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Taxonomy and Distribution of Nonnapsylla (Siphonaptera: Stephanocircidae:
Craneopsyllinae)
Author(s) :Marcela Lareschi, Analía G. Autino, M. Mónica Díaz, and Rubén M. Barquez
Source: Journal of Parasitology, 97(5):954-955. 2011.
Published By: American Society of Parasitologists
DOI: 10.1645/GE-2738.1
URL: http://www.bioone.org/doi/full/10.1645/GE-2738.1

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Taxonomy and Distribution of Nonnapsylla Wagner, 1938 (Siphonaptera:

Stephanocircidae: Craneopsyllinae)

Marcela Lareschi, Analı́a G. Autino*, M. Mónica Dı́az*�, and Rubén M. Barquez*�, Centro de Estudios Parasitológicos y de Vectores, CEPAVE
(CCT-La Plata, CONICET-UNLP) Calle 2 #584, 1900 La Plata, Argentina; *Facultad de Ciencias Naturales e Instituto Miguel Lillo, Universidad
Nacional de Tucumán y PIDBA (Programa de Investigaciones de Biodiversidad Argentina), Miguel Lillo 205, San Miguel de Tucumán (4000),
Tucumán, Argentina; �CONICET (Consejo Nacional de Investigaciones Cientı́ficas y Técnicas), Argentina. e-mail: mlareschi@cepave.edu.ar

ABSTRACT: The genus Nonnapsylla Wagner, 1938 (Craneopsyllinae)
includes only 1 species, Nonnapsylla rothschildi Wagner, 1938. Two
subspecies are recognized by the shape of the dorsal margin of the helmet,
apical shape of the spines in helmet and genal combs, length of first and
fourth segments of the maxillary palpus, and shape of the hilla of the
spermatheca. The validity of the characters used to separate the 2 subspecies
of Nonnapsylla rothschildi Wagner, 1938 is discussed based on information
from the literature, as well as from direct observation of type specimens and
specimens collected in northwestern Argentina. We conclude that
Nonnapsylla rothschildi wagneri Johnson, 1957 should be considered as a
synonym of the nominal subspecies. We also extend the southern limits of
the distribution of the species and report its presence in Argentina.

Helmeted fleas belong to the Stephanocircidae; they are distinguished
from other fleas by the division of the anterior portion of the head, which
forms a ‘‘true helmet’’ with combs along its posterior margin, and by the
presence of a genal comb (Johnson, 1957; Whiting et al., 2008). The family
includes 2 subfamilies with 9 genera and 51 species. Stephanocircinae
contains only 1 genus, associated with marsupials from the Australian
region, while Craneopsyllinae has 8 genera associated with marsupials and
rodents from South America (Hopkins and Rothschild, 1956; Johnson,
1957; Whiting et al., 2008). Nonnapsylla Wagner, 1938 (Craneopsyllinae)
includes only 1 species, Nonnapsylla rothschildi Wagner, 1938; its
description was on based of 4 females collected from the bird
Opisthocomus hoazin Illiger, 1811 (Cuculiformes, Opisthocomidae), an
accidental host from Rı́o Yapacani, Santa Cruz, Bolivia. The genus is
differentiated from other Craneopsyllinae based on the helmet, which is
angled on its anterior margin and divided from the rest of the head only
dorsally (Wagner, 1938; Hopkins and Rothschild, 1956; Johnson, 1957).
Johnson (1957) described the subspecies Nonnapsylla rothschildi wagneri
based on 8 females and 2 males collected from Galea musteloides Meyen,
1832 (Rodentia, Caviidae) at 2 localities in Peru (Pisacoma and 4 km
northwest of Pomata, Puno Department); these fleas are distinguished by
the shape of the dorsal margin of the helmet, the apical shape of the spines
in the helmet and genal combs, the length of first and fourth segments of
the maxillary palpus, and shape of the hilla of the spermatheca.

Beaucournu and Gallardo (1989) reported N. r. wagneri based on
specimens (a male and 3 females) collected on Octodontomys gliroides
(Gervais and d’Orbigny, 1844) (Rodentia, Octodontidae) and Abrocoma
cinerea Thomas, 1919 (Rodentia, Abrocomidae) from Iquique, Parnicota
(Tarapacá Region), Chile. Differences in the phallosome of these
specimens, compared with those described by Johnson (1957), were
mentioned, but they were not detailed (Beaucournu and Gallardo, 1991).
However, the latter authors suggested that the characters used to
differentiate females of the 2 subspecies were weak and that probably
N. r. wagneri is a synonym of the nominal subspecies.

Here, we add N. rothschildi to the flea fauna of Argentina and analyze
the strength of the diagnostic characters used in the literature to
differentiate these subspecies. Fleas were collected at the following 2
localities in Jujuy Province in northwestern Argentina: Cuesta del Hurón,
29 km west of Cienegüillas on provincial route 64 (22u069S, 66u039W;
3,835 m, Santa Catalina Department), and Curques, 24 km north of
Susques, on provincial route 74 (23u169S, 66u279W; 4,100 m, Susques
Department) (Fig. 1). The Jujuy Province is bordered by the Argentine
province of Salta on the east and south, Bolivia to the north, and Chile to
the west. The mammal hosts were captured alive, and fleas were collected

by examining their pelage. Fleas were fixed in 96% ethanol and prepared;
they were photographed following conventional techniques for their study
using light microscopy. Voucher specimens were deposited at the section
of Annexes of the Colección Mamı́feros Lillo (CML), Universidad
Nacional de Tucumán, Argentina. For comparison purposes, the holotype
and allotype of N. r. wagneri, deposited at the Collection of Division of
Insects of the Field Museum of Natural History (Chicago, Illinois), were
examined as well as drawn using a microscope equipped with a drawing
tube; descriptions and figures (Wagner, 1938; Hopkins and Rothschild,
1956; Johnson, 1957) of the 2 subspecies were analyzed and compared.

The following 14 specimens of fleas (11 females and 3 males) were
identified as N. rothschildi because they perfectly fit with diagnostic
characters of the species: Curques, 24 km north of Susques, on provincial
route 74: 2 females on Eligmodontia puerulus (Philippi, 1896) (Rodentia:
Cricetidae) (MMD 7974-1/2); Cuesta del Hurón, 29 km west of Cienegüillas
on provincial route 64: 1 male on Galea musteloides Meyen, 1832 (Caviidae)
(CML 7136), 7 females and 2 males on Octodontomys gliroides (Gervais and
d’Orbigny, 1844) (Octodontidae) (2 females on CML 7143-1/2 and 5 females
and 2 males on CML7144-1/7), and 2 females on Phyllotis xanthopygus
(Waterhouse, 1837) (Cricetidae) (CML 7969, 7970).

In these specimens, the following characters, frequently used to
differentiate the 2 subspecies in the literature, were observed: males and
females with helmets angled on their dorsal margins, with 10 dorsal spines

DOI: 10.1645/GE-2738.1

FIGURE 1. Distribution of Nonnapsylla. (1) Rı́o Yapacani (Santa Cruz
Department), Bolivia; (2) Pisacoma (Puno Department), Perú; (3) 4 km
NW of Pomata (Puno Department), Perú; (4) Iquique, Parnicota
(Tarapacá region), Chile; (5) Cuesta del Hurón, 29 km west of Cienegüillas
on provincial route 64 (Santa Catalina Department, Jujuy), Argentina
(22u069S, 66u039W; 3,835 m); (6) Curques, 24 km north of Susques, on
provincial route 74 (Susques Department, Jujuy), Argentina (23u169S,
66u279W; 4,100 m). Localities 1 to 4 are written as in the literature; 5 and 6
were sampled by us, so the geographic coordinates are given as they were
recorded in the field.

J. Parasitol., 97(5), 2011, pp. 954–955

F American Society of Parasitologists 2011

954



in females, and 8 (CML 7136) or 9 (CML 7144-6) in males (CML 7144-7
with helmet broken); the first spines are rounded, while the latter are
rather square (Fig. 2A, B); maxillary palpus with the first segment slightly
longer than the fourth (Fig. 2C, E); genal comb with 5 spines, the first
rounded apically, while the other 4 are somewhat square (Fig. 2A, B); the
bulga of the spermatheca rounded (Fig. 2D); phallosome similar to figure
and description given by Johnson (1957).

The distribution of N. rothschildi is known through just a few records
from Peru (Johnson, 1957) and Chile (Beaucournu and Gallardo, 1989) for
N. r. wagneri, and from Bolivia for the nominal subspecies (Wagner, 1938).
Nonnapsylla rothschildi is reported for the first time in Argentina, making
this country the only one where all 7 genera of Craneopsyllinae have been
recorded (Autino and Lareschi, 1998; Beaucournu and Castro, 2003;
Colombetti et al., 2008). Moreover, this is the first report of cricetid rodents
associated with Nonnapsylla. The new records of the present study extend
the southern limits of N. rothschildi to 23u169S and allow us to refine the
distributional limits of the species to an area in midwestern South America
between 23u169 and 16u469S and 64u309 and 70u119W (Fig. 1).

Our specimens of N. rothschildi fit well with the diagnosis from the
original description by Wagner (1938), as well as with the characters
observed in the holotype and allotype of N. r. wagneri. It is notable that
some characters reported for the type specimens do not coincide with
those of the original description given by Johnson (1957) as diagnostic
characters to differentiate the 2 subspecies. For example, in the Argentine
specimens, the helmet is angled in both males and females, as in the types
(Fig. 3A, B), but it is not ‘‘sharply rounded medially’’ in females, as
described by Johnson (1957). Spines of the helmet are variable in the shape
of their tips—the first ones are rounded, and the last ones are square—in
accordance with the types (Fig. 3A, B), but they differ from the drawings
and descriptions given by Johnson (1957, page 64), who said that they are
‘‘somewhat sinuate, not straight dorsally, the dorsal most spine is sharply
rounded, not squared as in Wagner’s figure.’’ The length of the first
segment of the maxillary palpus is slightly longer than the fourth, as in the
types, and it is not ‘‘obviously longer’’ as expressed by Johnson (1957).
Spines from the genal comb and hilla of the spermatheca in the Argentine
specimens are similar to those of the types and agree with the description
of N. r. wagneri; we maintain that no differences can be observed between

these specimens and the drawings given by Wagner (1938). Thus, since no
differences can be found among the characters of N. rothschildi given by
Wagner (1938) and the types of N. r. wagneri, and both compare favorably
with the specimens from Argentina, we consider that at least some of the
diagnostic characters used by Johnson (1957) to differentiate 2 subspecies
are inadequate. Consequently, we conclude that N. rothschildi is a
monotypic species and that only 1 subspecies should be recognized.

We wish to thank Pablo J. Martinez and David A. Flores for their
collaboration in the fieldwork during the doctoral research developed by
M. M. Dı́az, and Luis Giambelluca and Marı́a Cristina Estivariz (both
CEPAVE) for their help with photographs and drawings. Specimens
deposited at the Collection of the Division of Insects of the Field Museum
of Natural History (Chicago, Illinois) were studied by the senior author
during a short visit funded by the Systematic Society of Biology (US), and
CONICET/Comisión Fulbright (Argentina); we thank James H. Boone
and Carl Dick for their assistance. The study was partially funded by
CONICET (Consejo Nacional de Investigaciones Cientı́ficas y Técnicas)
PID 4963, with a fellowship to MMD, and 11N618 (Facultad de Ciencias
Naturales y Museo, Universidad de la Plata, Argentina).

LITERATURE CITED

AUTINO, A. G., AND M. LARESCHI. 1998. Siphonaptera. In Biodiversidad
de artrópodos argentinos. Una perspectiva biotaxonómica, J. J. Morrone,
and S. Coscarón (eds.). Ed. Sur, La Plata, Argentina, p. 279–290.

BEAUCOURNU, J. C., AND D. CASTRO. 2003. Contribution á un inventaire
des puces d’Argentina. Beiträge zur Entomologie 53: 449–479.

———, AND M. H. GALLARDO. 1989. Contribution à la faune du Chili.
Puces nouvelles de la moité nord. Bulletin de la Société Entomolo-
gique de France 94: 181–188.

———, AND ———. 1991. Catalogue provisoire des puces du Chili
(Insecta: Siphonaptera). Bulletin de la Société Française de Para-
sitologie 9: 237–270.

COLOMBETTI, P. L., A. G. AUTINO, G. L. CLAPS, M. I. CARMA, AND M.
LARESCHI. 2008. Primer registro de Cleopsylla townsendi (Siphonap-
tera: Stephanocircidae: Craneopsyllinae) en la Argentina. Revista de
la Sociedad Entomológica Argentina 67: 179–182.

HOPKINS, G. H., AND M. ROTHSCHILD. 1956. An illustrated catalogue of
the Rothschild Collection of fleas (Siphonaptera) in the British
Museum (Natural History). Volume II. Coptopsyllidae, Vermipsylli-
dae, Stephanocircidae, Ischnopsyllidae, Hypsophthalmidae and
Xiphiopsyllidae. British Museum of Natural History, London,
U.K., 445 p.

JOHNSON, P. T. 1957. A classification of the Siphonaptera of South
America with descriptions of new species. Memoirs of the Entomo-
logical Society of Washington 5: 1–298.

WAGNER, J. 1938. Eine neue Art ‘‘helmtragender’’ Flöhe aus Bolivien.
Zeitschrift für Parasitenkunde 10: 132–137.

WHITING, M. F., A. S. WHITING, M. W. HASTRITER, AND K. DITTMAR.
2008. A molecular phylogeny of fleas (Insecta: Siphonaptera): Origins
and host associations. Cladistics 24: 1–31.

FIGURE 2. Optical micrographs of specimens of Nonnapsylla
rothschildi collected in Argentina; bar 5 50 mm. (A) Head of the female
(CML 7144-1). (B) Head of the male (CML 7136). (C) Detail of maxillary
palpus of the female (CML 7144-1). (D) Spermatheca (CML 7969). (E)
Detail of maxillary palpus of the male (CML 7144-6).

FIGURE 3. Drawings of type specimens of Nonnapsylla rothschildi
wagneri; bar 5 50 mm. (A) Holotype. (B) Allotype.

RESEARCH NOTES 955