Sucking lice (Insecta, Anoplura) from indigenous Sulawesi rodents : a new species of Polyplax from a montane shrew rat, and new information about Polyplax wallacei and P. eropepli

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Number 3052, 19 pp., 9 figures, 1 table

10024
December 14, 1992

Sucking Lice (Insecta, Anoplura) from
Indigenous Sulawesi Rodents: a New Species of
Polyplax from a Montane Shrew Rat, and New

Information About Polyplax wallacei and P. eropepli

LANCE A. DURDEN! AND GUY G. MUSSER?

ABSTRACT

Polyplax melasmothrixi, a new species of po-
lyplacid sucking louse, is described from Melas-
mothrix naso, a small-bodied shrew rat known
only from tropical upper montane rain forest in
Central Sulawesi, Indonesia. The male of Polyplax
wallacei is described from specimens collected from
Bunomys chrysocomus trapped in tropical lowland
evergreen rain forest in Central Sulawesi. A further
specimen of Polyplax eropepli, a taxon previously
known only from the type series, is documented

from Eropeplus canus from tropical upper mon-
tane rain forest also in Central Sulawesi. Host and
habitat associations for these three species of suck-
ing lice are discussed. Polyplax melasmothrixi and
P. eropepli are both known only from montane
habitats in Central Sulawesi and both appear to
be host specific (to M. naso and E. canus, respec-
tively). Contrastingly, P. wallacei parasitizes two
species of Bunomys in lowland forests and is known
from North and Central Sulawesi.

INTRODUCTION

Melasmothrix naso, Bunomys chrysoco-
mus, and Eropeplus canus are three murine
rodents found only in forests on the Indo-
nesian island of Sulawesi (Musser, 1987;

Musser and Holden, 1991). The shrew rat,

M. naso, and the large-bodied E. canus have

been recorded only from montane rainforest
formations in the mountainous central part

' Assistant Professor and Assistant Curator, Institute of Arthropodology and Parasitology, Georgia Southern Uni-
versity, Landrum Box 8056, Statesboro, Georgia 30460. Research Associate, Department of Entomology, Smithsonian

Institution, Washington, DC 20560.

? Archbold Curator, Department of Mammalogy, American Museum of Natural History.

Copyright © American Museum of Natural History 1992

ISSN 0003-0082 / Price $2.60

2 AMERICAN MUSEUM NOVITATES

of the island. Bunomys chrysocomus also oc-
curs in montane forest but is most frequently
encountered at lower altitudes in lowland
tropical evergreen rain forest. These three
species of rats are phylogenetically distant
from each other, but they share, in addition
to their endemicity to Sulawesi, an important
trait: they are all parasitized by species of
sucking lice (Anoplura) belonging to the ge-
nus Polyplax.

The descriptions of these lice, along with
notes about the rodent hosts and their hab-
itats, form the body of the present report. We
describe and illustrate a new species of Po-
lyplax based on specimens removed from the
fur of Melasmothrix naso. We also describe
the male of Polyplax wallacei, a louse origi-
nally known only by female examples (Dur-
den, 1987); this ectoparasite has been col-
lected from Bunomys chrysocomus trapped
in lowland evergreen rain forest in north-
eastern and Central Sulawesi, and from Bun-
omys fratrorum, which occurs only in the
northeast arm of the island. Finally, we doc-
ument an additional specimen of Polyplax
eropepli, which until now was only known
from the type series; this louse is apparently
a specific parasite of Eropeplus canus.

Durden and Musser (1991) have summa-
rized the taxa of sucking lice and their hosts
so far recorded from Sulawesi. The present
report extends our knowledge beyond that
summary and is one of a series intended to
document the diversity of ectoparasites, some
of which possess unusual or unique morpho-
logical traits, found on the array of murine
host species that are endemic to the forests
of Sulawesi. This and future papers will con-
tain descriptions of new or poorly known spe-
cies of ectoparasites that have recently been
removed from the skins (both dry and pre-
served in fluid) of Sulawesian rodents housed
in major museum collections.

More than 40 indigenous species of rats
and mice have been recorded from Sulawesi
and its offshore islands (Musser and Holden,
1991). It is important to document the di-
versity of ectoparasites and their host asso-
ciations. Results will form another set of data
that may be significant for testing hypotheses
of zoogeographical relationships among hosts
and their ectoparasites (Durden and Traub,
1990).

NO. 3052

Information derived from these descrip-
tive surveys will also be significant in a wider
context. Eventually, phylogenetic data will be
available for both hosts and parasites, and
hypotheses focusing on host-parasite coevo-
lution can be formulated and tested. The re-
lationships between ectoparasites and their
hosts have always intrigued biologists. Re-
cently, a growing body of literature has de-
scribed coevolution between groups of hosts
and parasites based on comparisons of their
phylogenies (Brooks, 1988; Hafner and Nad-
ler, 1988, 1990; Hellenthal and Price, 1991;
May and Anderson, 1983; Page, 1991; Toft
and Karter, 1990) as well as apparent coevo-
lution that in reality reflects host-induced
morphology of the parasite (Downes, 1990).
Preceding this more complex level of analysis
are surveys simply describing the parasites
and documenting their host species; the re-
sults that we present here reflect that level of
inquiry.

ABBREVIATIONS AND PROCEDURES

Specimens examined and referred to here
are deposited in the collections of the Amer-
ican Museum of Natural History, New York
(AMNH); the Natural History Museum,
London (BMNH); Lance A. Durden (LAD);
Museum Zoologicum Bogoriense, Bogor, In-
donesia (MZB); South Australian Museum,
Adelaide, Australia (SAM); and the National
Museum of Natural History, Smithsonian In-
stitution, Washington, D.C. (USNM).

Other abbreviations used in the descriptive
portions of this work are as follows:

DAcHS Dorsal accessory head setae
DAnHS Dorsal anterior head setae
DCAS Dorsal central abdominal setae
DMHS _ Dorsal marginal head setae
DMsS Dorsal mesothoracic setae
DPHS Dorsal principal head setae

DPoCHS Dorsal posterior central head setae

DPrS Dorsal paratergal setae
DPTS Dorsal principal thoracic setae
DPtS Dorsal prothoracic setae

OrS Oral setae

SHS Sutural head setae

SpAtHS Supraantennal head setae
StAS Sternal abdominal setae

TeAS Tergal abdominal setae
VPaHS _ Ventral preantennal head setae
VPHS Ventral principal head setae
VPrS Ventral paratergal setae

1992

Descriptive format and terminology follow
Kim (1966), Kim and Ludwig (1978), and
Durden and Musser (1991). Drawings of en-
tire lice conventionally illustrate dorsal mor-
phology to the left of the midline and ventral
features to the right. Measurements were
made using a calibrated eyepiece micrometer
inserted into a high-power phase contrast mi-
croscope. Values of measurements and
weights are reported in millimeters (mm) and
grams (g). All rodent hosts discussed here
(Melasmothrix naso, Eropeplus canus, Bun-
omys chrysocomus, and Bunomys fratrorum)
are members of the subfamily Murinae, which
Carleton and Musser (1984) place in the fam-
ily Muridae of Rodentia.

ACKNOWLEDGMENTS

We are grateful to Michael D. Carleton
(USNM) and Paula D. Jenkins (BMNH) who
arranged for various murine study skins un-
der their curation to be examined for ecto-
parasites. Lance Durden’s fieldwork in Su-
lawesi was funded by a grant from the
Committee for Research and Exploration of
the National Geographic Society; he was
sponsored in Indonesia by the Lembaga IImu
Pengetahuan Indonesia (LIPI) and the Royal
Entomological Society of London. Guy Mus-
ser’s field collections in Sulawesi were partly
supported by the Celebes Fund of the Amer-
ican Museum as well as Archbold Expedi-
tions, Inc.; his sponsorship in Indonesia came
from LIPI and MZB. He was also assisted by
members of United States Navy Medical Re-
search Unit No. 2 (NAMRU-2) in Jakarta.
Donna Moore Smith skillfully prepared the
drawings included in this paper. Drs. Richard
G. Robbins and Nixon Wilson critically re-
viewed an earlier draft of the manuscript.

Polyplax melasmothrixi, new species

HOLOTYPE: Male collected by L. A. Durden
from pelt of adult male Melasmothrix naso
(AMNH 225100) collected by G. G. Musser
at 2255 m on Gunung Nokilalaki (1°16’S,
120°10’E), Central Sulawesi, Indonesia on 29
April, 1975. Deposited in the MZB.

REFERRED SPECIMENS: Seven additional
adult lice (2 males, 5 females) all removed
by L. A. Durden from pelts of M. naso from
animals trapped on Gunung Nokilalaki by G.

DURDEN AND MUSSER: SUCKING LICE—SULAWESI RODENTS 3

G. Musser in March and April, 1975. Allo-
type female and one paratype female from
the same host individual as holotype; three
paratype females ex male M. naso (AMNH
225091) at 2255 m; one paratype male ex
male M. naso (AMNH 225105) at 2285 m;
one male ex female M. naso(AMNH 225106)
at 2285 m. In addition, five immature lice
were removed from AMNH 225106. Allo-
type female: MZB. Paratypes: BMNH, LAD,
USNM.

DISTRIBUTION: Known by four collections
from Melasmothrix naso collected at 2225
and 2285 m (tropical upper montane rain
forest) on or close to the summit of Gunung
Nokilalaki (2285 m), Central Sulawesi (Su-
lawesi Tengah), Indonesia.

ETYMOLOGy: The species is named for the
type host genus.

Dracnosis: Polyplax melasmothrixi is dis-
tinctive. It is easily distinguished from all
other species of Polyplax by the unusual ap-
pearance of the male genitalia and by the
setation of gonopods VIII and IX in the fe-
male. The following combination of char-
acters also serves to distinguish this species:
(1) DPHS, DAcHS, and DPoCHS all stout;
DPHS and DAcHS displaced posterolater-
ally; (2) the shape of the thoracic sternal plate;
(3) the presence of some short stout setae, in
addition to longer stout setae, on the abdom-
inal tergites and sternites; (4) the shape of the
paratergal plates and the lengths of the setae
borne on these plates.

DESCRIPTION: Male (fig. 1). Length of ho-
lotype 1.02 mm (mean for series 1.00; range
0.96-1.03; N = 3). Head, thorax, and ab-
domen well sclerotized.

Head. About as wide as long with blunt
anterior apex; 2 DAnHS, 1 DMHS, 1 Ors, 1
SHS (2 present on 1 side of 1 specimen), 1
SpAtHS, and 2 VPaHS on each side; DPHS
relatively short and stout, extending about
half distance to thoracic spiracle, with 1 stout
DAcHS distal to DPHS on each side; both
DPHS and DAcHS displaced posteromargin-
ally; 1 stout peglike DPoCHS on each side;
VPHS long and fairly stout. Antennae 5-seg-
mented with basal segment much larger than
second segment, wider than long; third an-
tennal segment with anterior projection as
typical for males of Polyplax spp.

Thorax. Broad with almost parallel outer

4 AMERICAN MUSEUM NOVITATES

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Fig. 1. Polyplax melasmothrixi, male. A, hab-
itus; B, thoracic sternal plate; C, paratergal plates;
D, genitalia.

Cc

margins; thoracic sternal plate (fig. 1B) ta-
pering to slightly prolonged, rounded, pos-
terior apex, with central, blunt, anterior pro-
jection, and slightly concave, lateral margins;
mesothoracic spiracle moderate in size (0.018
mm in diameter); DPTS short (0.098 mm);
1 DMsS and 1 DPtS on each side. Legs. Mid

NO. 3052

and hind coxae subtriangular; forelegs small,
each with narrow acuminate claw; mid and
hind legs progressively larger and with cor-
respondingly larger acuminate claws.

Abdomen. Slightly wider than thorax; one
broad plate per segment dorsally (on seg-
ments 2-8) and ventrally (on segments 4—7);
two narrow plates ventrally on each of seg-
ments 2 and 3; curving, concave plate ven-
trally on segment 8; most plates with finely
crenulated posterior margins; dorsal plates
on segments 2-7 each with 6-8 thickened
TeAS; dorsal plate on segment 8 with 2 long
TeAS; ventral plates 1-3 and 8 each with 3-
4 StAS; ventral plates 4-7 each with 6-7
thickened StAS; ventral plate 9 with 2 long
marginal StAS; some TeAS and StAS short
and stout. Paratergal plates (fig. 1C) present
on segments 2-8; all plates with 2 apical setae
each, with DPrS slightly longer than corre-
sponding VPrS of same plate: plates I and II
with short setae, plates III-V with setae of
intermediate length, and plates VI and VII
with longer setae; all plates subtriangular;
plates IV and V with both apical angles pro-
duced into points; plates II-VII each with
moderately sized spiracle.

Genitalia (fig. 1D). Basal apodeme deeply
concave posteriorly and with differential scle-
rotization; parameres flanged medially,
notched anteriorly, and appearing to articu-
late with posterior projections of basal apo-
deme; pseudopenis short, extending approx-
imately to apices of parameres, and with three
sclerotized patches.

Female (fig. 2). Length ofallotype 1.18 mm
(mean for series 1.18; range 1.14—-1.24; N =
5).

Head, thorax, and legs. As in male unless
indicated otherwise. Third antennal segment
not modified; thoracic sternal plate (fig. 2B)
posteriorly more prolonged than in male.

Abdomen. With one broad plate dorsally
on each of segments 2, 3, and 8, and two
broad plates dorsally on each of segments 4—
7; two broad plates ventrally on each of seg-
ments 2-7; dorsal plates 1, 10, and 11 each
with four thickened TeAS; dorsal plates 2-9
each with 6-7 thickened TeAS; ventral plate
1 with two thickened StAS, ventral plate 2
with four thickened StAS, and ventral plates
3-12 each with 5-7 thickened StAS; StAS
elongated on ventral plate 12; some TeAS

1992 DURDEN AND MUSSER: SUCKING LICE—SULAWESI RODENTS 5

and StAS short and stout; most tergal and
sternal abdominal plates with fine crenula-
tions along posterior margins. Paratergal
plates (fig. 2C) present on segments 2-8 and
all bearing 2 apical setae each; DPrS longer
than corresponding VPrS on plates I, II, IV,
and V; plates I-III with short apical setae,
plates IV and V with setae of intermediate
length, and plates VI and VII with longer
setae; plates I and II subtriangular; plates IJ-
VI with both apical angles produced into
points; plates II~VII each with moderately
sized spiracle.

Genitalia (fig. 2D). Subgenital plate sub-
triangular; gonopods VIII each with one long
lateral seta and one short seta situated close
to center of gonopod; gonopods IX each with
one short stout seta but with smaller anterior
lobe also bearing short seta; vulvar fimbriae
distinct.

REMARKS: Morphologically, P. melas-
mothrixi is an unusual louse when compared
to its congeners. Some head setae and most
abdominal setae are stout, particularly in the
male. Both Polyplax phthisica Ferris and P.
smallwoodae Johnson, which are parasites of
African: murines (Lemniscomys spp.), simi-
larly possess stout abdominal setae (Johnson,
1960) but otherwise these lice do not appear
to be closely related to P. melasmothrixi.
Genitalic characters of P. melasmothrixi also
are unusual. The male pseudopenis is unique
in possessing three sclerotized distal patches.
Similarly, the proximally notched parameres
that appear to articulate with the distal arms
of the basal apodeme are very unusual. The
setation of the female gonopods VIII and IX
is also distinct.

These and other characters strongly suggest
that P. melasmothrixi is morphologically
unique and not closely related to any other
known species in the genus Polyplax. The
host species, M/. naso, is unique from two
perspectives. It is the only known member
of Melasmothrix. Its closest phylogenetic rel-
atives are the small-bodied shrew rats Ta-
teomys rhinogradoides and T. macrocercus,
also endemic to the mountains of Central Su-
lawesi, and they along with M. naso form a
monophyletic group without apparent close
relatives either on Sulawesi or elsewhere in
the Indo-Australian region (Musser, 1982;
Breed and Musser, 1991). We would also ex-

Fig. 2. Polyplax melasmothrixi, female. A,
habitus; B, thoracic sternal plate; C, paratergal
plates; D, genitalia.

pect the morphology of sucking lice from the
other two species of shrew rats to resemble
that characterizing Polyplax melasmothrixi;
unfortunately, no lice were found on any of
the preserved specimens of Tateomys.

NO. 3052

AMERICAN MUSEUM NOVITATES

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DURDEN AND MUSSER: SUCKING LICE—SULAWESI RODENTS

Fig. 4. Melasmothrix naso, the host of Polyplax melasmothrixi. An adult male caught during the
day on the summit of Gunung Nokilalaki in habitat similar to that shown in figure 3.

Host HAsiTaT: All three species of Su-
lawesian small-bodied shrew rats are known
to occur only in the mountainous central core
of Sulawesi. Records of Melasmothrix naso
are from 6000 ft at Rano Rano (1°30’S,
120°28’E) where the holotype (USNM
219752) was obtained (Miller and Hollister,
1921), and between 6400 and 7500 ft on the
southwest slope and near the summit of Gun-
ung Nokilalaki (1°16’S, 120°10’E) where 35
specimens were collected (Musser, 1982).

Cool, wet, and mossy forest above 6000 ft
is the general habitat where all examples of
M. naso have been encountered (fig. 3).
Among the characteristics of this formation
are the short canopy; small trees representing
species associations and morphologies dis-
tinctive to mountain environments; low di-
versity in species of trees, shrubs, palms, and
other plants; dense moss cover; and high fre-
quency and long duration of rain and mist.
Quantitative details, other descriptive obser-
vations, and other places where the shrew rats
were trapped are documented by Musser
(1982: 71).

Melasmothrix naso is small in body size
with an elongate head and muzzle, small eyes
and ears, chunky body, short tail, elongate
claws on the front feet, and velvety dark
chestnut fur (fig. 4). It is diurnal and terres-
trial, uses mossy and earthen runways con-
cealed beneath moss-covered rotting treefalls
or root entanglements, and apparently feeds
primarily on earthworms and fungus gnats
(Musser, 1982).

Polyplax wallacei Durden

Polyplax wallacei Durden, 1987: 812.

Polyplax wallacei Durden. Durden and Traub,
1990: 57 (zoogeography).

Polyplax wallacei Durden. Durden and Musser,
1991: 7 (faunal relationships).

HOLOTYPE: Female designated by Durden
(1987). In USNM. Allotype male removed
by L. A. Durden from pelt ofa male Bunomys
chrysocomus (AMNH 226924) collected by
G. G. Musser at 1000 m near Tomado (1°19’S,
120°03’E), Danau (Lake) Lindu, Central Su-
lawesi, Indonesia on 22 July, 1973. Depos-
ited in the MZB.

AMERICAN MUSEUM NOVITATES

NO. 3052

Fig. 5.

REFERRED SPECIMENS: Eight additional lice
(4 males, 4 females). Of these, an additional
male and one female were removed from the
same pelt from which the allotype was taken.
One male louse was removed from the pelt
ofa B. chrysocomus female (AMNH 223038)

Polyplax wallacei, male. A, habitus; B, thoracic sternal plate; C, paratergal plates; D, genitalia.

bearing the same locality data as the allotype
host but collected on 21 July, 1975. The re-
maining five specimens, most of which are
in relatively poor condition, were located in
the USNM slide-mounted Anoplura collec-
tions where they had been misidentified as

1992

Polyplax spinulosa Burmeister. These speci-
mens were also collected from Tomado; one
male and two female lice were collected on
22 January, 1972 and bear the field collection
number DJ.M-2366, while two male lice were
collected on 24 January, 1972 (field collec-
tion no. DJ.M-2455). Hosts for both of these
collections were originally labeled on the
slides as ““Rattus penitus penitus,” which we
have since determined to be Bunomys chry-
socomus. Both of these 1972 collections were
made by members of NAMRU-2 teams.
Paratypes: BMNH, LAD, MZB, USNM.

DISTRIBUTION: Known from both North
and Central Sulawesi (Sulawesi Utara and Su-
lawesi Tengah, respectively) from two species
of forest rats belonging to the genus Bunomys.
All known collections from North Sulawesi
are from Dumoga-Bone National Park
(0°34'N, 123°54’E), at an altitude of 230 m,
while those from Central Sulawesi all origi-
nate from Tomado near Danau (Lake) Lindu
(elevation 1000 m).

DIGANOsIs: The female of P. wallacei was
described and diagnosed by Durden (1987).
The male, described in this paper, has mor-
phological similarities to Polyplax spinulosa
Burmeister, a widespread parasite of domes-
tic Rattus. The male of P. wallacei can be
distinguished from P. spinulosa and from
other species of Polyplax by a combination
of the following traits: (1) distinctively shield-
shaped thoracic sternal plate with extended
anterolateral angles (fig. 5B); (2) the shapes
and lengths of the attached apical setae for
the paratergal plates (fig. 5C); (3) the shapes
and proportions of the genitalic structures (fig.
5D).

DESCRIPTION: Male (fig. 5). Length of al-
lotype 0.98 mm (mean for series 0.98; range
0.97-1.00; N = 5). Head, thorax, and ab-
domen well sclerotized.

Head. About as wide as long with anterior
apex broadly rounded; 2 DAcHS (both an-
teromedial to DPHS), 2 DAnHS, 2 DMHS,
1 DPoCHS, 3 OrS, 3 SHS, and 2 SpAtHS on
each side; DPHS almost extending to tho-
racic spiracle; VPHS short and close to in-
sertion of first antennal segment. Antennae
5-segmented with first segment much larger
than second, about as wide as long; third seg-
ment prolonged anterolaterally as is charac-
teristic of male Polyplax.

DURDEN AND MUSSER: SUCKING LICE—SULAWESI RODENTS 9

Thorax. Slightly broader than long; tho-
racic sternal plate (fig. 5B) distinctive with
rounded, central, anterior apex, angulate less
well-developed anterolateral angles, and ta-
pering to broadly rounded posterior apex;
mesothoracic spiracle small (0.016 mm in
diameter); DPTS moderate in length (0.115
mm), extending beyond insertions of first
DCAS; Legs. With subtriangular coxae; fore-
legs with small acuminate claw; hindlegs
slightly larger than midlegs, both with strong,
pointed claws.

Abdomen. Wider than thorax; two anterior
rows of 2 DCAS followed by one broad plate
dorsally on each of segments 2-8; dorsal plates
1-6 (on abdominal segments 2-7) each with
8—10 TeAS; dorsal plate 7 (on abdominal seg-
ment 8) with 4 TeAS; two narrow plates ven-
trally on each of segments 2 and 3, and one
broad plate ventrally on each of segments 4—
7; ventral plates 1 and 3—7 each with 5-7
StAS; ventral plates 2 and 8 each with 4 StAS;
some TeAS and StAS short. Paratergal plates
(fig. 5C) present on segments 2-8; plates I-V
subtriangular, each with two short apical se-
tae; plates VI and VII each with two long
apical setae; DPrS longer than corresponding
VPrS on plates I-III; plates I-VI with dorsal
angles developed into points; plates II-VII
each with moderately sized spiracle.

Genitalia (fig. 5D). Subgenital plate with
medial lacuna and one long lateral seta on
each side of plate immediately anterior to
lacuna; basal apodeme much longer than par-
ameres, forked posteriorly; parameres
smoothly curved; pseudopenis thick, extend-
ing appreciably beyond apices of parameres.

REMARKS: In addition to the new records
from Central Sulawesi recorded here, Durden
(1987) collected the following specimens of
P. wallacei from tropical lowland evergreen
rain forest (altitude 230 m) in Dumoga-Bone
National Park, North Sulawesi, during Feb-
ruary 1985:

2 females ex male B. chrysocomus (LAD no. 24),
2 females ex male B. chrysocomus (LAD no. 32),
2 females ex female B. chrysocomus (LAD no. 51),
4 females ex female B. chrysocomus (LAD no. 69),
1 female ex female Bunomys fratrorum (LAD no.
43),
1 female ex female B. fratrorum (LAD no. 53),
1 female ex female Taeromys sp. (LAD no. 73).

10 AMERICAN MUSEUM NOVITATES

Unfortunately, the last host specimen listed
(LAD no. 73) was not retained as a voucher
but our subsequent examination of collection
records indicates this animal was not a spe-
cies of Taeromys. Consequently, all reliable
known host records for P. wallacei pertain to
Bunomys chrysocomus and B. fratrorum.

Polyplax wallacei may be restricted to pop-
ulations of Bunomys living at low altitudes,
at least between 230 and 1000 m, the ele-
vations from which the louse has been re-
corded. The known hosts, however, are not
similarly ecologically bracketed. In Central
Sulawesi, samples of Bunomys chrysocomus
were collected by Musser from 300 m up
through lowland evergreen rainforest to the
interface with tropical lower montane rain
forest (about 1300 m) and higher into mon-
tane habitats between 1370 and 1515 m
(AMNH 223566-223568, 223570, 225148
are the montane voucher specimens). The
species has also been obtained in montane
formations up to 6000 ft on Gunung Tam-
bussissi (Musser, 1991), above 6000 ft on
Gunung Lehio (USNM 218673 and 218681),
at 2200 m on Pegunungan Latimojong
(AMNH 196578-196584) in the central part
of the island, and at 1500 and 2000 m at
Tanke Salokko on Pegunungan Mekongga in
the southeastern arm (Musser, 1991). Dur-
den unsuccessfully sought lice from the fur
of these montane samples, which consisted
of study skins and fluid-preserved specimens.

In contrast to Bunomys chrysocomus, which
occurs throughout most of Sulawesi (at least
where collectors have sampled) between about
300 and 1800 m (Musser, ms), B. fratrorum
has been found only in the northeastern arm
of Sulawesi east of the Gorontalo region. Most
samples come from lowlands and middle al-
titudes, but the species also reaches montane
forest environments as documented by sam-
ples from 1780 m on Gunung Maujat (SAM
m12644 and m12681) and 2000 m on the
slopes of Gunung Klabat (USNM 217581,
217582, and 217585). We have not surveyed
these animals for lice. That Polyplax wallacei
may be restricted to habitats at low and mid-
dle altitudes in tropical lowland evergreen
rain forest while their hosts are not so con-
fined is the obvious hypothesis to be tested
by sampling freshly caught rats collected in
montane situations.

NO. 3052

Additional survey of the other species of
Bunomys is needed to determine what kinds
of sucking lice they may harbor. For example,
in Central Sulawesi, B. andrewsi occurs in
tropical lowland evergreen rain forest, and B.
penitus is the common Bunomys in montane
habitats except where the mountain B. pro-
latus is found (Musser, 1991; Musser and
Holden, 1991). Samples of B. chrysocomus
have been taken at some of the same places
that yielded specimens of B. penitus and B.
andrewsi (Musser, MS), and close to B. pro-
latus (Musser, 1991). Is Polyplax part of their
ectoparasite fauna and, if so, do they all share
the same species of sucking louse? If Polyplax
wallacei is actually restricted to hosts living
at low and middle altitudes, we would not
expect to find it parasitizing either B. penitus
or B. prolatus but would not be surprised if
it occurred on B. andrewsi, which is mor-
phologically more similar to B. fratrorum than
to any other species of Bunomys (Musser,
MS).

Another facet of the association between
P. wallacei and Bunomys is the low frequency
of louse infestation on B. fratrorum. Durden
(1987: 814) noted that B. chrysocomus ap-
peared to be the primary host of P. wallacei
and that its occurrence on B. fratrorum “could
represent accidental infestations through eco-
logical associations.” This suggestion could
be tested by surveying freshly caught B. fra-
trorum specimens from localities where it does
not occur together with B. chrysocomus. Also,
if B. chrysocomus is really the primary host
and accidental infestation of other species is
a possibility, P. wallacei might also be found
on other species of Bunomys that are sym-
patric with B. chrysocomus.

Whether or not B. fratrorum is an acci-
dental or primary host of P. wallacei, because
the louse has been found on that host and
not on a murine in another genus suggests
evolutionary specificity between P. wallacei
and the genus Bunomys. Throughout its geo-
graphic range, B. chrysocomus occurs in the
same habitat and in Central Sulawesi was col-
lected at the same sites as Rattus hoffmanni,
Maxomys hellwaldii, and M. muschenbroe-
kii, other Sulawesian endemic murines, but
those hosts are parasitized by species of Ho-
plopleura, not Polyplax (Durden, 1990; Dur-
den and Musser, 1991).

1992 DURDEN AND MUSSER: SUCKING LICE—SULAWESI RODENTS 11

ey |
%

+

ca

Pe
i
SF

Fig. 6. Bunomys chrysocomus, the host of Polyplax wallacei. An adult female from Sungai Tokararu,

1150 m.

Hosts AND HABITAT: Bunomys chrysoco-
mus is of medium body size (see measure-
ments listed in tables 1 and 2 in Musser,
1991), with a tail shorter than head and body,
small eyes and ears, along snout, and covered
by dark, soft fur (fig. 6). Musser (fieldnotes
in AMNH) found it to be an opportunistic
feeder, subsisting on insects, earthworms,
snails, small frogs and lizards, and sometimes
fruit.

In Central Sulawesi, Musser encountered
B. chrysocomus most commonly in the wetter
and cooler parts of tropical lowland evergreen
rain forest (fig. 7), usually under dense cover
along the sides of wet ravines and small
streams. It was rarely collected higher on
slopes, ridgetops, or beneath open understo-
ry, where the ambient environment is slightly
drier than that characteristic of ravines and
streamside habitats. At higher elevations
within the boundaries of tropical lower mon-
tane rain forest, B. chrysocomus was also
found in wetter areas of the forest with good
cover.

In its morphology and external appear-
ance, B. fratrorum is a larger version of B.
chrysocomus (compare the values listed in
tables 1-3 in Musser, 1991), and examples
of it have often been misidentified as the
smaller species in museum collections. In ad-
dition to its much larger size, B. fratrorum
has a longer tail relative to head and body
length, the distal portion of the tail is white
in most specimens, and cusp patterns of mo-
lars are less complex. Bunomys fratrorum
likely occurs in similar habitat as B. chryso-
comus because specimens of both have been
collected at the same place (although no in-
formation on microhabitats was provided),
and they may have a similar diet. The stom-
ach of one specimen we studied contained
remains of insects and earthworms.

Polyplax eropepli (Ewing)

Eremophthirius eropepli Ewing, 1935: 209.
Polyplax eriopepli [sic] Ewing. Ferris, 1951: 207
(catalog).

AMERICAN MUSEUM NOVITATES NO. 3052

Fig. 7. Lowland evergreen rain forest along Sungai Tokararu at 1150 m, Central Sulawesi, 1973.
Examples of Bunomys chrysocomus were trapped beneath the dense streamside vegetation.

1992

Polyplax eropepli (Ewing). Johnson, 1958a: 47
(lectotype designated).

Polyplax eropepli (Ewing). Durden, 1°87: 814 (re-
description).

Polyplax eropepli (Ewing). Durden and Traub,
1990: 57 (zoogeography).

Polyplax eropepli (Ewing). Durden and Musser,
1991: 7 (faunal relationships).

REFERRED SPECIMEN: One female louse re-
moved by L. A. Durden from the pelt of an
Eropeplus canus female (USNM 219711)
trapped at 1800 m at Rano Rano, Central
Sulawesi, Indonesia, by H. C. Raven on 15
December, 1917 (Miller and Hollister, 1921).
Deposited in the USNM.

REMARKS: Until now, P. eropepli was
known only from four specimens (two males,
two females) removed from the same pelt
listed here for the new specimen (Ewing, 1935;
Ferris, 1951; Johnson, 1958a; Durden, 1987).
These four specimens are mounted on the
same microscope slide, which bears the
USNM type number 44906. All five P. ero-
pepli specimens are in poor condition, but
the new specimen has retained some intact
structures that are damaged in the type series.
Both female paralectotypes are missing all leg
segments other than the coxae; the new spec-
imen, however, has intact legs and these do
not differ significantly from those illustrated
by Durden (1987) for the male. Other differ-
ences noted for the new specimen are as fol-
lows: (1) a small, anterior, central projection
is present on the thoracic sternal plate; (2) on
paratergal plate VII, one apical seta remains
intact; it is much longer than the correspond-
ing setae on the paralectotypes suggesting se-
tal damage in those specimens; (3) on gon-
opods VIII, the lateral setae (broken on both
sides in the paralectotypes) are long, extend-
ing beyond the outline of the abdomen for
about 40% of their length; (4) on each gon-
opod IX, a single stout seta, rather than two
narrow setae, is present.

Polyplax eropepli has never been collected
from murine species other than Eropeplus
canus and it is probably host specific to that
animal. However, the uniqueness of this as-
sociation has to be tested by surveying Len-
omys meyeri for sucking lice. That murine is
endemic to Sulawesi and is the closest known
relative of E. canus, as revealed by shared
distinctive cranial and pelage traits (Musser,

DURDEN AND MUSSER: SUCKING LICE—SULAWESI RODENTS 13

1981) as well as spermatozoal morphology
(Breed and Musser, 1991). Lenomys is rep-
resented by few specimens in museums, and
unfortunately we have not found lice on any
of those we examined.

Host AND HABITAT: Eropeplus canus 1s a
member of the montane murine fauna en-
demic to Central Sulawesi (Musser and Hold-
en, 1991: 406). Although infrequently en-
countered by collectors, it is documented by
13 examples from the following places: Gun-
ung Nokilalaki (1°16’S, 120°10’E) at 5950 ft
(AMNH 225119 and 225120), 7200 ft
(AMNH 223553, 223554, 223557), 7250 ft
(AMNH 225121-225123), and 7500 ft
(AMNH 223555, 223556, 225124); Rano
Rano (1°30’S, 120°28’E) at 6000 ft (USNM
219711); Gunung Lehio (1°33’S, 110°53’E)
above 6000 ft (USNM 218707, holotype);
Pegunungan Quarles, Bulu Karua (2°56'S,
119°39’E) at 6000 ft (BM 40.386 and 40.387);
and Pegunungan Latimojong (3°30’'S,
121°23’E) at 2200 m (AMNH 196592).

Eropeplus canus is large and bulky with a
very long bicolored tail (fig. 8). Among five
adults from Gunung Nokilalaki, the range in
length of head and body is 230—255 mm,
length of tail is 270-296 mm, length of hind
foot is 46-48 mm, length of ear is 21-27 mm,
and weight is 190-315 g. Long and thick gray
fur covers the body and proximal portion of
the tail.

Tropical montane rain forest (generally de-
scribed by Whitmore, 1984) is the general
habitat of E. canus (fig. 9). Musser (1982) and
Musser and Dagosto (1987) provide descrip-
tions of this environment on Gunung Nok-
ilalaki. Musser trapped the rats on wet ground
near and within rock clusters covered with
moss and held together by tree roots, along-
side rotting trunks, and 15 ft above ground
on a woody vine. Fruits, leaves, palm hearts,
and some kinds of insects were eaten by cap-
tive rats (Musser, fieldnotes in AMNH).

ASSOCIATED MORPHOLOGICAL
DIVERGENCE IN
HOST AND PARASITE

Because evolutionary parallelism at the
levels of genera and species is a relationship
between sucking lice that is widespread (Kim,
1985, 1988), we might expect each species of

14 AMERICAN MUSEUM NOVITATES

NO. 3052

Fig. 8. Eropeplus canus, the host of Polyplax eropepli. An adult female from near the summit of

Gunung Nokilalaki, 7500 ft.

louse associated with the species of Melas-
mothrix, Eropeplus, and Bunomys to also be
unique, possibly at the generic level. Melas-
mothrix naso, the louse found on the shrew
rat, M. naso, is diagnosed by a set of char-
acters unique to it (stout setae, male pseu-
dopenis with three sclerotized distal patches,
proximally notched parameres that articulate
with distal arms of the basal apodeme, and
characteristic setation of female gonopods
VIII and IX), suggesting remote affinity with
any other described relative. However dis-
tinct it is, this louse does not appear to be
part of a monophyletic group different from
that formed by other species of Polyplax, a
new monotypic genus, for example.
Polyplax eropepli, the sucking louse spe-
cific to Eropeplus canus, is not as distinctive
in its morphology as is P. melasmothrixi.
Some characters of P. eropepli are shared with

P. spinulosa, a louse found on commensal
Rattus species (table 1; Durden, 1987; Dur-
den and Page, 1991) that occur on Sulawesi
but are not native to the island (Musser and
Holden, 1991). Polyplax wallacei, which par-
asitizes Bunomys chrysocomus and B. fratro-
rum, also resembles P. spinulosa in some fea-
tures and shares one trait (the general shape
of the thoracic sternal plate) with P. cutchicus,
a parasite of the Indian rat Cremnomys cutch-
icus (table 1; Durden, 1987).

These relationships suggest that the re-
ported evolutionary parallelism between
sucking lice and host taxa cited by Kim (1988)
does not describe the discordant degree of
morphological divergence that exists be-
tween host and louse among certain species
of Indo-Australian murine rodents. Melas-
mothrix naso is in a clade with two other
species of small-bodied shrew rats endemic

15

SUCKING LICE—~SULAWESI RODENTS

DURDEN AND MUSSER

1992

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TABLE 1

Principal Hosts and Known Geographical Distributions of Polyplax spp. Sucking Lice Parasitic on Murine

Rodents?
Louse species Host species Distribution Reference

P. abyssinica Arvicanthus niloticus N. & E. Africa Johnson (1960)
P. acomydis Acomys spinosissimus Mozambique Kim and Emerson (1970)
P. antennata Apodemus mystacinus E. Europe Smetana (1960)
P. arvicanthis Rhabdomys pumilio E. & Southern Africa Johnson (1960)
P. asiatica Nesokia indica Southern Asia & Near East Durden et al. (1990)
P. blanfordi Cremnomys blanfordi India Mishra (1981)
P. brachyrrhyncha Acomys cahirinus N. Africa to Pakistan Durden (1991)
P. bullimae Bullimus bagopus Philippines: Mindanao Johnson (1958b)
P. cummingsi Dasymys spp. Africa Durden (1991)
P. cutchicus Cremnomys cutchicus India Mishra (1981)
P. dacnomydi Dacnomys millardi China: Yunnan Chin (1990)
P. dolichura Acomys cahirinus Sudan Johnson (1962b)
P. eropepli Eropeplus canus Indonesia: Sulawesi Durden and Musser (1991)
P. expressa “Rattus sp.” Philippines: Luzon Johnson (1964)
P. gracilis Micromys minutus Eurasia Beaucournu (1968)
P. grammomydis Grammomys dolichurus Southern Africa Johnson (1960)
P. hoogstraali Acomys spp. Egypt Johnson (1960)
P. humae Cremnomys blanfordi® Pakistan Khan and Khan (1985)
P. indica Golunda ellioti India Mishra (1981)
P. insulsa Leopoldomys sabanus Malaysia, Natuna Is. Johnson (1964)

L. edwardsi¢ China Chin (1980)
P. kondana Millardia kondana India Mishra (1981)

Millardia meltada Pakistan Durden et al. (1990)
P. melasmothrixi Melosmothrix naso Indonesia: Sulawesi Present report
P. meridionalis Acomys spinosissimus4 Botswana Johnson (1962a)
P. oxyrrhyncha Acomys spp. N. & E. Africa Durden (1991)
P. phloeomydis Phloeomys cumingi Philippines: Luzon Cuy (1982)
P. phthisica Lophuromys spp. E. & Central Africa Durden (1991)
P. praomydis Aethomys namaquensis S. Africa & Namibia Johnson (1960)
P. pricei Niviventer confucianus® N. Thailand Kim (1968)
P. serrata Mus musculus Cosmopolitan Durden et al. (1990)
P. smallwoodae Lophuromys spp. W. & Central Africa Durden (1991)
P. solivaga Aethomys chrysophilus S. Africa Johnson (1962b)
P. spinulosa Rattus norvegicus, Cosmopolitan Durden (1991)

R. rattus

Rattus argentiventer S.E. Asia to New Guinea Durden (1987)

Rattus exulans Indo-Pacific region Durden (1987)
P. tarsomydis Tarsomys apoensis Philippines: Mindanao Ewing (1935)
P. thamnomydis Grammomys rutilans Central Africa Johnson (1960)
P. vicina Rattus loseaf China Blagoveshchensky (1972)
P. visenda Leopoldomys sabanus Vietnam Blagoveshchensky (1972)
P. wallacei Bunomys chrysocomus Indonesia: Sulawesi Durden (1987)
P. waterstoni Mastomys spp., Southern Africa Durden (1991)

Praomys spp.

@ An additional 39 species of Polyplax are known. Collectively, they parasitize mammals belonging to eight other

subfamilies of Muridae (in the sense of Carleton and Musser, 1984) and to the families Sciuridae and Soricidae.

6 There are no substantiated records of Cremnomys blanfordi from Pakistan. This murine is endemic to peninsular

India and to Sri Lanka. We assume that the correct type host of Polyplax humae is either Millardia gleadowi or M.
meltada.

© We have not seen the louse or host material reported by Chin (1980) for P. insulsa from L. edwardsi.

4 The type host of Polyplax meridionalis was listed as Acomys cahirinus by Johnson (1962a). However, A. spinosissi-
mus is the only murine of the genus Acomys that is known from Botswana (Dippenaar and Rautenbach, 1986).

¢ The type host of Polyplax pricei was listed as Rattus niveiventer [sic] (=Niviventer niviventer). This murine does
not occur in northern Thailand (Marshall, 1977) and we have since determined the type host to be Niviventer
confucianus.

f We have not seen the louse or host material reported by Blagoveschensky (1972).

1992

to Sulawesi, Tateomys rhinogradoides and T.
macrocercus. Eropeplus canus and Lenomys
meyeri form a separate clade that is phylo-
genetically distant from the group of shrew
rats (Musser, 1982; Breed and Musser, 1991).
Bunomys and Rattus are part of a clade that
includes genera that are native not only to
Sulawesi but to other islands and continents
in the Indo-Australian region (Musser and
Heaney, 1992; Kitchener et al., 1991), and
one that is not phylogenetically closely relat-
ed to either the shrew rat or Eropeplus-Len-
omys clades. Similarly, the Indian rat, Crem-
nomys cutchicus, is not closely related to any
of the endemic Sulawesi murines. Yet it and
species in the other three clades are parasit-
ized by species of sucking lice in the same
supposed monophyletic group, Polyplax.
These and other known examples of murine/
Polyplax host-parasite associations are listed
in table 1. Most of the lice shown in table 1
are clearly host specific (monoxenous). How-
ever, because diverse murine hosts with var-
ious Old World distributions (except for two
species that have accompanied their cos-
mopolitan hosts around the globe) are par-
asitized, host-parasite coevolution in the strict
sense (i.e., Fahrenholz’ rule) does not appear
to have occurred at the genus and species
levels. These observations are based on mor-
phological characters and need to be com-
pared with results from analyses of other kinds
of systems, molecular traits for example.
Some other ectoparasites of Sulawesi rodents
may show clearer host-parasite morpholog-
ical coevolution. For example, many unde-
scribed genera and species of Sulawesian ro-
dent fleas are known, and distinct flea genera
appear to parasitize distinct murine genera
in many cases (Durden and Traub, 1990; R.
Traub, personal commun.).

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