R

Medical and Veterinary Entomology (1999) 13, 447±452

Abrocomaphthirus hoplai, a new genus and species of

sucking louse from Chile and its relevance to
zoogeography
L . A . D U R D E N and J . P . W E B B J r *
Institute of Arthropodology and Parasitology, Georgia Southern University, U.S.A. and *Orange County Vector Control District,
California, U.S.A.

Abstract. Both sexes of Abrocomaphthirus hoplai, new genus and new species
(Anoplura: Polyplacidae), are described and illustrated. The endemic Chilean
chinchilla rat Abrocoma bennetti Waterhouse (Rodentia: Abrocomidae) is the type
host. The de®nition of the family Polyplacidae is amended to accommodate the
new genus. Polyplax longa (Werneck), also referred to in the literature as
Neohaematopinus longus Werneck, is reassigned to Abrocomaphthirus. The host of
A. longus comb.n., is Abrocoma cinerea Thomas, another chinchilla rat, which
inhabits parts of Argentina, Bolivia, Chile and Peru. The erection of
Abrocomaphthirus as a distinct genus has important zoogeographical and
evolutionary implications. The tenuous anomaly of P. longa being the sole native
representative of the genus Polyplax in South America, possibly with African
af®nities, is now refuted. Instead, partial colonization of the neotropics by native
species of both Polyplax and Neohaematopinus appears to have been relatively
recent and from North America. The phylogenetic af®nities of Abrocomaphthirus
are unknown, but it appears to be closely related to other, more ancient, native
South American polyplacid louse genera, such as Cuyana, Eulinognathus,
Galeophthirus, and Lagidiophthirus. Arguments are presented in support of an
ancestral zoogeographical link to Africa for these louse genera.
Key words. Abrocomaphthirus hoplai, A. longus, Polyplax, Anoplura,
Phthiraptera, new combination, new genus and species, neotropics, sucking lice,
zoogeography, Chile, Peru.

Introduction (Burmeister), which parasitize commensal Old World mice

and rats, respectively, have accompanied their hosts in their
Although Vanzolini & Guimaraes (1955a, b) and Traub (1980) colonizations throughout of much of the world, including
provided lucid accounts of the zoogeography and host North and South America (Durden & Musser, 1994a).
associations of neotropical mammal lice, one louse species, Although historical faunal connections between Africa and
Polyplax longa (Werneck), originally described as South America are documented (Goldblatt, 1994), if Polyplax
Neohaematopinus longus Werneck, did not easily conform had accompanied ancestral African rodents that colonized
with their hypotheses. Polyplax is a speciose sucking louse South America, it seems likely that more than one native
genus currently with 79 recognized species (Durden & Musser, species of Polyplax would now be established on the latter
1994a; Chin, 1995) and with most representatives inhabiting continent. Furthermore, collection records of native New
the Old World (Africa and Eurasia) but with three species World species of Polyplax are con®ned to North America and
native to North America (Kim et al., 1986). Two additional
Central America as far south as Panama (Durden & Musser,
species, Polyplax serrata (Burmeister) and P. spinulosa
1994a). In fact, a gradual reduction in the number of species of
Correspondence: Dr L. A. Durden, Institute of Arthropodology and native Polyplax from North America to Panama, from which
Parasitology, Georgia Southern University, PO Box 8056, Georgia just one species is known, suggests that this louse genus has
30460, U.S.A. E-mail: ldurden@gsvms2.cc.gasou.edu colonized the neotropics from the north. A similar north to

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448 L. A. Durden and J. P. Webb Jr

south colonization route has been demonstrated for Polyplax in small sclerites or completely lacking; 5 or 6 pairs of spiracles
eastern Asia (Durden & Traub, 1990). present; tergal and sternal plates usually highly developed and
The discovery of a new species of Chilean sucking louse at times reduced or lacking; sternal plate of segment 2 not
with close morphological af®nities to Polyplax longa, extending laterally to articulate with corresponding parater-
prompted us to compare these two lice and to analyse their gites. Male genitalia with well developed but variously shaped
morphological traits with respect to other polyplacid sucking basal apodeme, parameres and pseudopenis. Female genitalia
lice. Based on this study, we describe the new species and erect with well developed subgenital plate and gonopods VIII and
a new genus to accommodate both species of lice. Next, we IX; spermatheca usually indistinct.
brie¯y discuss the zoogeographical signi®cance of these
taxonomic actions. Finally, we consider any medical or
veterinary importance that these lice may have. Abrocomaphthirus Durden & Webb, gen.n.

Small Anoplura. Head produced into rounded apex;

Materials and Methods antennae 5-segmented, not sexually dimorphic. Thoracic
sternal plate well developed with broadly rounded anterior
Lice were collected from rodents in central Chile by Robert E. and lateral margins and slightly extended posterior margin;
Martin in 1975 and later forwarded to us for analysis. thoracic phragmata distinct; dorsal mesothoracic setae (DMsS)
Specimens were cleared in 10% potassium hydroxide, absent. Forelegs small and weak with small acuminate claw;
dehydrated in a series of ethanol changes of ascending mid- and hindlegs progressively larger, each with larger
strength, further cleared in xylene and then slide-mounted in acuminate claw. Abdomen with narrow tergites and sternites,
Canada balsam, as described by Kim et al. (1986). The new or tergites absent in male; paratergites well developed;
material, and types of Polyplax longa on loan from the paratergites II±V each with 1 or both posterolateral angles
Museum of Comparative Zoology (MCZ), Harvard University, extended; 5 abdominal spiracles ± on paratergites II±VI. Dorsal
were examined and compared by compound light microscopy. lateral abdominal setae (DLAS) and ventral lateral abdominal
In the descriptions that follow, abbreviations established by setae (VLAS) present in both sexes. Male basal apodeme,
Kim & Ludwig (1978) are used but are spelled out when ®rst parameres and pseudopenis well developed; male subgenital
mentioned in this paper. Measurements were made using a plate weakly sclerotized. Female genitalia with small but well
calibrated eyepiece graticule. sclerotized subgenital plate; gonopods VIII and IX distinct;
vulvar ®mbriae indistinct.

Results
Included species. A. hoplai sp.n.; A. longus (Werneck)
Although the new louse genus is clearly a typical polyplacid as comb.n.
characterized by Kim & Ludwig (1978), representatives of the
new genus have ®ve abdominal spiracles rather than six. This Hosts. Chinchilla rats (Rodentia: Abrocomidae) in Andean
trait necessitates amending the characters that de®ne the family South America.
Polyplacidae. Although the number of abdominal spiracles Collection data and deposition of types. (1) For A. hoplai,
present is generally constant for higher taxa of sucking lice, see species description below; (2) For A. longus, types ex
with six being the plesiomorphic (primitive) state, an Abrocoma cinerea Thomas, Puno Department, Peru (Werneck,
apomorphic (derived) reduction in this number occurs in 1948; Ferris, 1951, 1953; Durden & Musser, 1994a), types
representatives of some families such as the Enderleinellidae deposited in the MCZ, Harvard University.
(Kim & Ludwig, 1978). Thus, although the number of Type species. A. longus (Werneck).
abdominal spiracles is a fundamental and constant trait in Etymology. Named for the type host genus, Abrocoma.
most families of sucking lice, intrafamilial variation of this
number is not unprecedented.
Abrocomaphthirus hoplai Durden & Webb, sp.n. (Figs 1, 2)

Family Polyplacidae ± amended description (adapted from Male (Fig. 1)

Kim & Ludwig, 1978) Length of holotype, 0.76 mm. Head, thorax and abdomen
moderately sclerotized.
Medium to small Anoplura. Head with 5-segmented Head. Longer than wide with slightly extended, rounded
antennae, usually sexually dimorphic; postantennal angles apex; 3 dorsal ApHS (apical head setae), 2 ventral ApHS, 2
variable. Thorax with mesothoracic phragmata usually evident; dorsal anterior head setae (DAnHS), 1 supraantennal head seta
sternal plate usually well developed and rarely absent; notal pit (SpAtHS), 2 sutural head setae (SHS), 2 dorsal marginal head
absent. Forelegs always small and slender, each with setae (DMHS), 1 dorsal anteror central head seta (DAnCHS), 1
acuminate claw; mid- and hindlegs subequal in size and shape dorsal posterior central head seta (DPoCHS), 1 dorsal principal
or hindlegs larger than midlegs. Abdomen with paratergites head seta (DPHS), 1 dorsal accessory head seta (DAcHS), 1
usually highly developed and occasionally represented by ventral principal head seta (VPHS), and 1 ventral preantennal

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A new genus and species of louse 449

anteriorly, followed posteriorly by a row of 7 VCAS, then

successively by 6 sternal abdominal setae (StAS), 8 inter-
sternal abdominal setae (InStAS), 8 StAS, 8 InStAS, 6 StAS, 5
StAS, 5 StAS, 4 VCAS, and 2 VCAS; 1 VLAS present on each
of segments 3±7; dorsally, 2 rows of 4 dorsal central
abdominal setae (DCAS) anteriorly, followed successively by
3 rows of 8 DCAS, 2 rows of 6 DCAS, and 1 row of 2 DCAS;
2 DLAS present on each of segments 3±6; ventral abdominal
setae stouter than dorsal abdominal setae. Paratergal plates
(Fig. 1c) present on segments 2±8; plates II-VI each with small
spiracle; plate I with deep medial depression; plates II±IV each
with extended ventral and dorsal apices; plate V with extended
ventral apex; plates VI and VII small and indistinct; plate I
with 2 long dorsal paratergal setae (DPrS), plate II with DPrS
and ventral paratergal seta (VPrS) both long, plate III with
short DPrS and long vPrS, plates IV and V lacking setae, plates
VI and VII each with 2 very long setae.
Genitalia (Fig. 1d). Slightly distorted as shown in ®gure;
basal apodeme slightly longer than parameres and pseudope-
nis, concave posteriorly; parameres stout, well sclerotized, and
with blunt medial indentation; pseudopenis extending approxi-
mately to apex of parameres; subgenital plate (Fig. 1a) poorly
sclerotized and indistinct, U-shaped with small medial
projection on each side.

Female (Fig. 2)
Length of allotype, 0.98 mm; length of paratype female,
0.94 mm; mean length, 0.96 mm. Head, thorax and abdomen as
in male unless indicated otherwise.
Head. 2±3 ApHS present on each side; head distinctly
broader posterior to antennae.
Thorax. Thoracic sternal plate (Fig. 2b) distinctly extended
posteriorly to truncate apex; mesothoracic spiracle diameter,
0.015 mm; DPTS length, 0.08 mm. Legs with fore- and mid
Fig. 1. Abrocomaphthirus hoplai, new genus and species; male (a) coxae subtriangular.
whole louse showing dorsal morphology to the left of the midline Abdomen. Distinctly broader than thorax, with 14 narrow
and ventral features to the right; (b) thoracic sternal plate; (c) plates ventrally and 12 dorsally; ventrally, sternites 1 and 2
paratergal plates; (d) genitalia. broader than other sternites, and sternites 1, 10, 11, 13 and 14
shorter than adjacent sternites; 4 StAS on sternite 1, then 6±8
StAS on sternites 2±9 and 12, and 4±5 StAS on sternites 10,
head seta (VPaHS) on each side; medial SHS slightly longer 11, 13 and 14; 5 rows of 1±2 VLAS present; dorsally, tergites 1
than lateral SHS; DPHS and DAcHS borne on ¯ap-like and 2 each with 4 TeAS; tergites 3±9 each with 6±8 TeAS,
structure. Antennae 5-segmented with basal segment distinctly tergites 10 and 11 each with 4 TeAS, and tergite 12 with 2
larger than other segments, slightly wider than long; third TeAS; 1 DCAS posterior to tergite 12; 11 rows of 1±4 DLAS
antennal segment unmodi®ed. present. Paratergal plates (Fig. 2c) as in male but 1 long DPrS
Thorax. Broad with rounded margins; thoracic sternal plate present on plate II, 1 long VPrS and 1 short DPrS on plate IV,
(Fig. 1b) broadly rounded anteriorly and laterally but with and 1 short seta on plate V.
extended posterior apex; mesothoracic spiracle small Genitalia (Fig. 2d). Small but well sclerotized subgenital
(0.0125 mm in diameter); dorsal principal thoracic seta plate slightly extended anterolaterally on each side, with
(DPTS) moderate in length (0.075 mm); no other thoracic medial indentation posteriorly and 1 small seta on each side;
setae present except on legs. Legs with subtriangular fore gonopods VIII each with 3 apical setae of moderate length;
coxae; forelegs small with narrow acuminate claw; mid- and gonopods IX small, with 2 fairly stout setae (appearing as 3
hind legs progressively larger with correspondingly larger due to the presence of dorsal seta in the same region); vulvar
acuminate claws. ®mbriae indistinct.
Abdomen. Slightly wider than thorax with leathery integu- Collection data. All three type specimens were recovered
ment; 1 narrow sternite ventrally on segments 3, 4 and 6; 2 from different individuals of the chinchilla rat, Abrocoma
narrow sternites ventrally on segment 5; no tergites dorsally; bennetti Waterhouse, trapped by R. E. Martin in Santiago
ventrally, 1 row of 6 ventral central abdominal setae (VCAS) Province, Chile, at an elevation of 900 m. The precise type

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450 L. A. Durden and J. P. Webb Jr

Polyplax longa (Werneck): Ferris, 1953.

Polyplax longus (Werneck): Durden & Musser, 1994a, b.
This louse shares many characters with Abrocomaphthirus
hoplai, sp.n. It also parasitizes a congeneric chinchilla rat in
the same approximate geographical region of the neotropics
(southern Peru). Clearly, this louse is a congener of A. hoplai,
sp.n. For this reason, we have reassigned it to the new genus as
Abrocomaphthirus longus, comb.n.
Since it was ®rst described in 1948, the precise generic
assignment of this louse has been debated. Clearly, it does not
belong to the genus Neohaematopinus because it lacks
morphological characters of that genus such as: (1) a
posteriorly emarginate second tergite on abdominal segment
2; (2) a posteriorly emarginate thoracic sternal plate; (3) a
specialized third antennal segment in the male; (4) character-
istic male genitalia. Although Ferris (1953) reassigned this
louse from Neohaematopinus to Polyplax, a genus with which
it undoubtedly shares several characters, he expressed
reservations about retaining this louse in Polyplax. He stated,
`There remains a question as to what genus it should be
referred or as to whether or not a new genus should be named
for it.' (Ferris, 1953; p. 53). Despite sharing some morpholo-
gical characters with Polyplax, both A. longus and A. hoplai
have ®ve abdominal spiracles rather than the six that
distinguish Polyplax, Neohaematopinus, and other polyplacids.
Males of Polyplax also have a highly modi®ed third antennal
segment, which remains unmodi®ed in males of
Abrocomaphthirus. However, because Abrocomaphthirus is a
typical polyplacid genus in all other respects, we have taken
what we consider to be the most prudent action by slightly
amending the de®nition of the family Polyplacidae in order to
accomodate it. Although other traits of Abrocomaphthirus such
Fig. 2. Abrocomaphthirus hoplai, new genus and species; female (a) as the narrow abdominal tergites and sternites, the relatively
whole louse showing dorsal morphology to the left of the midline large number of DLAS and VLAS and the shape of the
and ventral morphology to the right; (b) thoracic sternal plate; (c)
reproductive structures are unusual, a few species of Polyplax
paratergal plates; (d) genitalia.
share one or more of these characters. However,
Abrocomaphthirus also shares certain morphological charac-
ters with other native neotropical polyplacid genera, namely
locality is 2.5 km north-east of Cerro Manquehue at the site of
Cuyana, Eulinognathus, Galeophthirus and Lagidiophthirus,
a Trappist monastery (33°20¢ S, 70°33¢30¢¢ W). The holotype
and we believe that it is phylogenetically more closely related
male of A. hoplai was collected from an adult female
to these genera than to Polyplax. Its morphological similarity
chinchilla (®eld no. REM 1229) on 1 August 1975, the
to Polyplax may be due, in part, to convergent evolution.
allotype female louse from a subadult female chinchilla (REM
Interestingly, Eulinognathus, as currently interpreted, is well
1189) on 10 April 1975 and the paratype female louse from an
represented in both South America and Africa.
adult female chinchilla (REM 1208) on 1 June 1975.
Deposition of types. Holotype male and allotype female
deposited in the U.S. National Museum of Natural History,
Smithsonian Institution, Washington DC. Paratype female Discussion
deposited in the Natural History Museum, London.
Etymology. This new species is named for our trusted friend Selection pressures associated with parasitizing South
and colleague, Professor Cluff E. Hopla, a renowned authority American abrocomid rodents have resulted in the evolution
on ectoparasites and vector-borne diseases. of several taxa of ectoparasites. In addition to the new sucking
louse genus described here, a unique family of chewing lice,
the Abrocomophagidae, is associated with abrocomids
Abrocomaphthirus longus (Werneck, 1948), comb.n. (Emerson & Price, 1976). Further, the ixodid tick Ixodes
sigelos Keirans, Clifford & Corwin, and the rhopalopsyllid
Neohaematopinus longus Werneck, 1948. ¯eas Delostichus coxalis (Rothschild) and Tetrapsyllus cor®dii
Neohaematopinus longus Werneck: Ferris, 1951. (Rothschild) parasitize A. bennetti and some of its close

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A new genus and species of louse 451

ecological rodent associates in Chile (Keirans et al., 1976; Musser, 1994a). Therefore, if A. longus did belong to the genus
Smit, 1987). Polyplax, there would again be a sizeable gap between Panama
Interestingly, some convergent morphological evolution and southern Peru, from which no native species of Polyplax
seems to have occurred between the sucking louse, A. hoplai, are known. Because native species of Polyplax do not appear
and the chewing louse, Abrocomophaga chilensis Emerson & to have colonized the neotropics any further south than
Price, both of which parasitize A. bennetti. Both lice have Panama, they may not have dispersed further into the region
narrow abdominal plates, relatively abundant lateral abdominal after the Panamanian landbridge formed.
setae, and ®ve abdominal spiracles (on segments 3±7), Unlike Vanzolini & Guimaraes (1955a, b), who did not
presumably re¯ecting evolutionary responses to the same consider continental drift in their zoogeographical analyses,
selection pressures associated with parasitizing A. bennetti. Traub (1980) believed this form of vicariance zoogeography to
The taxonomic action of erecting a new genus for the be important in shaping the distributions of vertebrates and
sucking lice associated with abrocomids has zoogeographical their parasites. He advocated that some neotropical sucking
implications. Foremost, no native species of Polyplax are now lice had African af®nities, highlighting the fact that
known to inhabit South America. Three noteworthy papers Eulinognathus is well represented on both continents and that
have discussed the zoogeography of neotropical mammal lice. three closely related monotypic native polyplacid genera,
The authors of the earlier of these, Vanzolini & Guimaraes Cuyana, Galeophthirus and Lagidiophthirus, also inhabit
(1955a, b), treated A. longus as a member of the polyplacid South America. Although Traub (1980) acknowledged that
genus Neohaematopinus and stated that it re¯ected a recent any physical connection between Africa and South America
invasion of this genus from North America after the predated the evolution of the Anoplura, he believed that certain
Panamanian landbridge formed 2.5±3 million years ago, i.e. ancestral African rodents accompanied by their lice, could
during the `great American interchange' (Vrba, 1992). have `island-hopped' between the two continents when they
Although Neohaematopinus does indeed appear to have were physically closer in the geological past. Given the lack of
colonized Central and South America from North America, rigorous alternative scenarios, this certainly seems to be a
which is inhabited by 10 native species belonging to this genus plausible hypothesis for Eulinognathus and its allies (Cuyana,
(Kim et al., 1986; Durden, 1991), a sharp reduction in the Galeophthirus and Lagidiophthirus) because these louse
number of species in this genus is evident in the neotropics. genera are all absent from North America and other land-
Thus, although 10 species of Neohaematopinus are native to masses with potential relevance to this issue such as
the United States, seven species are known from Mexico, two Antarctica, Australia and India (Durden & Musser, 1994a).
from other Central American countries and just one (N. Because Abrocomaphthirus is also allied to this cluster of
semifasciatus Ferris) from Venezuela (Johnson, 1972b; neotropical polyplacid genera, it seems likely that they share
Durden, 1991; Durden & Musser, 1994a). This, and the ancestral African af®nities. In fact, based on currently
documented Nearctic rodent invasions of South America available data, this seems to be the most likely route of
during the great American interchange, provide circumstantial neotropical colonization by these sucking lice.
evidence that Neohaematopinus has recently in®ltrated the The evolutionary and zoogeographical history of South
neotropics from the north. Thus, if A. longus did belong to the American caviomorph rodents such as the abrocomids may
genus Neohaematopinus as originally believed (Werneck, also favour ancient African origins. Two competing views of
1948), there would be a signi®cant gap between Venezuela late-Eocene caviomorph colonization of South America exist
and southern Peru in which Neohaematopinus is apparently (Vrba, 1992; Wyss et al., 1993); one advocates dispersal of
absent. Although polyplacid louse extinctions could have ancestral forms from North America prior to the formation of
occurred in this region, the above explanation of the derivation the Panamanian landbridge, whereas the other favours
of the extant zoogeography of Neohaematopinus seems more dispersal from Africa. Fossil evidence presented by Wyss
plausible. Further, 30 of the 32 recognized species of et al. (1993) promotes the African connection. Interestingly,
Neohaematopinus parasitize sciurid rodents and the remaining the derivation of the extant neotropical polyplacid sucking
two parasitize sigmodontine rodents (woodrats, Neotoma spp.), louse fauna supports both ancient African origins (for
with none associated with other rodent families such as the Abrocomaphthirus, Cuyana, Eulinognathus, Galeophthirus
abrocomids (Durden, 1991; Durden et al., 1997). Regardless, and Lagidiophthirus) as well as more recent North American
the neotropical distribution for this genus described by origins (for Neohaematopinus and Polyplax). The louse data
Vanzolini & Guimaraes (1955a) is refuted. therefore support the contention that there were two major
Similarly, when A. longus was thought to belong to the waves of rodent colonization into South America; an ancient
genus Polyplax (Ferris, 1953; Durden & Musser, 1994a, b), it one from Africa and a recent one from North America. Future
represented a zoogeographical anomaly because no other phylogenetic analyses of neotropical rodents and their lice,
native species of the latter genus are known south of Panama in particularly with respect to host-parasite cospeciation, should
the neotropics. Similar to the case for Neohaematopinus, a provide a test of these hypotheses.
reduction in the number of native species of Polyplax occurs It is presently unknown whether A. hoplai and A. longus
from the United States (three species), southwards to have any medical or veterinary importance. However, some
Guatemala and Honduras (two species, including an unde- sucking lice of rodents are enzootic vectors of zoonotic
scribed species at hand) and Panama (one species ± P. pathogens such as the causative agents of murine typhus and
auricularis Kellogg & Ferris) (Johnson, 1972a; Durden & epidemic typhus (Traub et al., 1978; Durden & Musser,

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452 L. A. Durden and J. P. Webb Jr

1994b). Related species of sucking lice are vectors of the notes on a fourth species (Insecta; Anoplura). Microentomology, 18,
agents of murine eperythrozoonosis and haemobartonellosis to 19±22.
their rodent hosts (Durden & Musser, 1994b). If rodent Goldblatt, P., ed. (1994) Biological Relationships Between Africa and
ectoparasites are vectors of hantaviruses as advocated by some South America. Yale University Press, New Haven.
Johnson, P.T. (1972a) On the rodent-infesting Anoplura of Panama.
researchers (e.g. Meng et al., 1996), then South American lice
Great Basin Naturalist, 32, 121±136.
such as A. hoplai and A. longus could be enzootic vectors of
Johnson, P.T. (1972b) Sucking lice of Venezuelan rodents, with
these pathogens. Increasing recognition of hantaviral infection remarks on related species (Anoplura). Brigham Young University
of humans in Chile has recently been documented (Espinoza Science Bulletin, Biological Series, 17 (5), 1±62.
et al., 1998; Toro et al., 1998). Keirans, J.E., Clifford, C.M. & Corwin, D. (1976) Ixodes sigelos, n. sp.
(Acarina: Ixodidae), a parasite of rodents in Chile, with a method for
preparing ticks for examination by scanning electron microscopy.
Acknowledgements Acarologia, Paris, 18, 217±225.
Kim, K.C. & Ludwig, H.W. (1978) The family classi®cation of the
We thank Robert E. Martin (McMurry University, Abilene, Anoplura. Systematic Entomology, 3, 249±284.
Texas, U.S.A.) for submitting lice for analysis and Stefan P. Kim, K.C., Pratt, H.D. & Stojanovich, C.J. (1986). The Sucking Lice of
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Pennsylvania State University Press, University Park.
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