Neotrop. Helminthol., 6(1), 2012

2012 Asociación Peruana de Helmintología e Invertebrados Afines (APHIA)

ISSN: 2218-6425 impreso / ISSN: 1995-1043 on line

ORIGINAL ARTICLE / ARTÍCULO ORIGINAL

FIRST REPORT OF NEMATODE PARASITES OF PHYSALAEMUS ALBONOTATUS

(STEINDACHNER, 1864) (ANURA: LEIUPERIDAE) FROM CORRIENTES, ARGENTINA

PRIMER REPORTE DE NEMATODES PARÁSITOS DE PHYSALAEMUS ALBONOTATUS

(STEINDACHNER, 1864) (ANURA: LEIUPERIDAE) DE CORRIENTES, ARGENTINA

Cynthya Elizabeth González & Monika Inés Hamann

Abstract

Thirty-four adults of Physalaemus albonotatus (20 males and 14 females) were collected near the city

of Corrientes, Province of Corrientes, Argentina (27°28'S - 58°50'W) between January 2001 and

September 2004. Adults of four nematode species, Strongyloides sp., Oswaldocruzia sp., Cosmocerca

podicipinus, Cosmocerca parva and larvae of two other species, Spiroxys sp. and Physaloptera sp.

were recovered from the digestive tract, lungs and gastric mucosa. We present morphologic characters

(some with scanning electron microscopy), morphometric information and range extensions for these

nematode species. This is the first report of nematode parasites of P. albonotatus from Argentina.

Key words: Amphibians - Argentina - Corrientes - nematode parasites - Physalaemus albonotatus.

Resumen

Palabras clave: anfibios - Argentina - Corrientes - nemátodos parásitos, Physalaemus albonotatus.

Treinta y cuatro adultos de Physalaemus albonotatus (20 machos y 14 hembras) fueron colectados en

las proximidades de la ciudad de Corrientes, provincia de Corrientes, Argentina (27°28'S - 58°50'W)

entre enero del 2001 y septiembre del 2004 para el estudio de su fauna nematológica. Un total de cuatro

especies de nemátodos en estado adulto Strongyloides sp., Oswaldocruzia sp., Cosmocerca

podicipinus, Cosmocerca parva y dos especies en estado larval, Spiroxys sp. y Physaloptera sp.,

fueron halladas en el tracto digestivo, pulmón y mucosa gástrica. En este estudio se presentan datos

métricos y morfológicos de las especies halladas (algunos de ellos observados con microscopio

electrónico de barrido). Éste es el primer reporte de nemátodos parásitos para P. albonotatus en

Argentina.

Suggested citation: González CE & Hamann MI. 2012. First report of nematode parasites of Physalaemus albonotatus

(Steindachner, 1864) (Anura: Leiuperidae) from Corrientes, Argentina. Neotropical Helminthology, vol. 6, nº 1, pp. 9 -23.

Centro de Ecología Aplicada del Litoral, Consejo Nacional de Investigaciones Científicas y Técnicas.

Ruta Provincial Número 5, km 2,5. C. P. 3400. Corrientes, Argentina.

E-mail: cynthyaelizabethg@hotmail.com

MATERIAL AND METHODSINTRODUCTION

The nematode parasites of Argentinean

amphibians were analyzed mainly for bufonids

from northwestern (Ramallo et al., 2007a, b, 2008)

and for leptodactylids and bufonids from

northeastern (González & Hamann, 2006a, b,

2007a, b, 2008, 2009; Hamann et al., 2006a, b),

while in the family Leiuperidae were made few

studies of this topic. To our knowledge, in

Argentina, the following leiuperids has been

studied for nematode fauna: Physalaemus

biligonigerus (Cope, 1861) from Cordoba

province (Gutiérrez et al., 2005) and,

Pseudopaludicola falcipes (Hensel, 1867), P.

boliviana Parker, 1927 and Physalaemus

santafecinus Barrio, 1965 from Corrientes

province (Duré et al., 2004; González & Hamann,

2004, 2009, 2010a). About the species

Physalaemus albonotatus (Steindachner, 1864)

this study is absent. Thus, the purpose of the

present study is to report for the first time

nematode species that found in adults specimens of

P. albonotatus from Corrientes, Argentina.

This amphibian is found in Mato Grosso and Mato

Grosso do Sul (Brazil), Paraguay, and the Chacoan

regions of Bolivia and Argentina (Frost, 2011). In

Argentina is common in natural environments but

it can also be found in agricultural areas (Duré et

al., 2008). It can survive in both dry and moist

substrata, i.e. near to the shore of temporary,

semipermanent and permanent ponds, and also in

the flooded grass. It has a diet dominated by

formicids and isopterans and using an active

foraging as strategy for prey capturing (Duré,

2004). There are no major threats to this species, is

categorized as Least Concern by Aquino et al.

(2004).

We described for the first time five genus of

nematode parasites found in different organs of

this amphibian host and presented new metrical

and morphological data obtained with scanning

electron microscopy.

Thirty-four adults of P. albonotatus (20 males and

14 females) were collected near the city of

Corrientes, Province of Corrientes in Argentina

(27°28'S - 58°50'W) between January 2001 and

September 2004. Frogs were hand-captured,

mainly at night, using the sampling technique

defined as “visual encounters survey” (Crump &

Scott, 1994). Amphibians were transported live to

the laboratory, killed in a chloroform (CHCL )

solution. At necropsy, hosts were sexed

and the alimentary

canal, lungs, liver, kidneys, urinary bladder,

musculature and integument examined for

parasites by dissection. Nematodes were observed

in vivo, counted and killed in hot distilled water and

preserved in 70% ethyl alcohol, cleared in

glycerine or lactophenol and examined as

temporary mounts. Cyst were counted and isolated

from host tissues. Larvae were removed from cysts

using preparation needles and studied either in

vivo. Some specimens were studied by scanning

electron microscopy (SEM); these specimens were

dehydrated in ethanol series, dried using the

critical point technique, coated with gold, and

examined with a JSM-5800 scanning electron

microscope. Measurements are presented in

micrometers (µm) unless otherwise indicated as

mean values and standard deviation followed by

range values in parenthesis. The systematic

determination of the nematodes was carried out

following the guidelines given by Anderson et al.

(2009) and Gibbons (2010). Prevalence and mean

intensity were calculated according to Bush et al.

(1997).

The nematode specimens studied were deposited

in the Helminthological Collection of the Centro de

Ecología Aplicada del Litoral (CECOAL),

Corrientes, Argentina. The present study has

complied with all the regulations and ethical and

legal considerations for the capture and use of

animals established by the National Council of

Scientific Research and Technical of Argentina

(determined by examination of gonads and

external nuptial features),

Nematode parasites of Physalaemus albonotatus

González & Hamann

Neotrop. Helminthol., 6(1), 2012

A total of 74 nematodes were collected from this

leiuperid. These nematodes belonged to six

species of the following families: Strongyloididae

Chitwood and McIntosh, 1934: Strongyloides sp.,

Molineidae (Skrjabin and Schulz, 1937) Durette-

Desset and Chabaud, 1977: Oswaldocruzia sp.,

Cosmocercidae Travassos, 1925: Cosmocerca

podicipinus Baker and Vaucher, 1984 and

Cosmocerca parva Travassos, 1925,

Gnathostomatoidea Railliet, 1985: Spiroxys sp.

and, Physalopteridae Leiper, 1908: Physaloptera

sp. Each larva was spirally coiled inside the

brownish capsule.

RESULTS Strongyloides sp.

(Fig. 1A-B)

Prevalence, number of parasites: 2.9% (1 of 34

amphibians infected), 1.

Site of infection: large intestine.

Accession number: CECOAL 02113236 (1 gravid

female).

Description: Based on 1 female specimen. Body

slender, elongate, length 2.25mm, maximum

width 55.0. Stoma simple, elongate, surrounded by

six small lobes, each with one inconspicuous

papilla. Esophagus filiform, 710.0 long. Nerve

ring 170.0 from anterior end. Excretory pore not

observed. Vulva with well developed lip situated at

Figure 1A-E. Fig. 1A. Strongyloides sp. Female, total view. Fig. 1B. Strongyloides sp. Posterior end, ventral view.

Fig. 1C. Oswaldocruzia sp. Female, anterior end. Fig. 1D. Oswaldocruzia sp. Detail of anterior end. Fig. 1E.

Oswaldocruzia sp. Posterior end, lateral view.

Nematode parasites of Physalaemus albonotatus

1.6mm from anterior end. Uteri opposed. Oval

eggs, 50.0 in length x 34.0 wide. Anus 85.0 from

posterior end; sharply pointed tail.

Remarks: The posterior end of these nematodes

has been used as a diagnostic character for the

species of this genus that parasitize anurans.

Moravec and Kaiser (1995) have proposed two

types: Type A: Strongyloides species that have a

sharply pointed tail and, Type B: species that have

a rounded tail. Specimen analyzed in this study

might belong to the first type due to the

morphology of the posterior end. In the present

study, female nematode showed greater length

(2.25mm) that female of S. carinii Pereira, 1935

(1.3-1.7mm) and S. pereirai Travassos, 1932

(1.56-1.91mm) and, the distance of anus to

posterior end was greater (85.0) that were reported

for these species (S. carinii: 60.0; S. pereirai: 64.0-

72.0) analyzed by Pereira (1935) and Travassos

(1932), respectively.

Geographical distribution: In amphibians from

Neotropical Realm, species of this genus were

found in leptodactylids and bufonids from Brazil

(Travassos, 1932, 1933; Pereira, 1935; Luque et

al., 2005; Santos & Amato, 2010), in bufonids

from Cuba (Pérez-Vigueras, 1942) and, in

strabomantids from St. Vincent (Moravec &

Kaiser, 1995). This is the first report of this genus

for Argentinean amphibians.

Oswaldocruzia sp.

(Fig. 1C-E)

Prevalence, number of parasites: 2.9% (1 of 34

amphibians infected), 1.

Site of infection: small intestine.

Accession number: CECOAL 96040901 (1

immature female).

Description: Based on 1 female specimen. Body

9.0 mm x 150.0, with maximum width at level of

midbody. Lateral alae undeveloped. Cephalic

vesicle present with anterior swelling, total length

of cephalic vesicle 87.4, rising approximately 6.9

above body wall in the widest first part. Claviform

esophagus 540.0 length. Nerve ring, excretory

pore and vulva 170.0, 282.0 and 6.37mm from

anterior end, respectively. Amphidelphic. Tail

207.0 in length, terminating in flexible filament

approximately 11.0 in length. Female without

eggs.

Remarks: The identification of the species of this

genus is done by analyzing the caudal bursa of

males, the morphology of the spicules and the

constitution of synlophe (Ben Slimane et al.,

1996); because were not found males the specific

identification was not possible.

Geographical distribution: Oswaldocruzia is a

genus with a widely distribution in Neotropical

Realm. Ben Slimane et al. (1996) reported species

of this genus in leptodactylids from Chile,

leptodactylids and bufonids from Brazil,

leptodactylids from Paraguay, bufonids, hylids

and leptodactylids from Guyana, bufonids from

Venezuela, leptodactylids, bufonids and hylids

from Ecuador and, in hylids, leptodactylids and

bufonids from Cuba and Puerto Rico. In Argentina,

previous reports in amphibians hosts were:

Oswaldocruzia mazzai Travassos, 1935 in

bufonids from Jujuy province (Travassos, 1935,

1937), Oswaldocruzia sp. in leptodactylids from

Corrientes province (González & Hamann, 2006b)

and, O. proencai Ben Slimane and Durette-Desset,

1995 in bufonids from Corrientes (González &

Hamann, 2008) and Salta provinces (Ramallo et

al., 2007b). This is the first report of Argentinean

leiuperid as host of genus Oswaldocruzia.

Cosmocerca podicipinus Baker and Vaucher, 1984

(Fig. 2A-B)

Prevalence, number of parasites: 73.5% (25 of 34

amphibians infected), 59.

Site of infection: lung, large intestine and small

intestine.

Accession number: CECOAL 03074242 (1 male, 4

females); 02092915 (2 males, 4 females).

Description: Nematodes with conspicuous sexual

dimorphism; body cuticle provided with

transverse striations. Lateral alae present in both

sexes. Mouth with 3 small v-shaped lips, dorsal lip

with 2 sessile papillae, each ventrolateral lip with 1

ventral sessile papilla and 1 amphid; three

González & Hamann

Figure 2A. Cosmocerca podicipinus. Posterior end of

male, lateral view. pl: plectanes; la: lateral alae. Figure 2B. Cosmocerca podicipinus. Posterior end of

mail; detail of plectanes. rp: rosette papillae.

Neotrop. Helminthol., 6(1), 2012

additional pairs of cephalic papillae. The anterior

end of the esophagus presents three tooth-like

projections covered with a thick cuticle, also called

cuticular flap. Esophagus with short pharynx,

cylindrical corpus, and posterior bulb containing

valves. On the ventral surface of the preanal region

of male there are two longitudinal rows of comb-

like crests (plectanes). In this species there are 5

pairs of plectanes; the plectanes of each row are

fusioned by sclerotized supports very marked (Fig.

2A-B). Somatic papillae numerous, in two ventral,

two subventral, two dorsal and two subdorsal

rows, extending from cervical region to posterior

end. The metric characters of the males and

females of these nematode species are present in

Table 1.

Remarks: Some species of genus Cosmocerca

have a fixed number of pairs of plectanes in the

posterior end of males; C. chilensis Lent and

Freitas, 1948 and C. rara Freitas and Vicente, 1966

have 6 pairs, C. cruzi Rodrigues and Fabio, 1970,

C. travassosi Rodrigues and Fabio, 1970 and C.

podicipinus have 5 pairs, C. longispicula Moravec

and Kaiser, 1994 and C. vrcibradici Bursey and

Goldberg, 1998 and C. uruguayensis Lent and

Freitas, 1948 have 7 pairs. Other species have a

variable number of pairs of these structures: C.

brasiliensis Travassos, 1925 have 8-11 pairs, C.

paraguayensis Moravec and Kaiser, 1994 have 4-5

pairs and C. parva Travassos, 1925 have 4-7 pairs

(Goldberg et al., 2005). Thereon, specimens of C.

podicipinus examined in this study show the same

pattern as previous studies (5 pairs of plectanes);

nevertheless, in some of immature specimens, the

plectanes of each row do not fuse.

The metrical characteristic of C. podicipinus

compared with original description realized by

Baker and Vaucher (1984) with specimens

collected in leptodactylids from Paraguay shows

that males and females specimens analyzed from P.

albonotatus are smaller than those studied by these

authors (Baker & Vaucher, 1984: females:

2.86–7.33mm; males: 1.88–3.17mm vs. present

study: females: 2.58–5.68mm; males:

1.35–2.95mm). In males, the length of spicules and

gubernaculum are smaller than those found in

Paraguayan specimens (Baker & Vaucher, 1984:

spicules: 82-188; gubernaculum: 87-152 vs.

present study: spicules: 57-97; gubernaculum: 62-

141). Compared with specimens collected in a

leiuperid sympatric host, P. santafecinus studied

by González and Hamann (2010a), males analyzed

in this study have total length of body, length of

gubernaculum and length of spicules greater that

males analyzed from P. santafecinus (P.

Nematode parasites of Physalaemus albonotatus

Table 1. Morphological features and measurements of Cosmocerca podicipinus and Cosmocerca parva in Physalaemus

albonotatus; measurements are presented in micrometers unless otherwise indicated as mean values and standard deviation

followed by range values in parenthesis.

Cosmocerca podicipinus Cosmocerca parva

Females

(n= 10) Males

(n= 10) Females

(n= 10) Males

(n= 10)

Total length

(2.77 6.83) (1.15 1.72)

Width

(139 180)

Pharynx

4.13 0.87mm

(2.58 5.68)

220.5 33.1

(185 300)

35.6 7.7

(28 51)

34.3 6.3

(28 42)

2.16 0.56mm

(1.35 2.95)

201.8 57.3

(100 280)

26.6 4.1

(20 33)

18.1 3.4

(11 23)

(200 350)

(35 48)

(30 41)

(29 32)

(19 23)

Muscular

esophagus 276.5 25.0

(234 306)

38.5 4.7

(31 46)

204.2 27.9

(172 270)

22.9 3.5

(17 30)

(294 360)

(37 49)

(198 232)

(21 30)

Bulb 84.6 11.4

(69 103)

91.3 11.1

(80 115)

55.6 7.4

(46 70)

118)(87

(90 125)

(50 66)

x 6.4

(49 63)

Nerve ring * 209.5 27.0

(180 270)

55.1 7.7

(44 70)

173.0 33.6

(122 235)

(192 250)

(140 155)

Excretory pore * 336.6 49.8

410)(230 265.2 46.0

(198 350)

(330 390)

(160 240)

Anus from

posterior end 413.7 45.1

(350 510) 153.1 74.4

(103 360)

(102 140)

(225 370)

Gubernaculum - 96.3 24.5

(62 141) -

(78 107)

Spicules

- 85.3 10.7

(57 97) -

(92 130)

Plectanes - 5 pairs - 4 6 pairs

Adanal papillae - 3 pairs -

Vulva from

anterior end 2.13 0.44mm

(1.32 2.91) -

(2.0 2.5) -

Eggs 95.4 12.4

(78 122)

61.0 7.7

(53 76)

(89 118)

(49 68)

* from anterior end.

González & Hamann

Neotrop. Helminthol., 6(1), 2012

santafecinus: total length: 1.3–2.3mm;

gubernaculum: 75-92; spicules: 69-89 vs. present

study: total length: 1.35-2.95mm; gubernaculum:

62-141; spicules: 57-97); females, however, have a

total length less than those studied in P.

santafecinus (P. santafecinus: 4.27–7.8mm vs.

present study: 2.58–5.68mm).

Geographical distribution: In Neotropical Realm,

the genus Cosmocerca has a widely distribution in

amphibians hosts. Cosmocerca podicipinus was

found in families Bufonidae, Aromobatidae,

Strabomantidae and Leptodactylidae from Peru

(Bursey et al., 2001), in Leptodactylidae (Baker &

Vaucher, 1984) and Hylidae from Paraguay

(McAllister et al., 2010b), Bufonidae and

Dendrobatidae from Colombia (Goldberg &

Bursey, 2003), Hylidae (Campião et al., 2010;

Goldberg et al., 2007) and Leptodactylidae from

Brazil (Campião et al., 2009; Goldberg et al.,

2009), Dendrobatidae from Panama (Martinez &

Maggenti, 1989), Eleutherodactylidae,

Leptodactylidae, Hylidae (Goldberg & Bursey,

2002; Goldberg et al., 2002a) and Ranidae

(Goldberg & Bursey, 2002; Cabrera-Guzmán et

al., 2007) from Mexico, Bufonidae,

Leptodactylidae, Microhylidae, Strabomantidae,

Craugastoridae (Bursey & Brooks, 2010), Ranidae

(Bursey & Goldberg, 2005, 2006, 2007; Bursey &

Brooks, 2010), Brachycephalidae (Goldberg &

Bursey, 2008a), and Hylidae (Goldberg & Bursey,

2008b) from Costa Rica. In Argentina, it was found

in families Leiuperidae (González & Hamann,

2004, 2009a, 2010a), Leptodactylidae (González

& Hamann, 2006b; Hamann et al., 2006a, b;

Schaefer et al., 2006), Bufonidae (González &

Hamann, 2006a, 2007a, b, 2008), Cycloramphidae

(González & Hamann, 2009b) and Hylidae

(González & Hamann, 2008, 2011; Hamann et al.,

2010). This is the first record of C. podicipinus in P.

albonotatus from Argentina.

Cosmocerca parva Travassos, 1925

(Fig3. A-B)

Prevalence, number of parasites: 8.8% (3 of 34

amphibians infected), 10.

Site of infection: large intestine.

Figure 3A. Cosmocerca parva. Posterior end of male,

lateral view. rp: rosette papillae.

Figure 3B. Cosmocerca parva. Detail of anterior

extremity of male. cp: cephalic papillae; a: amphid; cf:

cuticular flap.

Nematode parasites of Physalaemus albonotatus

Accession number: CECOAL 02113235 (1

female), CECOAL 02103078 (1 female, 1 male).

Description: The Table 1 shows the metric

characters of the males and females of C. parva. In

this study, the number of plectanes and rosette

papillae in the posterior end of male varied

between 4 and 6 pairs, each rosette papillae with

two complete circles of punctations directed

perpendicular to the body surface; the plectanes of

each row are not fused to each other as in the

previous species (Fig. 3A). The anterior extremity

of this species presents the same number and

arrangement of cephalic papillae than C.

podicipinus (Fig. 3B).

Remarks: For C. parva some studies show, in the

same host, a fixed number of pairs of plectanes, in

other hosts, the number of pairs of plectanes for

this species was variable (see González &

Hamann, 2010a). In this study we found males

with new pattern of number of plectanes (4-6

pairs). On the other hand, adanals papillae present

the same number and arrangement to those found

in P. santafecinus (1 preanal pair, 1 lateral pair and

1 postanal pair); in addition, 1 unpaired papillae on

the anterior lip of anus.

Studies realized with SEM in males of C. parva

showed rosette papillae surrounded by 12-16

punctations in specimens collected in R. granulosa

(Mordeglia & Digiani, 1998); surrounded by 12-

15 interior punctations and 12-15 exterior

punctations in specimens collected in R.

schneideri (González & Hamann, 2008) and,

rosette papillae surrounded by 10-11 interior

punctations and 12-14 exterior punctations in

specimens collected in P. santafecinus (González

& Hamann, 2010a). Males of C. parva of this study

shows 11 interior punctations and 12 exterior

punctations in each one of rosette papillae.

Compared with original description realized by

Travassos (1925) with specimens collected in

Pernambuco State, Brazil, males of C. parva from

P. albonotatus present a total length and the length

of gubernaculum smaller than Brazilian

specimens, but length of spicules is greater

(Travassos, 1925: total length: 3.5mm;

gubernaculum: 120-140; spicules: 80 vs. present

study: total length: 1.15-172mm; gubernaculum:

78-107; spicules: 92-130). Compared with

specimens studied in a leiuperid sympatric host, P.

santafecinus, by González and Hamann (2010a),

the males of C. parva in this study are smaller and

females are greater than those analyzed in P.

santafecinus (P. santafecinus: males:

1.53–2.57mm; females: 3.43–5.92mm vs. present

study: males: 1.15–1.72mm; females:

2.77–6.83mm). Also, the length of the spicules was

higher in specimens collected from P. albonotatus

(P. santafecinus: 57-98 vs. present study: 92-130).

Geographical distribution: In Neotropical Realm,

C. parva was found in Brazil in the following

families: Bufonidae (Santos & Amato, 2010),

Leptodactylidae (Fabio, 1982), Leiuperidae and

Hylolidae (Travassos, 1925); in Mexico from

Ranidae (Paredes-Calderón et al., 2004), in

Guyana from Dendrobatidae and Hylidae

(McAllister et al., 2010a), in Paraguay from

Leptodactylidae, Bufonidae (Baker & Vaucher,

1984, McAllister et al., 2010b) and Hylidae (Masi

Pallares & Maciel, 1974), in Peru from Bufonidae,

Aromobatidae, Hylidae, Leiuperidae,

Leptodactylidae, Eleutherodactylidae,

Microhylidae (Bursey et al., 2001),

Strabomantidae and Dendrobatidae (McAllister et

al., 2010c), in Trinidad and Tobago from

Leptodactylidae and Eleutherodactylidae

(Goldberg et al., 2002b), in Costa Rica from

Brachycephalidae (Goldberg & Bursey, 2008a),

Hylidae (Goldberg & Bursey, 2008b),

Craugastoridae, Bufonidae, Eleutherodactylidae

and Ranidae (Bursey & Brooks, 2010), in

Colombia from Dendrobatidae (Sánchez et al.,

2010). In Argentina, it was found in the families

Bufonidae (Mordeglia & Digiani, 1998; González

& Hamann, 2006a, 2007a, b, 2008),

Leptodactylidae (González & Hamann, 2006b;

Hamann et al., 2006a, b, Schaefer et al., 2006),

Cycloramphidae (González & Hamann, 2009b),

Hylidae (González & Hamann, 2008, Hamann et

al., 2009, 2010) and Leiuperidae (González &

Hamann, 2006b, 2010a). This is the first record of

C. parva in P. albonotatus from Argentina.

González & Hamann

Neotrop. Helminthol., 6(1), 2012

Spiroxys sp. (Larvae)

(Fig. 4A-B)

Prevalence, number of parasites: 2.9% (1 of 34

amphibians infected), 1.

Site of infection: serous of stomach.

Accession number: CECOAL 10020901 (larvae).

Description: Based on 1 specimen. Small

nematodes with fine cuticular transverse striations.

Length of body 2.2 mm, maximum width 80.0.

Cephalic end provided with two triangular lateral

pseudolabia. Two cephalic papillae and one small

amphid present on either side at level of base of

pseudolabia. Walls of oral opening weakly

sclerotized. Stoma weakly developed, very short.

Esophagus starting at level of base of pseudolabia.

Esophagus divided into narrow, anterior muscular

part (length 330.0) and broader posterior glandular

part (length 750.0). Excretory pore not observed.

Intestine brownish, straight. Tail conical, 90.0

long, with rounded tip. Elongated cyst, measures

were 850.0 x 370.0.

Remarks: The identification of this larva is based

mainly on the characteristic large lateral triangular

pseudolabia. The principal differences compared

with it found in Pseudis paradoxa from Corrientes,

Argentina studied by González and Hamann

(2010b) are the form of cyst (almost spherical in P.

paradoxa, 380-400 x 350-375) and the smaller

metrics characteristics (e.g. length of body: 1.97

mm; length of muscular esophagus: 100.0; length

of glandular esophagus: 620.0; length of tail:

82.0).

Figure 4A-C. Fig. 4A. Spiroxys sp. Anterior end, lateral view. Fig. 4B. Spiroxys sp. Posterior end, ventrolateral

view. Fig. 4C. Physaloptera sp. Anterior end, ventral view.

Nematode parasites of Physalaemus albonotatus

Geographical distribution: In amphibian hosts of

Neotropical Realm, the genus Spiroxys is

previously reported in serous of stomach of hylid

P. paradoxa from Argentina (González &

Hamann, 2010b).

Physaloptera sp. (Larvae)

(Fig. 4C)

Prevalence, number of parasites: 2.9% (1 of 34

amphibians infected), 2.

Site of infection: gastric mucosa.

Accession number: CECOAL 06030612 (2

larvae).

Description: Based on 2 specimens. Larval body

whitish, 2.56±0.14mm (2.55-2.57) long and

117.5±3.5 (115.0-120.0) wide, with transversely

annulated cuticle. Terminus of head with two

lateral lips and cephalic collar formed by inflated

cuticle. On either lip a sclerotized support and one

terminal tooth present at upper margin. Each lip

bearing two cephalic papillae and one amphid.

Muscular esophagus 167.5±3.5 (165.0-170.0)

long by 32.0±4.2 (29.0-35.0) wide; length of

glandular esophagus 0.87±0.03mm (0.87-0.88) by

51.0±1.4 (50.0-52.0) wide. Nerve ring encircling

muscular esophagus at its posterior half, 115.0±7.1

(110.0-120.0) from anterior end. Excretory pore

near anterior end of glandular esophagus,

145.0±7.1 (140.0-150.0) from anterior end of

body. Tail conical, 110.0±7.1 (105.0-115.0) long.

Remarks: Compared with larvae that found in

other hosts of the same area, specimens analyzed

from P. albonotatus showed the same

morphological characteristics, but in terms of size,

were smaller [total length in P. santafecinus: 3.13-

3.7mm (González & Hamann, 2010a); in R.

fernandezae: 4.17mm (González & Hamann,

2007a); in R. granulosa: 4.6-5.6 (González &

Hamann, 2006a); in R. schneideri: 5.9mm

(González & Hamann, 2008)]. Also, the distance

of the excretory pore and nerve ring from anterior

end in specimens of this study are smaller that

observed in P. santafecinus (nerve ring: 161.0-

182.0, excretory pore: 168.0-204.0; González &

Hamann, 2010a), in R. shcneideri (nerve ring:

275.0, excretory pore: 360.0; González &

Hamann, 2008), in R. granulosa (nerve ring:

210.0-250.0, excretory pore: 240.0-340.0;

González & Hamann, 2006a), in R. fernandezae

(nerve ring: 162.0, excretory pore: 212.0;

González & Hamann, 2007a).

Geographical distribution: In amphibians of

Neotropical Realm larvae of Physaloptera were

found in families Cycloramphidae (Boquimpani-

Freitas et al., 2001), Leptodactylidae (Goldberg et

al., 2009), Hylidae, Bufonidae and Leiuperidae

(Vicente et al., 1990) from Brazil; in Bufonidae,

Aromobatidae, Hylidae, Leiuperidae,

Eleutherodactylidae, Leptodactylidae and

Microhylidae (Bursey et al., 2001) from Peru, in

Leptodactylidae (Goldberg et al., 2002b) from

Trinidad and Tobago, Bufonidae (Galicia-

Guerrero et al., 2000; Bursey & Goldberg, 2001;

Goldberg et al., 2002a; Espinoza-Jiménez et al.,

2007), Hylidae (Bursey & Goldberg, 2001;

Goldberg et al., 2002a) and Ranidae (Bursey &

Goldberg, 2001) from Mexico and, in

Brachycephalidae (Goldberg & Bursey, 2008a)

and Hylidae (Goldberg & Bursey, 2008b) from

Costa Rica. In it was found in families

Leiuperidae (Gutiérrez et al., 2005; González &

Hamann, 2010a), Leptodactylidae (González &

Hamann, 2006b), Hylidae and Bufonidae

(González & Hamann, 2007a, 2008).

Of a total of nine species of the genus Physalaemus

from Argentina, P. albonotatus, P. biligonigerus, P.

cuqui Lobo, 1993, P. cuvieri Fitzinger, 1826, P.

fernandezae (Müller, 1926), P. gracilis

(Boulenger, 1883), P. henselii (Peters, 1872), P.

riograndensis Milstead, 1960, P. santafecinus

(Lavilla et al., 2000), the study of P. albonotatus

represents the third report to the knowledge of the

nematode fauna of this genus of amphibians

(Gutiérrez et al., 2005; González & Hamann,

2010a).

Nematode parasites in the genus Physalaemus

from Argentina include Rhabdias sp. and

Physaloptera sp. in P. biligonigerus (Gutiérrez et

Argentina

DISCUSSION

González & Hamann

Neotrop. Helminthol., 6(1), 2012

al., 2005), C. podicipinus, C. parva, Aplectana

hylambatis (Baylis, 1927) Travassos, 1931 and

Physaloptera sp. in P. santafecinus (González &

Hamann, 2010a). Also, reports from Brazil

(Vicente et al., 1990) included Aplectana sp., A.

lopesi Silva, 1954, A. membranosa (Schneider,

1866) Miranda, 1924, C. parva, Oxyascaris

oxyascaris Travassos, 1920 and Physaloptera sp.

in P. signifer (Girard, 1853) and, Aplectana sp., C.

parva, O. oxyascaris and Physaloptera sp. in P.

soaresi Izecksohn, 1965. In the present study five

genera of nematode parasites (Strongyloides,

Oswaldocruzia, Cosmocerca, Spiroxys and

Physaloptera) are reported for the first time in P.

albonotatus. Additionaly, the genus Strongyloides

is reported for the first time in Argentinean

amphibians.

The genera Strongyloides, Oswaldocruzia and

Cosmocerca have a direct life cycle. The definitive

host becomes infected by skin penetration of

infective stage. Female nematodes expelled eggs

that develop in the external environment into first-

stage larvae and moult twice to the infective third

stage. Migration to the lungs is apparently a

necessary part of the development in species of

genus Cosmocerca (Anderson, 2000). Nematodes

of genera Spiroxys and Physaloptera have an

indirect life cycle. They need to complete the cycle

an intermediate host (arthropod) which is then

consumed by the predaceous final host (Anderson,

2000). Adults stage of genus Spiroxys parasite

stomach of different species of freshwater turtles.

Female nematodes expelled eggs with the turtle´s

faeces and the first-stage larvae develop in them.

Then, it is ingested by a copepod and, in the

haemocoel of this invertebrate, attains its third

stage (Anderson, 2000). On the other hand,

physalopterines are usually found firmly attached

to the gastric mucosa with the aid of large dentate

pseudalabia and a collarette which presses into the

mucosa. In this way the larval parasite may persist

for varying periods of time and be available to the

predaceous final hosts (reptiles, birds, mammals)

(Anderson, 2000). The presence of these larvae

indicates that the amphibians could play a role in

the transmission of these nematodes, serving as

Anderson, RC. 2000.

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correspondencia:

Cynthya Elizabeth González

Centro de Ecología Aplicada del Litoral; Consejo

Nacional de Investigaciones Científicas y

Técnicas, Ruta Provincial Número 5, km 2,5

(3400), Corrientes, Argentina.

E-mail / correo electrónico:

cynthyaelizabethg@hotmail.com

Author for correspondence / Autor para

Received November 9, 2011.

Accepted January 27, 2012.