: Lunaschi & Drago (2012).
Strigea meridionalis Lunaschi & Drago, 2009
Host: Buteogallus meridionalis (Latham,
1790) (Accipitridae).
Site of infection: small intestine.
Locality: Bellaco wetland (26°17′S,
59°06′W), Formosa Province.
Material deposited: Holotype MLP-He 5883;
paratypes MLP-He 5884.
Reference: Lunaschi & Drago (2009b).
Strigea microbursa Pearson & Dubois, 1985
Host: Buteogallus meridionalis (Latham,
1790) (Accipitridae).
Site of infection: small intestine.
Locality: Bellaco wetland (26°17′S,
59°06′W), Formosa Province.
Material deposited: MLP-He 5886.
Reference: Lunaschi & Drago (2009b).
Strigea vaginata (Brandes, 1888) Szidat, 1928
Host: Coragyps atratus (Bechstein, 1793)
(Cathartidae).
Site of infection: intestine.
Locality: La Marcela farm (26°17'S;
59°06'W), Pirané, Formosa Province.
Material deposited: MLP-He 5964.
Reference: Drago & Lunaschi (2011a).
Strigea orbiculata Lunaschi & Drago, 2013
Host: Taraba major (Vieillot, 1816)
(Thamnophilidae).
Site of infection: small intestine.
Locality: La Marcela farm (26°17'35''S;
59°08'38''W), Pirané, Formosa Province.
Material deposited: Holotype MLP-He 6672;
paratypes MLP-He 6673.
Reference: Lunaschi & Drago (2013).
Strigea proteolytica Drago, Lunaschi &
Draghi, 2014
Host: Buteogallus urubitinga (Gmelin, 1788)
(Accipitridae).
Site of infection: small intestine.
Locality: La Marcela farm (26°17'35”S;
59°06'38”W), Pirané, Formosa Province.
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Update of checklist of Digenean parasites of wild birds from Argentina
Material deposited: Holotype MLP-He 6709;
paratypes MLP-He 6710, voucher specimens:
MLP-He 6711.
Reference: Drago et al. (2014).
Genus Parastrigea Szidat, 1928
Comments: The citations of P. brasiliana
reported by Boero et al. (1972a) and Labriola
& Suriano (1998) listed in Lunaschi et al.
(2007) were regarded erroneous (see Drago &
Lunaschi 2011a).
Parastrigea macrobursa Drago & Lunaschi
2011
Host: Buteogallus urubitinga (Gmelin, 1788)
(Accipitridae).
Site of infection: small intestine.
Locality: La Marcela farm (26°17''S;
59°06''W), Pirané, Formosa.
Material deposited: Holotype MLP-He 6272;
paratypes MLP-He 6273, MLP-He 6274;
voucher specimens: MLP-He 6275 (in cross-
sections), MLP-He 6276.
Reference: Drago & Lunaschi (2011b).
Superfamily Echinostomatoidea Looss, 1899
Family Echinostomatidae Looss, 1899
Genus Drepanocephalus Dietz, 1909
Drepanocephalus spathans Dietz, 1909
Host: Phalacrocorax brasilianus (Gmelin,
1789) (Phalacrocoracidae).
Site of infection: intestine.
Localities: Lacombe lagoon (35º49'S;
57º49'W), Lezama, Buenos Aires Province; La
Marcela farm (26°17'S; 59°06'W), Pirané and
Clorinda (25°17'S; 57°43'W), Formosa
Province.
Material deposited: MLP-He 6422, MLP-He
6423.
Reference: Drago et al. (2011).
Genus Edietziana Özdikmen, 2013
Edietziana serrata (Diesing, 1850) Özdikmen,
2013
Host: Aramus guarauna (Linnaeus, 1766)
(Aramidae).
Site of infection: small intestine.
Locality: La Marcela farm (26°17'35”S;
59°08'38”W), Pirané, Formosa Province.
Material deposited: MLP-He 6604.
Reference: Lunaschi et al. (2015).
Genus Echinostoma Rudolphi, 1809
Echinostoma revolutum “group”
Host: Sturnus vulgaris Linnaeus, 1758
(Sturnidae).
Site of infection: intestine.
Locality: Bernal (34°41'′44''S; 58°16'0''W),
Buenos Aires Province.
Material deposited: MLP-He 6732.
Reference: Valente et al. (2014).
Genus Himasthla Dietz, 1909
Himasthla escamosa Díaz & Cremonte, 2004
Host: Larus atlanticus Olrog, 1958 (Laridae).
Site of infection: intestine.
Locality: Bahía Blanca estuary (38°48'S;
62°15'W), Buenos Aires Province.
Material deposited: MLP-He 5735 (indicated
as 5800-5804).
Reference: La Sala et al. (2009).
Host: Larus dominicanus Lichtenstein, 1823
(Laridae).
Site of infection: intestine.
Localities: Fracasso Beach (42°25'S;
64°07'W), San José Gulf, and Puerto Madryn
Beach (42°47'S; 65°02'S), Nuevo Gulf,
Chubut Province.
Material deposited: MLP-He 5233, MLP-He
5234, CNP-Par. 6.
Reference: Díaz et al. (2011).
Genus Lyperorchis Travassos, 1921
Lyperorchis lyperorchis Travassos, 1921
(Syn. Lyperorchis inexpectabilis Digiani,
1997)
Host: Aramus guarauna (Linnaeus, 1766)
(Aramidae).
Site of infection: cloaca.
Localities: Punta Blanca (34°56'S; 57°41'W),
Magdalena, Buenos Aires Province; La
Marcela farm (26°17'35”S; 59°08'38”W),
Pirané, Formosa Province.
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Material deposited: MLP-He 3323/2; MLP-He
6603.
Reference: Lunaschi et al. (2015).
Comments: Lyperorchis inexpectabilis
Digiani, 1997 was listed in Lunaschi et al.
(2007) parasitizing A. guarauna from Buenos
Aires Province. Later, this species was
synonymized with Lyperorchis lyperorchis by
Lunaschi et al. (2015).
Genus Nephrostomum Dietz, 1909
Nephrostomum limai Travassos, 1922
Host: Syrigma sibilatrix (Temminck, 1824)
(Ardeidae).
Site of infection: intestine.
Locality: La Marcela farm (26°17'35”S;
59°08'38”W), Pirané, Formosa Province.
Material deposited: MLP-He 6787.
Reference: Lunaschi et al. (2015).
Genus Paryphostomum Dietz, 1909
Paryphostomum segregatum Dietz, 1909
Host: Coragyps atratus (Bechstein, 1793)
(Cathartidae).
Site of infection: intestine.
Locality: La Marcela farm (26°17'S;
59°06'W), Pirané, Formosa Province.
Material deposited: MLP-He 6010.
Reference: Drago & Lunaschi (2011a).
Host: Phalacrocorax brasilianus (Gmelin,
1789) (Phalacrocoracidae).
Site of infection: intestine.
Localities: Clorinda (25°17'S; 57°43'W),
Formosa Province and Lacombe lagoon
(35º49'S, 57º49'W), Lezama, Buenos Aires
Province.
Material deposited: MLP-He 6419, MLP-He
6420.
Reference: Drago et al. (2011).
Paryphostomum parvicephalum (Rietschel &
Werding, 1978) Kostadinova, Vaucher &
Gibson, 2002
Host: Phalacrocorax brasilianus (Gmelin,
1789) (Phalacrocoracidae).
Site of infection: intestine.
Locality: La Marcela farm (26°17'S;
59°06'W), Pirané, Formosa Province.
Material deposited: MLP-He 6421.
Reference: Drago et al. (2011).
Genus Petasiger Dietz, 1909
Petasiger argentinensis Lunaschi & Drago
2010
Hosts: Podiceps major (Boddaert, 1783);
Rollandia rolland (Quoy & Gaimard, 1824)
(Podicipedidae).
Site of infection: small intestine.
Locality: Lacombe lagoon (35°49΄S;
57°49΄W), Lezama, Buenos Aires.
Material deposited: Holotype MLP-He 6061;
paratypes MLP-He 6062; voucher specimens
MLP-He 6063; MLP-He 6064.
Reference: Lunaschi & Drago (2010).
Petasiger sp.
Host: Ardea cocoi Linnaeus, 1766 (Ardeidae).
Site of infection: intestine.
Lacombe lagoon (35º49'S, 57º49'W), Lezama,
Buenos Aires Province.
Material deposited: MLP-He 6012.
Reference: Drago & Lunaschi (2011a).
Petasiger sp.
Host: Coragyps atratus (Bechstein, 1793)
(Cathartidae).
Site of infection: intestine.
Locality: La Marcela farm (26°17'S;
59°06'W), Pirané, Formosa Province.
Material deposited: MLP-He 6011.
Reference: Drago & Lunaschi (2011a).
Genus Stephanoprora Odhner, 1902
Stephanoprora aylacostoma Ostrowski de
Núñez & Quintana, 2008
Natural definitive host: unknown.
Experimental definitive host: Gallus gallus
(Linnaeus, 1758) (Phasianidae).
Site of infection: posterior intestine.
Life cycle: Hi : sporocysts, rediae and
1
cercariae in gills of Aylacostoma chloroticum
Hylton Scott, 1954 (natural host). Hi :
2
metacercariae encysted on gill filaments and
gill chamber of Moenkhausia dichroura (Kner,
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Update of checklist of Digenean parasites of wild birds from Argentina
1858), Astyanax erythropterus (Holmberg,
1891) and Hyphesobrycon serpae (Durbin in
Eigenmann 1908) (natural hosts), and
Cnesterodon decemmaculatus (Jenyns, 1842)
and Poecilia reticulata Peters, 1859
(experimental hosts).
Locality: Paraná River in Heller Peninsula
(27°20'S; 55°55'W), Misiones Province.
Material deposited: MACN-Pa 441/1-6.
Reference: Ostrowski de Núñez & Quintana
(2008).
Stephanoprora podicipei Etchegoin &
Martorelli, 1997
Host: Larus dominicanus Lichtenstein, 1823
(Laridae).
Site of infection: intestine.
Localities: Fracasso Beach (42°25'S;
64°07'W), San José Gulf, and Puerto Madryn
Beach (42°47'S; 65°02'S), Nuevo Gulf,
Chubut Province.
Material deposited: MLP-He 6307; CNP-Par
9.
Reference: Díaz et al. (2011).
Comments: Stephanoprora podicipei was
listed as synonymous of Stephanoprora
uruguayense Holcman-Spector & Olagüe,
1989 in Lunaschi et al. (2007). Díaz et al.
(2011) regarded to S. podicipei as a valid
species, through a personal communication of
Dr. Ostrowski de Núñez.
Stephanoprora uruguayense Holcman-
Spector & Olagüe, 1989
Natural definitive hosts: see Lunaschi et al.
(2007).
Experimental definitive hosts: Gallus gallus
(Linnaeus, 1758) (Phasianidae), Mus musculus
Linnaeus, 1758 (Muridae).
Site of infection: posterior intestine.
Material deposited: MACN-Pa 440/1-4 from
G. g. domesticus, 440/5 from M. musculus.
Life cycle: Hi : sporocysts in gills, rediae and
1
cercariae in hepatopancreas and gonads of
Heleobia parchappei (d'Orbigny, 1835) from
Buenos Aires City, rediae and cercaria in
digestive gland and gonad of Heleobia
australis (d'Orbigny, 1835) from Buenos Aires
Province. Hi : metacercariae in gills of
2
Cnesterodon decemmaculatus (Jenyns, 1842)
from Buenos Aires City, Galaxias maculatus
(Jenyns, 1842) and Aplochiton zebra Jenyns,
1842 from Patagonia.
References: Ostrowski de Núñez (2007),
Viozzi et al. (2009), Fernández et al. (2010,
2012), Alda & Martorelli (2014).
Family Philophthalmidae Travassos, 1918
Genus Parorchis Nicoll, 1907
Parorchis sp.
Host: Larus dominicanus Lichtenstein, 1823
(Laridae).
Site of infection: cloaca, rectum, bursa
Fabricii.
Localities: Fracasso Beach (42°25'S;
64°07'W), San José Gulf and Puerto Madryn
Beach (42°47'S; 65°02' S), Nuevo Gulf,
Chubut Province.
Material deposited: MLP-He 6308, CNP-Par
15.
Reference: Díaz et al. (2011).
Family Psilostomidae Looss, 1900
Genus Psilochasmus Lühe, 1909
Psilochasmus oxyurus (Creplin, 1825) Lühe,
1909
Host: Himantopus melanurus Vieillot, 1817
(Recurvirostridae).
Site of infection: intestine.
Locality: Bahía Blanca estuary (38º49'S;
62º06'W), Buenos Aires Province.
Material deposited: MLP-He 5984.
Life cycle: Hi and Hi rediae and
1 2 :
metacercariae in Heleobia australis
(d'Orbigny, 1835) from Bahía Blanca estuary.
References: Alda et al. (2011a), Alda &
Martorelli (2014).
Genus Ribeiroia Travassos, 1939
Ribeiroia ondatrae (Price, 1932) Price, 1942
Host: Phalacrocorax brasilianus (Gmelin,
1789) (Phalacrocoracidae).
Site of infection: stomach.
Localities: Lacombe lagoon (35º49'S;
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57º49'W), Lezama, Buenos Aires Province; La
Marcela farm (26°17'S; 59°06'W), Pirané and
Clorinda (25°17'S; 57°43'W), Formosa
Province.
Material deposited: MLP-He 6416, MLP-He
6417.
Reference: Drago et al. (2011).
Superfamily Gorgoderoidea Looss, 1899
Family Dicrocoeliidae Looss, 1899
Genus Athesmia Looss, 1899
Athesmia heterolecithodes (Braun, 1899)
Looss, 1899
Host: Nothura maculosa (Temminck, 1815)
(Tinamidae)
Site of infection: not reported
Locality: San Jaime de la Frontera (Juan B.
Arruabarrena), Entre Rios Province.
Material deposited: not reported.
Reference: Byrd et al. (1967), Bump & Bump
(1969).
Host: Nothura darwinii Gray, 1867
(Tinamidae).
Site of infection: not reported.
Locality: Uriburu, La Pampa Province.
Material deposited: not reported
Reference: Byrd et al. (1967).
Hosts: Cariama cristata (Linnaeus, 1766)
(Cariamidae); Guira guira (Gmelin, 1788)
(Cuculidae); Milvago chimachima (Vieillot,
1816) (Falconidae); Rostrhamus sociabilis
(Vieillot, 1817) (Accipitridae); Theristicus
c a u d a t u s ( B o d d a e r t , 1 7 8 3 )
(Threskiornithidae).
Site of infection: liver bile ducts.
Locality: La Marcela farm (26°17'35”S;
59°08'38”W), Pirané, Formosa Province.
Material deposited: MLP-He 5576, MLP-He
6788; MLP-He 6789.
References: Lunaschi & Drago (2009a),
Lunaschi et al. (2015).
Genus Lubens Travassos, 1919
Lubens lubens (Braun, 1901) Strom, 1940
Host: Taraba major (Vieillot, 1816)
(Thamnophilidae).
Site of infection: intestine.
Locality: La Marcela farm (26°17'35''S;
59°08'38''W), Pirané, Formosa Province.
Material deposited: MLP-He 6730.
Reference: Lunaschi et al. (2014).
Genus Lyperosomum Looss, 1919
Lyperosomum oswaldoi (Travassos, 1919)
Travassos, 1944
Host: Taraba major (Vieillot, 1816)
(Thamnophilidae).
Site of infection: bile ducts.
Locality: La Marcela farm (26°17'35''S;
59°08'38''W), Pirané, Formosa Province.
Material deposited: MLP-He 6674.
Comments: This species was reported by
Boero et al. (1972b) parasitizing Molothrus
bonariensis (Gmelin, 1789) (Passeriformes,
Icteridae) (Lunaschi et al. 2007). Later, this
report was considered erroneous by Lunaschi
& Drago (2013).
Reference: Lunaschi & Drago (2013).
Lyperosomum sp.
Host: Larus dominicanus Lichtenstein, 1823
(Laridae).
Site of infection: pancreas.
Localities: Fracasso Beach (42°25'S;
64°07'W), San José Gulf and Puerto Madryn
Beach (42°47'S; 65°02'S), Nuevo Gulf,
Chubut Province.
Material deposited: MLP-He 6306, CNP-Par
16.
Reference: Díaz et al. (2011).
Genus Zonorchis Travassos, 1944
Zonorchis sp.
Host: Nothura maculosa (Temminck, 1815)
(Tinamidae)
Site of infection: liver.
Locality: not reported.
Material deposited: not reported.
Reference: Bump & Bump (1969).
Superfamily Gymnophalloidea Odhner, 1905
Family Gymnophallidae Odhner, 1905
Genus Bartolius Cremonte, 2001
Bartolius pierrei Cremonte, 2001
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Host: Larus dominicanus Lichtenstein, 1823
(Laridae).
Site of infection: intestine.
Localities: Fracasso Beach (42°25'S;
64°07'W), San José Gulf; Puerto Madryn
Beach (42°47'S; 65°02'S), Nuevo Gulf,
Chubut Province.
Material deposited: MLP-He 6310; CNP-Par
12.
Reference: Díaz et al. (2011).
Genus Gymnophallus Odhner, 1900
Gymnophallus australis Szidat, 1962
Host: Larus dominicanus Lichtenstein, 1823
(Laridae).
Site of infection: intestine.
Localities: Fracasso Beach (42°25'S;
64°07'W), San José Gulf, and Puerto Madryn
Beach (42°47'S; 65°02'S), Nuevo Gulf,
Chubut Province.
Life cycle: Hi : metacercariae in central
2
extrapallial space, between the valve and
mantle of Perumytilus purpuratus (Lamarck,
1819), Mytilus edulis (Linnaeus, 1758) and
Aulacomya atra (Molina, 1782) from
Comodoro Rivadavia, Chubut Province.
Material deposited: adults from natural host
MLP-He 6311 and CNP-Par 14. Experimental
adults: MACN-Pa 437/1-4 and BMNH
2007.2.21.17. Metacercariae: MACN-Pa
438/1-4 and BMNH 2007.2.21.18.
References: Cremonte et al. (2008), Díaz et al.
(2011).
Superfamily Microphalloidea Ward, 1901
Family Microphallidae Ward, 1901
Genus Levinseniella Stiles & Hassall, 1901, in
Ward (1901)
Levinseniella cruzi Travassos, 1920
Hosts: Larus atlanticus Olrog, 1958 (Laridae);
Himantopus melanurus Vieillot, 1817
(Recurvirostridae).
Site of infection: intestine.
Locality: Bahía Blanca estuary, (38°48'S;
62°15'W; 38º49'S; 62º06'W) Buenos Aires
Province.
Life cycle: Hi1 sporocysts in digestive gland
:
and gonad of Heleobia australis (d'Orbigny,
1835) from Bahía Blanca estuary; Hi :
2
metacercariae in gonads of Cyrtograpsus
angulatus Dana, 1851 and Neohelice
granulata (Dana, 1851) from Bahía Blanca
estuary.
Material deposited: MLP-He 5734 (indicated
as 5800-5804); MLP-He 5986.
Reference: La Sala et al. (2009), Alda et al.
(2011a-b), Alda & Martorelli (2014).
Genus Maritrema Nicoll, 1907
Maritrema bonaerense Etchegoin &
Martorelli, 1997
Host: Larus atlanticus Olrog, 1958 (Laridae).
Experimental host: Gallus gallus (Linnaeus,
1758) (Phasianidae).
Site of infection: intestine.
Locality: Bahía Blanca estuary (38°48'S;
62°15'W), Buenos Aires Province.
Life cycle: Hi : sporocysts and cercariae in
1
Heleobia australis (d'Orbigny, 1835) from
Bahía Blanca estuary; Hi : metacercariae in
2
gills, haemocoel, muscle and ovary of
Cyrtograpsus angulatus Dana, 1851 and
Neohelice granulata (Dana, 1851) from Bahía
Blanca estuary.
Material deposited: Adult from natural host
MLP-He 5731 (indicated as 5800-5804);
sporocysts and cercariae MLP-He 6525;
metacercariae MLP-He 6526; experimental
adult MLP-He 6527.
References: La Sala et al. (2009), Alda et al.
(2011b, 2013), Alda & Martorelli (2014).
Maritrema formicae Díaz, Gilardoni &
Cremonte, 2012
Host: Larus dominicanus Lichtenstein, 1823
(Laridae).
Site of infection: small intestine.
Locality: Fracasso Beach (42°25'S; 64°07'W),
Peninsula Valdes, Chubut Province.
Material deposited: Holotype MLP-He 6501,
paratypes MLP-He 6502, voucher specimens
MLP-He 6503; CNP-Par 13.
Comments: These specimens were reported as
Maritrema sp. by Díaz et al. (2011).
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Reference: Díaz et al. (2011, 2012).
Maritrema madrynense Díaz & Cremonte,
2010
Host: Larus dominicanus Lichtenstein, 1823
(Laridae).
Site of infection: small intestine.
Locality: Puerto Madryn coast (42°46'S,
65°02'W), Fracasso Beach (42°25'S;
64°07'W), San José Gulf, and Puerto Madryn
Beach (42°47'S; 65°02'S), Nuevo Gulf,
Chubut Province.
Material deposited: Holotype MLP-He 6087,
paratypes MLP-He 6088, voucher: specimens
MLP-He 6089, 6090 (metacercariae and
experimentally obtained adults).
Life cycle: Hi : metacercariae in body cavity of
2
Cyrtograpsus altimanus Rathbun, 1914 from
Puerto Madryn coast.
References: Díaz & Cremonte (2010), Díaz et
al. (2011).
Maritrema orensense Cremonte & Martorelli,
1998
Host: Larus atlanticus Olrog, 1958 (Laridae).
Experimental definitive host: Gallus gallus
(Linnaeus, 1758) (Phasianidae).
Site of infection: intestine.
Locality: Bahía Blanca estuary (38°48'S;
62°15'W), Buenos Aires Province.
Life cycle: Hi : sporocysts and cercariae in
1
Heleobia australis (d'Orbigny, 1835) from
Bahía Blanca estuary. Hi : metacercariae in
2
gills of Cyrtograpsus angulatus Dana, 1851
and Neohelice granulata (Dana, 1851) from
Bahía Blanca estuary.
Material deposited: Adult from natural host:
5732 (indicated as 5800-5804); experimental
adult MLP-He 6524; cercariae MLP-He 6522;
metacercariae MLP-He 6198, MLP-He 6523.
References: La Sala et al. (2009), Alda et al.
(2011b, 2013), Alda & Martorelli (2014).
Maritrema patagonica Rauque, Flores &
Brugni, 2013
Natural definitive host: unknown.
Experimental definitive host: Gallus gallus
(Linnaeus, 1758) (Phasianidae).
Site of infection: intestine.
Life cycle: Hi : metacercariae encysted in gill
2
axis of Aegla neuquensis Schmitt, 1942 and
Aegla riolimayana Schmitt, 1942.
Locality: Aluminé River, 38°55'S; 71°10'W
(Type locality), Limay River, Rucachoroi
Lake, Caleufu River, Nahuel Huapi Lake,
Comallo River, Ñireco River (Patagonia).
Material deposited: MACN-Pa 557/1,
Holotype from experimental infection in
Gallus gallus; MACN-Pa 557/2, Paratype
from G. g. domesticus; MLP-He 6657,
Paratypes from G. g. domesticus; UNCo-Pa
48/1-2; Paratypes from G. g. domesticus;
MACN-Pa 557/3a, 557/3b, 557/4, 557/5,
Voucher specimens; MLP-He 6658 - 6660
Voucher specimens; UNCo-Pa 217/1, . 218/1-
4, 220/1, voucher specimens.
Reference: Rauque et al. (2013).
Genus Megalophallus Cable, Connor &
Balling 1960
Megalophallus deblocki Kostadinova,
Vaucher & Gibson, 2006
Hosts: Rostrhamus sociabilis (Vieillot, 1817);
Buteogallus meridionalis (Latham, 1790)
(Accipitridae).
Site of infection: intestine.
Locality: La Marcela farm (26°17'35”S;
59°06'38”W), Pirané, Formosa Province.
Material deposited: MLP-He 6720, MLP-He
6721.
Reference: Drago et al. (2014).
Microphallus Ward, 1901
Microphallus simillimus (Travassos, 1920)
Martorelli, 1991
Natural definitive hosts: see Lunaschi et al.
(2007).
Life cycle: Hi and Hi : sporocysts and
1 2
metacercariae in gonad and digestive gland of
Heleobia australis (d'Orbigny, 1835) from
Bahía Blanca estuary, Buenos Aires Province.
Reference: Alda et al. (2010), Alda et al.
(2011c), Alda & Martorelli (2014).
Genus Odhneria Travassos, 1921
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Odhneria odhneri Travassos, 1921
Host: Larus atlanticus Olrog, 1958 (Laridae).
Site of infection: intestine.
Locality: Bahía Blanca estuary (38°48'S;
62°15'W), Buenos Aires Province.
Material deposited: MLP-He 5733 (indicated
as 5800-5804).
Reference: La Sala et al. (2009).
Host: Larus dominicanus Lichtenstein, 1823
(Laridae).
Site of infection: intestine.
Localities: Fracasso Beach (42°25'S;
64°07'W), San José Gulf and Puerto Madryn
Beach (42°47'S; 65°02'S), Nuevo Gulf,
Chubut Province.
Material deposited: MLP-He 6309, CNP-Par
10.
Reference: Díaz et al. (2011).
Family Prosthogonimidae Lühe, 1909
Genus Prosthogonimus Lühe, 1899
Prosthogonimus ovatus (Rudolphi, 1803)
Lühe, 1899
Host: Phalacrocorax brasilianus (Gmelin,
1789) (Phalacrocoracidae).
Site of infection: bursa Fabricii.
Locality: Lacombe lagoon (35º49'S;
57º49'W), Lezama, Buenos Aires Province.
Material deposited: MLP-He 6418.
Reference: Drago et al. (2011).
Family Stomylotrematidae Poche, 1926
Genus Stomylotrema Looss, 1900
Stomylotrema vicarium Braun, 1901
Hosts: Busarellus nigricollis (Latham, 1790);
Buteogallus meridionalis (Latham, 1790)
(Accipitridae); Theristicus caudatus
(Boddaert, 1783) (Threskiornithidae).
Site of infection: cloaca.
Locality: La Marcela farm (26°17'35”S;
59°06'-59°08'W).
Material deposited: MLP-He 5743, MLP-He
5577, MLP-He 6790.
References: Lunaschi & Drago (2009a),
Lunaschi et al (2015).
Superfamily Ophistorchioidea Looss, 1899
Family Heterophyidae Leiper, 1909
Genus Ascocotyle Looss, 1899
Ascocotyle diminuta Stunkard & Haviland,
1924
Host: Ardea alba Linnaeus, 1758 (Ardeidae).
Site of infection: intestine.
Locality: Lacombe lagoon (35º49'S;
57º49'W), Lezama, Buenos Aires Province.
Material deposited: MLP-He 5976.
Reference: Drago & Lunaschi (2011a).
Ascocotyle (Ascocotyle) felippei Travassos,
1928
Host: Himantopus melanurus Vieillot, 1817
(Recurvirostridae).
Site of infection: caeca.
Locality: Bahía Blanca estuary, (38º49'S;
62º06'W) Buenos Aires Province.
Material deposited: MLP-He 5985.
Reference: Alda et al. (2011a).
Genus Pygidiopsis Looss, 1907
Pygidiopsis crassus Ostrowski de Núñez,
1995
Host: Himantopus melanurus Vieillot, 1817
(Recurvirostridae).
Site of infection: small intestine.
Locality: Bahía Blanca estuary, Buenos Aires
Province.
Material deposited: MPL He 5983.
Reference: Alda et al. (2011a).
Superfamily Paramphistomoidea Fischoeder,
1901
Family Zygocotylidae Ward, 1917
Genus Zygocotyle Stunkard, 1917
Zygocotyle lunata (Diesing, 1836) Stunkard,
1917
Natural hosts: see Lunaschi et al. (2007).
Experimental hosts: Gallus gallus (Linnaeus,
1758) (Phasianidae), Mus musculus Linnaeus,
1758 (Muridae).
Life cycle: Hi : sporocysts, rediae and
1
cercariae in Biomphalaria peregrina
(d'Orbigny, 1835) from Buenos Aires City and
Biomphalaria tenagophila (d'Orbigny, 1835)
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from Lerma Valley, Salta Province (natural
hosts), and Biomphalaria straminea (Dunker,
1848) (experimental host). Metacercariae
encyst on the substrate.
References: Ostrowski de Núñez et al. (2011).
Superfamily Pronocephaloidea Looss, 1899
Family Notocotylidae Lühe, 1909
Genus Catatropis Odhner, 1905
Catatropis chilinae Flores & Brugni, 2003
Natural host: unknown.
Experimental hosts: see Lunaschi et al. (2007).
Life history: Hi : rediae in Chilina dombeiana
1
(Bruiguiere, 1789) from Mascardi Lake, Río
Negro Province.
Reference: Flores & Semenas (2008).
Genus Notocotylus Diesing, 1839
Notocotylus biomphalariae Flores & Brugni,
2005
Natural host: unknown.
Experimental hosts: see Lunaschi et al. (2007).
Life history: Hi : rediae in Biomphalaria
1
peregrina (d'Orbigny, 1835) from Fantasma
pond, Patagonia.
Reference: Flores et al. (2010).
Inventory of digenean parasites of wild birds
from Argentina
The inventory work of digeneans from birds in
Argentina began in 1909 with few and isolated
reports on their taxonomy. It was not until 4
decades later that the formal survey work
started with Lothar Szidat. In the 1970s,
Argentinean parasitologists described a
considerable number of digenean species
parasitizing birds, in separate and isolated
reports, mostly new species from particular
host species collected in specific localities. In
the following two decades the number of
records decreases, however since 1996 to
present it had a significant increase, reaching
112 species including studies related to
taxonomic and ecological aspects. However,
the species accumulation curve shows that the
asymptote has not been reached yet (Fig. 1).
The most represented families are
Ec hi n os to ma t id ae , S t ri ge id a e a nd
Diplostomidae, with 26, 18 and 15 species,
respectively (Fig. 2).
Host-parasite relations
Neotropical Helminthology. Vol. 9, Nº2, jul-dec 2015
Update of checklist of Digenean parasites of wild birds from Argentina
Figure 1. Species accumulation curve of digenean species from wild birds in Argentina from the decade of the 1900s to the
present.
Of the almost 1000 bird species known in
Argentina, only 70 (7%) has been reported as
definitive natural hosts of digenean. The
families of birds more studied are Laridae,
Ardeidae and Accipitridae, with 32, 29 and 17
digenean species reported.
The accumulation curve of host-paraste
relations (digenean species and wild bird
species) is similar to the accumulation species
curve up to year 2000, because in most of the
papers the authors describe each species
parasitizing a particular host species collected
in specific localities. The increase of papers on
ecology and biogeography of digeneans over
the last two decades cause a rise in reports on
host-parasite relations, reaching 190 reports.
However, the host-parasite relations
accumulation curve shows that the asymptote
has not been reached yet (Fig. 3).
Ecoregions richness
340
Neotropical Helminthology. Vol. 9, Nº2, jul-dec 2015 Drago & Lunaschi
Figure 2. Number of species per family reported in Argentina.
Figure 3. Accumulation curve of digenean species and wild birds relations in Argentina from the decade of the 1900s to the
present.
341
Among the 15 continental ecoregions
recognized in Argentina, there are reports of
digeneans from birds in 7 of them, and a single
report in the South Atlantic Islands ecoregion
(Fig. 4). The highest richness of species and
host-parasite relations were reported in the
northeastern and the central part of the country,
mainly in the Humid Chaco, Pampa and Paraná
Flooded Savanna ecoregions. The Espinal,
Patagonian Steppe and Monte of Plains and
Plateaus ecoregions have low species richness
(Fig. 5).
Figure 4. Map of the ecoregions of Argentina indicating the number of digenean species recorded in wild birds.
Neotropical Helminthology. Vol. 9, Nº2, jul-dec 2015
Update of checklist of Digenean parasites of wild birds from Argentina
342
Neotropical Helminthology. Vol. 9, Nº2, jul-dec 2015 Drago & Lunaschi
Figure 5. Number of species and host-parasite relations in each ecoregion of Argentina.
Wild Bird Hosts
Order/Family
Species
Digenean species
References
Accipitriformes
Accipitridae
Busarellus
nigricollis
Posthodiplostomum macrocotyle
Stomylotrema vicarium
Tylodelphys brevis
Drago et al. (2014)
Lunaschi & Drago (2009a)
Drago et al. (2014)
Buteogallus meridionalis
Megalophallus deblocki
Neodiplostomum travassosi
Stomylotrema vicarium
Strigea elliptica
Strigea meridionalis
Strigea microbursa
Drago et al. (2014)
Drago et al. (2014)
Lunaschi & Drago (2009a)
Lunaschi & Drago (2009b)
Lunaschi & Drago (2009b)
Lunaschi & Drago (2009b)
Buteogallus
urubitinga
Parastrigea
macrobursa
Strigea
proteolytica
Drago & Lunaschi (2011b)
Drago et al. (2014)
Rostrhamus
sociabilis
Athesmia
heterolecithodes
Megalophallus
deblocki
Spaniometra
variolaris
Lunaschi & Drago (2009a)
Drago et al. (2014)
Drago et al. (2014)
Anseriformes
Anatidae
Cygnus
melancoryphus
Australapatemon
canadensis
Drago & Lunaschi (2010)
Cariamiformes
Cariamidae
Cariama
cristata
Athesmia
heterolecithodes
Brachylaima
yupanquii
Strigea
inflecta
Lunaschi et al. (2015)
Lunaschi & Drago (2012)
Lunaschi & Drago (2012)
Cathartiformes
Cathartidae Coragyps atratus Paryphostomum segregatum
Petasiger sp.
Strigea vaginata
Drago & Lunaschi (2011a)
Drago & Lunaschi (2011a)
Drago & Lunaschi (2011a)
Table 1. Host-parasite list since Lunaschi et al. (2007) up to present from Argentina.
Cont. Table 1
Ciconiiformes
Ciconiidae
Mycteria
americana
Tylodelphys
brevis
Drago & Lunaschi (2008)
Charadriiformes
Laridae
Larus
atlanticus
Himasthla
escamosa
Levinseniella
cruzi
Maritrema
bonaerense
Maritrema
orensensis
Odhneria
odhneri
La Sala et
al. (2009)
La Sala et
al. (2009)
La Sala et
al. (2009)
La Sala et
al. (2009)
La Sala et
al. (2009)
Larus
dominicanus
Bartolius
pierrei
Gymnophallus
australis
Himasthla
escamosa
Lyperosomum
sp.
Maritrema
formicae
Maritrema
madrynense
Odhneria
odhneri
Parorchis
sp.
Stephanoprora
podicipei
Díaz et
al. (2011)
Díaz et
al. (2011)
Díaz et
al. (2011)
Díaz et
al. (2011)
Díaz et
al. (2011, 2012).
Díaz & Cremonte (2010), Díaz et al. (2011)
Díaz et
al. (2011)
Díaz et
al. (2011)
Díaz et
al. (2011)
Recurvirostridae
Himantopus
melanurus
Ascocotyle
(A.) felippei
Levinseniella
cruzi
Psilochasmus
oxyurus
Pygidiopsis
crassus
Tanaisia
dubia
Alda et
al. (2011a)
Alda et
al. (2011a)
Alda et
al. (2011a)
Alda et
al. (2011a)
Lunaschi et
al. (in press).
Cuculiformes
Cuculidae
Guira
guira
Athesmia
heterolecithodes
Glaphyrostomum
propinquum
Lunaschi & Drago (2009a )
Lunaschi & Drago (2009a)
Falconiformes
Falconidae
Caracara
plancus
Strigea
falconis
brasiliana
Drago et
al. (2014)
Milvago
chimachima
Athesmia
heterolecithodes
Strigea
falconis
brasiliana
Lunaschi & Drago (2009a)
Drago et
al. (2014)
Gruiformes
Aramidae
Aramus
guarauna
Edietziana
serrata
Lyperorchis
lyperorchis
Lunaschi et
al. (2015)
Lunaschi et
al. (2015)
Passeriformes
Sturnidae
Sturnus
vulgaris
Echinostoma
revolutum
“group”
Valente et al. (2014)
Thamnophilidae
Taraba
major
Lubens
lubens
Lyperosomum
oswaldoi
Pojmanskia
riosae
Strigea
orbiculata
Lunaschi et
al. (2014)
Lunaschi & Drago (2013)
Lunaschi et
al. (2014)
Lunaschi & Drago (2013).
Tyrannidae
Pitangus
sulphuratus
Tanaisia
dubia
Lunaschi et
al. (in press)
Pelecaniformes
Ardeidae
Ardea
alba
Apharyngostrigea
ardearum
Ascocotyle
diminuta
Dolichorchis
lacombeensis
Posthodiplostomum
nanum
Drago & Lunaschi (2011a)
Drago & Lunaschi (2011a)
Drago & Lunaschi (2011a)
Drago & Lunaschi (2011a).
Ardea cocoi Apharyngostrigea ardearum
Dolichorchis lacombeensis
Petasiger sp
Posthodiplostomum sp..
Drago & Lunaschi (2011a)
Drago & Lunaschi (2011a)
Drago & Lunaschi (2011a)
Drago & Lunaschi (2011a)
Bubulcus ibis Apharyngostrigea ardearum Drago & Lunaschi (2011a)
Syrigma sibilatrix Nephrostomum limai Lunaschi et al. (2015)
Tigrisoma lineatum Clinostomatopsis sorbens
Clinostomum marginatum
Posthodiplostomum nanum
Sphincterodiplostomum musculosum
Lunaschi & Drago (2009a)
Lunaschi & Drago (2009a)
Drago & Lunaschi (2011a)
Drago & Lunaschi (2011a)
343
Tylodelphys elongata Drago & Lunaschi (2011a)
Cont. Table 1
Neotropical Helminthology. Vol. 9, Nº2, jul-dec 2015
Update of checklist of Digenean parasites of wild birds from Argentina
Threskiornithidae
Theristicus
caudatus
Athesmia
heterolecithodes
Stomylotrema
vicarium
Lunaschi et al. (2015)
Lunaschi et al. (2015)
Podicipediformes
Podicipedidae
Podiceps
major
Petasiger
argentinensis
Lunaschi & Drago (2010).
Rollandia
rolland
Petasiger
argentinensis
Lunaschi & Drago (2010)
Sphenisciformes
Spheniscidae
Spheniscus magellanicus
Cardiocephaloides
physalis
Díaz et
al. (2010).
Suliformes
Phalacrocoracidae Phalacrocorax brasilianus Austrodiplostomum mordax
Austrodiplostomum
ostrowskiae
Drepanocephalus
spathans
Hysteromorpha
triloba
Paryphostomum
parvicephalum
Paryphostomum
segregatum
Prosthogonimus
ovatus
Ribeiroia
ondatrae
Strigea
falconis
brasiliana
Tylodelphys
adulta
Drago et al. (2011)
Drago et al. (2011)
Drago et
al. (2011)
Drago et
al. (2011)
Drago et
al. (2011)
Drago et
al. (2011)
Drago et
al. (2011)
Drago et
al. (2011)
Drago et al. (2011)
Drago et
al. (2011)
Tinamiformes
Tinamidae
Nothura
darwinii
Athesmia
heterolecithodes
Byrd et
al. (1967)
Nothura
maculosa
Athesmia
heterolecithodes
Zonorchis
sp.
Byrd et
al. (1967)
Bump & Bump (1969)
Experimental Bird Hosts
Galliformes
Phasianidae Gallus gallus Australapatemon magnacetabulum
Davies & Ostrowski de Núñez (2012)
Maritrema
bonaerense
Maritrema
orensensis
Maritrema patagonica
Posthodiplostomum
sp.
Stephanoprora
aylacostoma
Stephanoprora
uruguayense
Zygocotyle
lunata
Alda et
al. (2013)
Alda et
al. (2013)
Rauque et
al. (2013)
Ritossa et
al. (2013)
Ostrowski de Núñez & Quintana (2008)
Ostrowski de Núñez (2007)
Ostrowski de
Núñez et al. (2011)
Anseriformes
Anatidae
Anas
platyrhynchos
Australapatemon
magnacetabulum
Davies & Ostrowski de Núñez (2012)
Experimental Mammal Hosts
Rodentia
Muridae
Mus
musculus
Posthodiplostomum
sp.
Stephanoprora
uruguayense
Zygocotyle
lunata
Ritossa et
al. (2013)
Ostrowski de Núñez (2007)
Ostrowski de Núñez et
al. (2011)
Intermediate Hosts
Class/Family Species Digenean species References
Gastropoda
Chilinidae Chilina dombeiana Catatropis chilinae Flores & Semenas (2008).
Cochliopidae Heleobia australis Levinseniella cruzi
Maritrema bonaerense
Maritrema orensensis
Microphallus simillimus
Psilochasmus oxyurus
Stephanoprora uruguayense
Alda & Martorelli (2014)
Alda et al. (2013), Alda & Martorelli (2014)
Alda et al. (2013), Alda & Martorelli (2014)
Alda & Martorelli (2014)
Alda & Martorelli (2014)
Alda & Martorelli (2014)
Heleobia parchappei Stephanoprora uruguayense Ostrowski de Núñez (2007)
Planorbidae Anisancylus obliquus Posthodiplostomum sp. Ritossa et al. (2013)
Biomphalaria peregrina Notocotylus biomphalariae
Zygocotyle lunata
Flores et al. (2010).
Ostrowski de Núñez et al. (2011)
Biomphalaria straminea Zygocotyle lunata
Ostrowski de Núñez et al. (2011).
Biomphalaria tenagophila Australapatemon magnacetabulum
Zygocotyle lunata
Davies & Ostrowski de Núñez (2012)
Ostrowski de Núñez et al. (2011)
Thiaridae Aylacostoma chloroticum Stephanoprora aylacostoma
Ostrowski de Núñez & Quintana (2008)
344
Neotropical Helminthology. Vol. 9, Nº2, jul-dec 2015 Drago & Lunaschi
Cont. Table 1
345
diplostomids, six species of the genus
Tylodelphys were described based on
metacercariae found parasitizing the brain, the
pericardial cavity or the visceral cavity of
freshwater fishes: Tylodelphys destructor
Szidat & Nani, 1951, Tylodelphys jenynsiae
Szidat, 1969 and Tylodelphys cardiophilus
Szidat, 1969, Tylodelphys barilochensis
Quaggiotto & Valverde, 1992, Tylodelphys
crubensis Quaggiotto & Valverde, 1992,
Tylodelphys argentinus Quaggiotto &
Valverde, 1992 (Szidat & Nani, 1951; Szidat,
1969; Quaggiotto & Valverde, 1992).
Nevertheless, only three species of adult are
known, T. adulta, T. brevis and T. elongata
(Lunaschi & Drago, 2004; Drago & Lunaschi,
2008, 2011a). Other diplostomids, such as
Lophosicyadiplostomum nephrocystis (Lutz,
1928) reported in anurans (Hamann &
Bivalvia
Mytilidae
Aulacomya
atra
Mytilus
edulis
Perumytilus
purpuratus
Gymnophallus
australis
Gymnophallus
australis
Gymnophallus
australis
Cremonte
et
al. (2008)
Cremonte
et
al. (2008)
Cremonte
et
al. (2008)
Hirudinea
Glossiphoniidae
Haementeria
eichhorniae
Helobdella
adiastola
Helobdella. triserialis
Australapatemon
magnacetabulum
Australapatemon
magnacetabulum
Australapatemon
magnacetabulum
Davies & Ostrowski de Núñez (2012)
Davies & Ostrowski de Núñez (2012)
Davies & Ostrowski de Núñez (2012)
Malacostraca
Aeglidae
Aegla
neuquensis
Aegla
riolimayana
Maritrema
patagonica
Maritrema
patagonica
Rauque et
al. (2013)
Rauque et
al. (2013).
Varunidae
Cyrtograpsus
altimanus
Maritrema
madrynense
Díaz & Cremonte (2010)
Cyrtograpsus
angulatus
Levinseniella
cruzi
Maritrema
bonaerense
Maritrema
orensensis
Alda et
al. (2011b)
Alda et
al. (2011b), Alda et al. (2013)
Alda et
al. (2011b), Alda et al. (2013)
Neohelice
granulata
Levinseniella
cruzi
Maritrema bonaerense
Maritrema orensensis
Alda et
al. (2011b)
Alda et al. (2011b), Alda et al. (2013)
Alda et al. (2011b), Alda et al. (2013)
Actinopterygii
Atherinopsidae Odontesthes bonariensis Austrodiplostomum cf. mordax Drago (2012)
Characidae Astyanax erythropterus
Hyphesobrycon serpae
Stephanoprora aylacostoma
Stephanoprora aylacostoma
Ostrowski de Núñez & Quintana (2008)
Ostrowski de Núñez & Quintana (2008)
Moenckhausia dichroura Stephanoprora aylacostoma Ostrowski de Núñez & Quintana (2008)
Galaxiidae Aplochiton zebra Stephanoprora uruguayense Fernández et al. (2012)
Galaxias maculatus Posthodiplostomum sp.
Stephanoprora uruguayense
Ritossa et al. (2013)
Viozzi et al. (2009), Fernández et al. (2010)
Poeciliidae
Cnesterodon decemmaculatus
Stephanoprora aylacostoma
Stephanoprora uruguayense
Ostrowski de Núñez & Quintana (2008)
Ostrowski de Núñez (2007)
Poecilia reticulata Stephanoprora aylacostoma Ostrowski de Núñez & Quintana (2008)
DISCUSSION
The actual number of digenean species
recorded from Argentinean wild birds amounts
112 species, distributed in 65 genera of 20
families. Reports from the last 8 years account
for 28% of all reported species. Also, the
reports of the host-parasite relations
experienced similar increase during tha last
years (32%), reaching a total of 190 reports vs
130 in 2007. The full life cycle was elucidated
for 25 species of digeneans and some aspects
about their life history have been studied in 14
species. In addition to the records mentioned
above, there are many records of metacercariae
from fishes and amphibians without
correspondence with their definitive hosts,
even in the most studied families. Among the
Neotropical Helminthology. Vol. 9, Nº2, jul-dec 2015
Update of checklist of Digenean parasites of wild birds from Argentina
González, 2009, Hamann et al., 2010) and
Diplostomum gymnoti Szidat, 1969 recovered
from fishes (Szidat, 1969), were never found
parasitizing birds. Among the strigeids,
metacercariae of Cotylurus, described as
Tetracotyle loricariae Szidat, 1969 were
reported parasitizing fishes (Szidat, 1969),
however the adult forms were never found in
birds. Moreover, numerous adult digeneans
were obtained experimentally, but their natural
hosts are unknow, such as Echinoparyphium
megacirrus Semenas, Brugni & Ostrowski de
Núñez, 1999, Stephanoprora aylacostoma
(Echinostomatidae), Pygidiopsis crassus
Ostrowski de Núñez, 1995, Pygidiopsis
australis Ostrowski de Núñez, 1996,
Ascocotyle (Ascocotyle) secunda Ostrowski de
Núñez, 2001, Ascocotyle (Ascocotyle) tertia
Ostrowski de Núñez, 2001 (Heterophyidae),
C a t a t ro p i s c h i l i n a e , N o t o c o t y l u s
biomphalariae (Notocotylidae), Maritrema
patagonica (Microphallidae); or has never
been found in Argentinean birds, such as
A u s t r a l a p a t e m o n m a g n a c e t a b u l u m
(Strigeidae), Ascocotyle (Phagicola) angeloi
Travassos, 1928 and Ascocotyle (Phagicola)
longa Ransom, 1920 (Heterophyidae).
The distribution pattern of digeneans from
birds in the different ecoregions does not
reflect the real diversity of this group of
helminth in Argentina, since large areas remain
unexplored. Also, the bird species examined in
each ecoregion is different. For example, in the
Humid Chaco, the ecoregion with the highest
species richness, the hosts more studied are the
accipitrids, being the strigeids the digeneans
more frequent (12 species). In the Pampa
ecoregion, the larids and ardeids are the best
inquired, and the echinostomatid the group
most recognized (10 species). In the Paraná
Flooded Savanna ecoregion, the ardeids and
rallids are the best studied, being the
echinostomatids the group most representative
(13 species). In the Espinal ecoregion, the
larids are the most studied, and the
microphallids (4 species) the most frequent
group. In the remaining ecoregions is not
possible to recognize the representive groups
because of the scarce number of reports. The
frecuency of particular families of digeneans
in each region appear to be related to the focus
of the investigations, mostly in connection to
the habits and feeding preferences of birds,
rather than to the geographical distribution.
Thus, echinostomatids and microphallids are
frecuently recorded in those areas where
studies were focused on aquatic birds, while
strigeids are dominant in areas where most of
the studies were conducted on birds with
terrestrial habits.
The limitations to complete the inventory of
wild birds in Argentina are related with the
scarce number of bird species studied (only 7
% of Argentinean birds were reported as host
of digeneans), and the existence of large areas
few explored or unexplored.
Cons ideri ng th e abo ve me ntion ed
observations, and that the curves of species
accumulation and host-parasite relations
indicate that the asymptote has not been
reached, we conclude that the inventory of
digeneans of wild birds in Argentina is far from
being completed, even in the most studied
areas.
346
Neotropical Helminthology. Vol. 9, Nº2, jul-dec 2015 Drago & Lunaschi
Alda, P, Bonel, N, Cazzaniga, NJ & Martorelli,
SR. 2010. Effects of parasitism and
environment on shell size of the South
American intertidal mud snail Heleobia
australis (Gastropoda). Estuarine,
Coastal and Shelf Science, vol. 87, pp.
305-310.
Alda, P, Bonel, N, Cazzaniga, NJ & Martorelli,
SR. 2011c. Variación morfométrica en
Heleobia australis causada por
parásitos trematodes y factores
ambientales. Amici Molluscarum,
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Received July 8, 2015.
Accepted September 26, 2015.
