ORIGINAL ARTICLE /ARTÍCULO ORIGINAL

A NEW SPECIES OF PHYSALOPTERA (NEMATODA: PHYSALOPTERIDAE) FROM

CERRADOMYS SUBFLAVUS (RODENTIA: SIGMODONTINAE) IN THE CERRADO

BIOME, BRAZIL

UNA NUEVA ESPECIE DE PHYSALOPTERA (NEMATODA: PHYSALOPTEROIDEA) DE

CERRADOMYS SUBFLAVUS (RODENTIA: SIGMODONTINAE) EN EL BIOMA

CERRADO, BRAZIL

1,2 1,2 3 4

Juliana São Luiz , Raquel Oliveira Simões , Eduardo Lopes Torres , Helene Santos Barbosa , Jeannie

5 6 7 2*

Nascimento Santos , Elane Guerreiro Giese , Fabiana Lopes Rocha & Arnaldo Maldonado Júnior

1Programa de Pós-Graduação em Biodiversidade e Saúde (PGBS), Instituto Oswaldo Cruz, Fundação Oswaldo Cruz, Av.

Brasil 4365, Manguinhos, Rio de Janeiro, RJ, 21045-900, Brazil

2Laboratório de Biologia e Parasitologia de Mamíferos Silvestres Reservatórios, Instituto Oswaldo Cruz, Fundação

Oswaldo Cruz, Av. Brasil 4365, Manguinhos, Rio de Janeiro, RJ, 21045-900, Brazil.

3Laboratório de Helmintologia Romero Lascasas Porto, Universidade do Estado do Rio de Janeiro, UERJ, Brazil Av.

Professor Manoel de Abreu, 444/50, Vila Isabel, Rio de Janeiro, RJ, 20511-070, Brazil.

4Laboratório de Biologia Estrutural, Instituto Oswaldo Cruz, Fundação Oswaldo Cruz, Av. Brasil 4365, Manguinhos, Rio

de Janeiro, RJ, 21045-900, Brazil.

5Laboratório de Biologia Celular e Helmintologia “Profa. Dra. Reinalda Marisa Lanfredi”, Instituto de Ciência Biológica,

Universidade Federal do Pará, UFPA, Augusto Corrêa, 01-Guamá, Belém 66075-110, PA, Brazil

6Laboratório de Histologia e Embriologia Animal, Instituto de Saúde e Produção Animal, Universidade Federal Rural da

Amazônia – UFRA, Av. Perimetral, 2501, CEP 66077-901, Montese, Belém, PA, Brasil

7Programa de Pós-Graduação em Ecologia e Monitoramento Animal, Universidade Federal da Paraíba- Campus IV

Litoral Norte. Centro de Ciências Aplicadas e Educação. Rua da Mangueira s/n Rio Tinto, João Pessoa, 58.297-000, PB,

Brazil.

*Correspondence to author: maldonad@ioc.fiocruz.br

Neotropical Helminthology, 2015, 9(2), jul-dec: 301-312.

ABSTRACT

Key words: nematode - rodent - taxonomic morphology - light microscopy - scanning electron microscopy

Physaloptera galvaoi n.sp. is described from the stomach of Cerradomys subflavus (Rodentia:

Sigmodontinae) collected in Serra da Canastra National Park (São Roque de Minas municipally,

Minas Gerais state, Brazil). The genus Physaloptera includes 13 species infecting rodents

worldwide. Among these, five occur in the Americas. The main characteristics of the new species

are size, shape and relation of spicule length to total body length, pre and postcloacal papillae

equidistant in males, and uterus with four to five branches in females. In relation to the

Physaloptera spp. that parasitize rodents in the Americas, the new species can be differentiated

from congeners as follows: Physaloptera bispiculata, Physaloptera murisbrasiliensis,

Physaloptera calnuensis, Physaloptera longispicula, and Physaloptera hispida because it is the

only one to have a short spicule and to present first and second pairs of papillae asymmetrically

displaced (left one anterior to right one) on male and tetra and pentadelphy on female, therefore

representing a new species for the genus. Thus, the specimens of nematodes parasites from

stomach of the C. subflavus represent a new species of Physalopteridae family named P. galvaoi

n.sp.

301

ISSN Versión impresa 2218-6425 ISSN Versión Electrónica 1995-1043

INTRODUCCIÓN

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Neotropical Helminthology. Vol. 9, Nº2, jul-dec 2015

Nematodes of the genus Physaloptera

Rudolphi, 1819 are parasites in the stomach of

reptiles, birds, mammals and occasionally

amphibians. They infect both wild animals and

pets, such as dogs and cats (Ortlepp, 1922;

Chabaud, 1975).

At present, 100 species of Physaloptera are

known (Pereira et al., 2012). Among these, 13

have been described infecting rodents

worldwide, including four species on South

America: P. bispiculata, Vaz and Pereira,

1935; P. murisbrasiliensis Diesing, 1860; P.

longispicula Quentin, 1968 (Vicente et al.,

1997); and P. calnuensis Sutton, 1989, with

only one in North America, P. hispida Schell,

1950.

The rice rat, subflavus (Wagner, Cerradomys

1842) (syn. Oryzomys subflavus) has

terrestrial habits and feeds on seeds, leaves and

arthropods (Souza et al., 2004). This rodent

was originally described in Lagoa Santa,

Minas Gerais and also occurs in the Brazilian

states of Góias, São Paulo and Bahia

(Langguth & Bonvicino, 2002; Percequillo et

al., 2008).

The vegetation of Serra da Canastra National

Park is characterized as an ecotone of Atlantic

Forest and cerrado biomes, with predominance

of high-altitude fields that harbor numerous

species of cerrado (savanna) fauna and flora.

Most of the Park consists of meadow

interspersed with forest, and the soil is acidic

and poorly drained. Grasses, sedges and

pipeworts predominate. The dry herbaceous

fields mainly contain grasses, while the

shrubby fields contain grasses and shrubs or

small trees that are widely spaced (Romero &

Nakajima, 1999).

In the present study, a new species of

Physaloptera (Spirurida: Physalopterinae) is

described found parasitizing the stomach of C.

subflavus in São Roque de Minas municipally

of Minas Gerais state.

RESUMEN

Palabras clave: microscopía de luz - microscopía electrónica de barrido - nematode - roedor - taxonomía morfológica.

Physaloptera galvaoi n.sp. fue registrada parasitando en el estómago de Cerradomys subflavus

(Rodentia: Sigmodontinae) capturados en el Parque Nacional Serra da Canastra (São Roque de

Minas, Minas Gerais, Brasil). El género Physaloptera tiene 13 especies que parasitan los

roedores en todo el mundo. Entre ellos, cinco fueron registrados en las Américas. Las principales

características de la nueva especie son el tamaño, la forma de la espícula y la relación de longitud

de la espícula con la longitud total del cuerpo, las papilas pre y pos cloacal equidistantes en los

espécimen masculinos, y el útero con cuatro hasta cinco ramas en las hembras. La nueva especie

difiere de las otras especies del género Physaloptera registradas en América como Physaloptera

bispiculata, Physaloptera murisbrasiliensis, Physaloptera calnuensis, Physaloptera

longispicula Physaloptera hispida y por tener: (1) una espícula pequeña, (2) primero y segundo

par de papilas asimétricamente desplazadas (la izquierda anterior a la derecha), (3) útero de

cuatro hasta cinco ramas (tetra y pentadelphy). Estos caracteres morfológicos representan a P.

galvaoi como una nueva especie para el género. Por lo tanto, los especimens de nemátodos

parásitos de estómago del hospededor C. subflavus representan una nueva especie de la familia

Physalopteridae llamado P. galvaoi n.sp.

São Luiz et al.

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Neotropical Helminthology. Vol. 9, Nº2, jul-dec 2015

images were captured using an Olympus DP-

12 digital camera.

Five male and five female nematode

specimens were separate for SEM. These

specimens were washed in 0.1 M Na-

cacodylate buffer, pH 7.2, post-fixed in 1%

OsO and 0.8% K Fe(CN) , dehydrated in

4 3 6

graded ethanol (30-100%) for 2 h, and dried by

the critical point method with CO (CPD 030,

Balzers, Switzerland). The samples were

mounted on aluminum stubs, coated with a 20-

nm layer of gold and examined with a Jeol JSM

6390LV scanning electron microscope

(operating at 15kV) at the Rudolf Barth

Electron Microscopy Platform of Oswaldo

Cruz Institute.

General: Body robust, elongated with fine

cuticular striations, tapering gradually at the

anterior end. Females more robust than males.

Cuticle in cephalic region reflecting to

posterior margins of pseudolabia, forming

cephalic collar (Figs. 2, 6 and 12). Oral

opening with two large lateral, semicircular

pseudolabia, each bearing three internal lateral

teeth, forming a tripartite structure and single

large external lateral triangulate tooth with

cuticular fold (Figs. 12, 13 and 15). Tripartite

internal tooth composed of unequal tips, two

rounded and one pointed triangular, with a pore

on internal surface (Fig. 15). Two pairs of

small teeth in the external lateral cuticular fold

on each side, totaling four small teeth. Each lip

has one pair of papillae in depression and one

amphid between them (Figs. 13 and 14). Three

well-delimited porous areas present on each lip

(Figs. 12 and 13 ). Cuticle with transverse

striations (Fig. 13). Esophagus long, divided

into anterior muscular and posterior glandular

parts (Figs. 1, 2 and 6). Nerve ring encircling

final portion of muscular esophagus (Fig. 1).

Male (holotype and six paratypes): 12.4 [10-

RESULTS

MATERIAL AND METHODS

Nematodes were collected from the stomach of

a naturally infected C. subflavus in the Serra da

Canastra National Park (20°13'28.30” S;

46°30'39.20” W), municipality of São Roque

de Minas, state of Minas Gerais, Brazil.

Collection was carried out in May 2010, using

a Tomahawk® trap (model 201; 16 x 5 x 5

inches or 40.6 x 12.7 x 12.7 cm) and a

Sherman® trap (model XLK; 3 x 3.75 x 12

inches or 7.6 x 9.5 x 30.5 cm). This study was

approved by the ethics committee of Oswaldo

Cruz Foundation/Fiocruz (CEUA L-0015/07).

Capture and sampling of small mammals was

authorized by Chico Mendes Institute for

C o n s e r v a t i o n o f B i o d i v e r s i t y

(ICMBIO/SISBIO, license number 18635-1).

For euthanasia, the animals were heavily

anesthetized with an intramuscular injection of

thiopental sodium, followed by intracardiac

administration of potassium chloride, under

the supervision of a licensed veterinarian.

Parasites were washed in saline and had their

anterior and posterior ends cut and fixed in hot

AFA ( 2% g l aci a l ac e ti c ac i d, 3 %

formaldehyde, and 95% of 70 ethanol) for

morphological identification. Drawings were

made using a drawing tube attached to a Zeiss

standard microscope. Measurements are in

millimeters unless otherwise stated for

holotype and allotype. The range (paratypes)

in brackets and mean in parentheses. Type

species of Physaloptera murisbrasiliensis

(CHIOC 9493) and representatives specimens

of Physaloptera bispiculata (CHIOC 36798)

colleted from Nectomys squamipes Brants,

1827, municipality of Sumidouro, Rio de

Janeiro deposited in the Helminth Collection

of the Oswaldo Cruz Institute (CHIOC), were

used for study.

The nematodes were clarified in 80% phenol

and mo u n t e d o n t e m porary slides.

Morphological analyses were conducted using

an Olympus BX-51 light microscope and

A new species of Physaloptera from Cerradomys

304

Figures 1-5. Physaloptera galvaoi n.sp. (1) Female anterior end showing muscular and glandular esophagus, nerve ring, vulva

opening. Scale bar: 100 µm. (2) Male anterior end showing tripartite tooth, cephalic papillae and cephalic collar. Scale bar: 100

µm. (3) Uterus with five uterine branches. Scale bar: 500 µm. (4) Male posterior end with four pairs of pedunculated papillae,

thirteen sessile papillae and one median protuberance-like papilla. Scale bar: 100 µm. (5) Left (L) and right (R) spicule. Scale bar:

100µm.

Neotropical Helminthology. Vol. 9, Nº2, jul-dec 2015 São Luiz et al.

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Figures 6-11. Light microscopy Physaloptera galvaoi n.sp. (6) Male anterior end and dorsal view. (7) Male posterior end, ventral

view showing pedunculated papillae (arrow) and sessile papillae (arrowhead); Detail of fifth pair sessile papilla (arrowhead) and

median protuberance-like papilla (*); (8) Ventral view of the precloacal papillae. Scale bar: 500 µm; (9) Ventral view of two pairs

postcloacal papillae (arrowhead). Scale bar: 100 µm; (10) Uterus with five uterine branches (arrowhead). (11) Female posterior

end showing anus (arrowhead). Scale bar: 100 µm.

Neotropical Helminthology. Vol. 9, Nº2, jul-dec 2015 A new species of Physaloptera from Cerradomys

306

Figures 12-19. Scanning electron microscopy Physaloptera galvaoi n. sp. (12). Female anterior end showing pseudolabia with

cephalic papillae (p), tripartite and small teeth (arrowhead), cervical region (cc), amphid (a) and porous areas (). (13). Detail

showing two pairs of lateral small teeth on pseudolabia (arrows). (14). Detail of small tooth (arrow) and cephalic papilla (p). (15).

Detail tripartite teeth (arrowhead), small tooth (arrow) and cephalic papilla (p). (16). Male posterior end in dorsal view. (17).

Male posterior end in ventral view showing two lateroventral folds (double arrow). (18). Male posterior end in ventral view, pairs

peduncullated papillae (pp), sessiles papillae (ss), cloacal openning and one median protuberance-like papilla (arrow). (19).

Female posterior end, showing the anal opening (arrowhead) and phasmids (arrow).

Neotropical Helminthology. Vol. 9, Nº2, jul-dec 2015 São Luiz et al.

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tail (Fig. 19). ). Eggs elliptical 0.04 [0.04-0.05

(0.05)] long and 0.03 [0.02-0.03 (0.03)] wide.

Anus opening 0.5 [0.43-0.50 (0.46)] from the

posterior end (Figs. 11 and 19; Table 2).

Taxonomic summary

Type host: Cerradomys subflavus (Wagner,

1842) (Rodentia: Cricetidae).

Site of infection: Stomach.

Type locality: São Roque de Minas

municipality, Serra da Canastra National Park,

Minas Gerais, Brazil (20 13'28.30" S;

46 30'39.20" W).

Prevalence of infection: 43% (3 rodents

infected/7 rodents collected)

Specimens deposited: Holotype male (CHIOC

no. 36758a) and allotype female (CHIOC

no.36758 b) were deposited in the Helminth

Collection of the Oswaldo Cruz Institute, Rio

de Janeiro, Brazil.

Etymology: New species is named in honor of

Dr. Cléber Galvão Ferreira for his

contributions to parasitology.

The genus Physaloptera was created by

Rudolphi (1819) having as type species

Physaloptera clausa parasitizing Erinaceus

dealbatus Swinhoe, 1870. Dujardin (1845)

suppressed the genus Physaloptera,

transferring its species to the genus Spiroptera.

Later, Diesing (1851) re-established

Physaloptera genus and included two new

species. Seurat (1914) highlighted the

taxonomic importance of uterus number

within the genus Physaloptera and

reorganized species groups according to the

presence of two or four uterus (Ortlepp, 1922).

Travassos (1920) maintained within

Physaloptera the species having up to two

uterus and created the Turgida genus from the

description of T. torresi, using as criterion the

presence of 10 uterus, and transferred

23 (15.5)] mm long, 0.64 [0.8-0.9 (0.8)] mm

wide at mid-body. Muscular and glandular

esophagus 0.5 [0.46-0.58 (0.5)] and 2.62 [3.0-

3.74 (3.4)] long, respectively. Nerve ring and

excretory pore 0.3 [0.2-0.4 (0.4)] and 0.5

[0.52-0.95 (0.8)] from the anterior end,

respectively (Fig. 1; Table 1). Male posterior

end with caudal fold not tapered at the end

(Figs. 4 and 7). Cloacal opening is crescent

shaped (Figs. 8 and 9). Twenty-one caudal

papillae, four pedunculated externolateral

papillae on each side of the caudal alae, three

sessile precloacal papillae different in size, a

single median papilla and one pair of papillae

situated on the same transverse line (Figs. 4, 7

and 8), and two pairs of sessile papillae slightly

posterior to cloacal opening (Figs. 4 and 9).

Three pairs of postcloacal papillae equidistant

to each other: first and second pairs

asymmetrically displaced (left one anterior to

right one), third pair symmetrical with a

median protuberance-like papilla (Figs. 4, 7,

17 and 18). Male tail 0.47 [0.28-0.61 (0.47)],

Posterior ventral end ornate with longitudinal

ridges, lateral region finely striated and central

region rough. Smooth dorsal surface caudal

alae (Fig. 16). Spicules are sub-equal and

dissimilar. Left spicule lanceolate, 0.26 [0.23-

0.28 (0.26)] long, right spicule stout and

slightly curved at tip, 0.21 [0.19-0.23 (0.21)]

long, representing 1.57% and 1.27% of total

body length (SpL/BL), respectively (Fig. 5,

Table 1).

Female (allotype and eight paratypes): 11.58

[16.4-31.3 (23.28)] mm long and 1.06 [1.10-

3.18 (1.44)] mm wide at mid-body. Muscular

and glandular esophagus 0.49 [0.42-0.71

(0.56)] and 1.44 [1.27-4.14 (2.7)],

respectively. Nerve ring and excretory pore

0.20 [0.20-0.28 (0.19)] and 0.7 [0.68-1.66

(1.17)] from the anterior end, respectively

(Figs. 1; Table 2). Uterus varied from four to

five uterine branches (Figs. 3 and 10). Vulva

opening 9.0 [6.42-11.14 (8.78)] from the

anterior end (Fig. 1). Rounded lateral

phasmids between the anus and tip of conical

DISCUSSION

Neotropical Helminthology. Vol. 9, Nº2, jul-dec 2015 A new species of Physaloptera from Cerradomys

308

P. murisbrasiliensis

Diesing, 1860

P. bispiculata

Vaz & Pereira, 1935

P. hispida

Schell, 1950

P. longispicula

Quentin, 1968

P. calnuensis

Suton, 1989

P. galvaoi

n. sp.

Present study

Lenght 22-28

25.0

30-42

34.4

17.1

10-23

Width 0.87-0.95

0.8

0.9-1.4

1.4

0.61

0.8-0.9

Glandular esophagus

3.2

4.2-5.3

5.3

2.99

3.0-3.74

Muscular esophagus

0.5

0.56-0.73

0.6

0.33

0.58-0.46

Nerve ring*

0.55-0.7

0.5

0.34

0.20-0.43

Excretore pore

0.87-1.2

0.83

0.80

0.52-0.95

Deirids -

0.67-0.91

0.7

0.95

Tail** -

0.87

1.9

0.58

0.28-0.61

Right spicule

0.40

0.39-0.55

0.71

0.31

0.19-0.23

SpL/BL 1.6%

1.6%

1.29%

2.06%

1.81%

1.57%

Left spicule

0.40

0.46

0.34-0.48

0.85

0.42

0.23-0.28

SpL/BL 1.6%

1.84%

1.13%

2.47%

2.45%

1.27%

Caudal papillae

protuberance-like papilla

Host Mus brasiliensis

(=Holochilus brasiliens)

Nectomys squamipes

Sigmodon hispidus

Thrichomys apereoides

(=Cercomys cunicularius)

Calomys laucha

Cerrdomys subflavus

Site of infection

Stomach

Locality Brazil

Brazil

USA

Brazil

Argentina

Brazil

p = present; np =not present by the authors; SpL/BL= relation spicule lenght by total body length; all measurements are in millimetres. *Distance from the anterior end; **Distance from the cloaca to the

posterior end; Number.

Table 1. Morphological features and measurements, vertebrate host and locality of Physaloptera spp. males infecting rodents from Americas.

Neotropical Helminthology. Vol. 9, Nº2, jul-dec 2015 São Luiz et al.

309

P. murisbrasiliensis

Diesing, 1860

P. bispiculata

Vaz & Pereira, 1935

P. hispida

Schell, 1950

P. calnuensis

Suton, 1989

P. galvaoi

n. sp.

Present study

Lenght 35-43 27-55 53-64 28.07 9.3-1.42

Width 1.1-1.75 1.0-1.9 1.9-2.0 0.99 1.10-3.18

Glandular esophagus - 3.8-5.4 5.0-7.0 3.32 1.27-4.14

Muscular esophagus - 0.54-0.7 0.64-0.69 0.39 0.42-0.71

Nerve ring* - - 0.71-0.74 0.38 0.20-0.28

Excretore pore - 1.2 0.84-1.08 0.78 0.68-1.66

Deirids - - 0.67-0.91 0.77 -

Vulva - 0.46 11-16 12.28 0.43-0.50

- - - 0.52 0.28-0.61

Eggs 0.045 x 0.026 0.05 x 0.036 0.040-0.052 x 0.023-0.030 0.039 x 0.019

0.043-0.50x

0.26-0.030

Uteri branch 2 2 2 - 4-5

Host

Holochilus brasiliensis

(= Mus brasiliensis)Nectomys squamipes Sigmodon hispidus Calomys laucha Cerrdomys subflavus

Site of infection Stomach Stomach Stomach Stomach Stomach

Locality Brazil Brazil USA Argentina Brazil

All measurements are in millimetres. *Distance from the anterior end; **Distance from the anus to the posterior end

Table 2. Morphological features and measurements, vertebrate host and locality of Physaloptera spp. females infecting rodents from Americas.

Tail**

Neotropical Helminthology. Vol. 9, Nº2, jul-dec 2015 A new species of Physaloptera from Cerradomys

310

and tetra and pentadelphy in female, therefore

representing a new species for the genus.

Some authors consider the presence of a

papilla-like protuberance in Physaloptera,

whereas in some original descriptions it is

depicted but not considered as a papilla, or

probably overlooked. In fact, P. galvaoi n. sp.

share this characteristic with P. bispiculata, P.

hispida, and P. calnuensis (Mafra & Lanfredi,

1998; Pereira et al., 2012). There are few

studies about SEM comparing Physaloptera.

SEM studies of Physaloptera have reported

the presence of small teeth in the external

lateral cuticular fold on each side of the

pseudolabia. Physaloptera rara Hall and

Wigdor, 1818 (Naem & Asadi, 2013),

Physaloptera clausa Rudolphi, 1819 (Gorgani

et al., 2013) and Physaloptera herthameyerae.

Torres et al. (2009) showed only one small

tooth on the external lateral cuticular fold on

each side of the pseudolabia, although P.

bispiculata Vaz and Pereira, 1935 (Mafra &

Lanfredi, 1998) and P. galvaoi n. sp. showed a

small tooth on the external lateral cuticular fold

on each side of the pseudolabia.

Thus, the presence of specific characteristics

such as size, shape and ratio of spicule length to

total body length, three pairs of postcloacal

papillae equidistant to each other: first and

second pairs asymmetrically displaced (left

one anterior to right one), third pair

symmetrical with a median papilla-like

protuberance in males, and uterus with four to

five branches in females, allow us to consider

the specimens from C. subflavus as a new

species.

Despite the consistency of the specific

morphological characters in Physaloptera

species, will be important for expanding

understanding adopt an integrated approach

including molecular data.

Physaloptera turgida Rudolphi, 1819 to the

new genus.

Yamaguti (1961) agreed with the systematic

classification of Physaloptera proposed by

Travassos (1920). More recently, Anderson et

al. (2009) accepted the presence of two to four

uterus as generic characteristics of

Physaloptera, while Turgida is characterized

by having more than four uterus. However,

Ortlepp (1922) stated that Physaloptera and

Turgida share generic traits, with the

differences being restricted to polidelphy.

Recently, Pereira et al. (2012) reorganized the

species of Physaloptera according to the

number of uterus and divided it into 58

didelphy species, two tridelphys and seven

tetradelphy. For other species, the type of

uterus is unknown. P. galvaoi sp. n. presents

m o r p h o l o g i c a l a n d m o r p h o m e t r i c

characteristics that allow classifying it within

Physaloptera genus, particularly by having 4

to 5 uterus (tetra and pentadelphy). The main

characteristics of P. galvaoi sp. n. include

spicules sub-equal and not long, three pairs of

postcloacal papillae equidistant to each other:

first and second pairs asymmetrically

displaced (left one anterior to right one), third

pair symmetrical with a median protuberance-

like papilla in males, and uterus with four or

five branches in females.

According to Schell (1950) and Sutton (1989) ,

in comparison with the other species of

Physaloptera that parasitize rodents

sigmodontinae and Eumysopinae in the

Americas, the new species can be

differentiated from congeners as follows: P.

bispiculata, P murisbrasiliensis, P. calnuensis,

P. longispicula, and P. hispida because it is the

only one to have short spicule and to present

fi r st and sec o n d pairs of p a pillae

asymmetrically displaced (left one anterior to

right one) in male. In addition, P. galvaoi n. sp.

resembles P. hispida when considered the ratio

between body length/size spicule. However P.

hispida displays twice the size P. galvaoi body,

Neotropical Helminthology. Vol. 9, Nº2, jul-dec 2015 São Luiz et al.

311

new species from Eastern Brazil.

American Museum Novitates, Nova

Iorque, vol. 3622, pp. 1-46.

Pereira, FB, Alves, PV, Rocha, BM, Lima, SS

& Luque, JL. 2012. A new Physaloptera

(Nematoda: Physalopteridae) parasite

of Tupinambis merianae (Squamata:

Teiidae) from southeastern Brazil.

Journal of Parasitology, vol. 98, pp.

1227-1235.

Vicente JJ, Rodrigues H O, Gomes DC & Pinto

RM. 1997. Nematóides do Brasil. Parte

V: nematóides de mamíferos. Revista

Brasileira de Zooogia, vol. 14 (supl.

1):1-452.

Romero R & Nakajima JN. 1999. Espécies

endêmicas do Parque Nacional da Serra

da Canastra, Minas Gerais. Revista

Brasileira de Botânica, vol. 22 , pp. 259-

265.

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