30 June 2005
PROC. ENTOMOL. SOc. WASH.
107(3).2005. pp. 604-622
ANOPHELES (ANOPHELES) LESTERI BAlSAS AND HU
(DIPTERA: CULICIDAE): NEOTYPE DESIGNATION AND DESCRIPTION
LEOPOLDO M. RUEDA, RICIIARD C. WILKERSON, AND CONG LJ
Department of Entomology. Walter Reed Army Institute of Research, 503 Robert Grant
Avenue. Silver Spring, MD 20910-7500. U.S.A. (LMR email: ruedapol@si.edu; RCW:
wilkersonr@si.edu: CL: Iicong@si.edu)
Abstract.-The Asian malaria vector. Anopheles (Allopheles) lesteri Baisas and Hu.
1936. is described with illustrations of Ihe larval and pupal stages. adult female, and the
male genitalia. Taxonomic and related literature records, diagnostic features. distribution,
rONA ITS2 sequence. bionomics. and medical importance of All. lested are included. A
neotypc female for the species from near the original type locality in Luzon. Philippines,
is designated.
Key Words:
Anopheles lesteri, Culicidae, taxonomy. description. neotype. mosquitoes.
malaria
Because of the recent increase of human
malaria cases in South Korea (Feighner et
al. 1998. Lee et al. 1998). there is a serious
need to clarify the identity of the possible
mosquito vectors. Misidentifications of vector species often lead to inadequate understanding of the epidemiology of disease
transmission and inadvertently affect control measures. Atlo/'heles (Allopheles) lesteri Baisas and Hu. 1936. may be the most
significant vector of malarial parasites in
Korea. Taiwan, Japan (particularly Okinawa). and central and northern China (Harrison 1973). It is one of the 27 species of
the Hyrcanus Group of Atlopheles (Atlol,heles) having an Oriental or eastern Palearctic distribution (Ramsdale 200 I. http://
www.mosquitocatalog.org). It may have a
potential role in malarial and filarial parasite transmission and disease outbreaks in
countries where it occurs. Recently. Wilkerson et al. (2003) demons(ratcd that All.
allthropophagus Xu and Feng. the mos( important vector malaria vector in eas(em
China. is actually a junior synonym of All.
lesteri.
Allopheles lester; was described by Baisas and Hu (1936: 214) as An. hyrca1llls var.
lested from I male (lot M 1-8) and I female
(lot M 1-12). with corresponding larval and
pupal exuviae, from San(u Mesa. Manila.
Luzon, Philippines (Jose P. Ingal. coli. 2
March 1936). Many specimens (or "colypes") were collected from Calauan, Laguna
in 1935. bu( type specimens were selected
from the Santa Mesa specimens collected in
1936. The syntypcs or type specimens
("types" and '·cotypes". collected from
Santa Mesa and Calauan. Laguna. Luzon).
were supposed to be deposited in the Philippine National Museum. Manila. Other
specimens from the same batch were to be
deposited in the National Museum of Natural History. Washington. DC. and Henry
Lester Institute of Medical Research.
Shanghai, China (Baisas and Hu 1936).
Knight and Stone (1977: 22) noted that (he
type specimens could not be found. Our inquiries were also unsuccessful in search of
these specimens in possible depositories in
Manila and Shanghai. Specimens of All.
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Anopheles (Anopheles) Lesteri Biases and Hu (Diptera: Culicidae):
Neotype Designation and Description
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605
VOLUME 107. NUMBER 3
lesteri found in the National Museum of
Natural History include 2 slides with the
following labels: slide # 1- right label light
yellow, with reddish border: "M 1-13 An.
hyr. var. lesteri Sta. Mesa, Manila Mar. 5.
1936", left label: "Anopheles (An.) lesteri
lesteri Baisas & Hu del. B.A. Harrison";
slide # 2-right label light yellow, with reddish border: "F 171-50 An. lesteri cotypemale Calauan, Laguna July 15, 1935."
However, larval and pupal exuviae, particularly those mounted on slide # 2, are in
bad conditions (i.e.• dark unknown mounting media, cracked specimens. etc.), and
most morphological characters are difficult
to see under the compound microscope.
Also. the original adult description (Baisas and Hu 1936) is not sufficiently complete
for accurate identification of the species.
and no detailed descriptions of the larval
and pupal stages or of the male terminalia
of specimens from the type locality (Luzon)
have been made. In view of this, it becomes
imperative to provide detailed descriptions
of various life stages and to designate a
neotype for this important species.
In the present paper, a neotype female
from near the original type locality is designated for An. lesteri and associated ribosomal DNA internal transcribed spacer 2
(rONA ITS2) sequence provided. Descriptions and illustrations are provided for the
adult female and male. pupa and larva of
this species from the same type locality.
MATERIALS AND METHODS
Except for wing spot nomenclature.
which is taken from Wilkerson and Peyton
(1990), for wing venation terms from Belkin (1962). and for pupal abdominal dark
marks. the terminology and abbreviations
of Harbach and Knight (1980, 1982) are
used for the morphological characters and
illustrations. Abbreviations used are as follow: E, egg; G, genitalia; L, larva; Le, larval exuviae; NE. non-existent; P. pupa; Pe,
pupal exuviae; var., variety. An asterisk following the abbreviation of a given life stage
indicates that at least part of the life stage
was illustrated in the publication cited. Collection codes of the most recent collections
consist of a country code in capital letters
followed by a collection number (e.g., PH
9-1 is an individual from collection 9 from
the Philippines; a specimen number lower
than 100 is used if there are associated larval and pupal exuviae, and equal or greater
than 100 if there are no associated larval
exuviae).
DNA isolation and sequencing. DNA
was isolated by phenol-chloroform extraction. as described by Wilkerson et a!.
( 1993), from a leg of the adult neotype female, a leg of a second female, and 2 entire
males. minus genitalia. from new type locality in Calauan, Laguna. Direct sequencing was carried out as described in Wilkerson et al. (2004) using their primers. The
beginning and end of the rDNA ITS2 was
estimated as in Cornel et a!. (1996).
TAXONOMIC TREATMENT
Anopheles (Anopheles) lesteri
Baisas and Hu
(Figs. ]-4)
Anopheles yesoensis Tsuzuki 190 I: 717
(nomen dubillm).
Anopheles je.mensis Tsuzuki 1902: 286 (nomen dubium).
Anopheles hyrcanlls var. lesteri Baisas and
Hu 1936: 229 (2, P*, L*, E*). Type:
Santa Mesa. Manila (Luzon), Philippines
(NE). other specimens!"cotypes": Calauan, Laguna (Luzon); Ohmori 1957:
209 (0*. E*); Ohmori 1959: 222 (P*).
Anopheles (Anopheles) lesteri: Otsuru and
Ohmori 1960: 47 (0*. 9*, P*, L. E*;
taxonomy; bionomics; distribution. Japan: Honshu [Hyogo, Mie, Niigata, Yamaguchi], Kyushu [Kumamoto. Kagosirna, Oita. Fukuoka, Saga. Nagasaki]);
Whang 1962: 39 (distribution, Korea:
Tansan, Wondang, Guidandong. Yongjueup); Reid 1968: 105 (type form); Cagampang-Ramos and Darsie 1970: 14
(identification key); Basio 1971: 36 (distribution, Philippines: widely found in
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PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Luzon including Manila. Pampanga);
Basio and Reisen 197 1:60 (L. distribution, Guam); Tanaka 197 I: 4 (distribution. Japan: Ryukyu Islands); Harrison
1973: 4 (taxonomy); Baisas 1974: 50
(9*. P, L*. E; taxonomy); Tanaka et al.
1979: 83 (0* 9*. P*. L*, E; taxonomy,
bionomics. distribution. Japan: Hokkaido. Honshu, Kyushu. Ryukyu Archipelago LAmami. Okinawa Gunto. Yaeyama
Gunto]); Rueda et al. 2004 (distribution.
China: Hong Kong).
Anopheles (Anopheles) lester; lester;: Chau
1982 (distribution, China: Hong Kong).
Anopheles (Anopheles) lester; anlhropophaglls Xu and Feng 1975: 81.97 (9*.0*.
P*, L*. E*; taxonomy).
Anopheles (Anopheles) an thropoplwglts:
Ma 1981: 1I (key; distribution, China:
Fukien. Kiangsi. Kiangsu. Kwangsi.
Kweichow. Shanghai. provinces south of
Yantze River); Wilkerson et al. 2003: I
(new synonym of lester;).
Other literature records.-Otsuru 1949:
139 (as possible malaria vector. Japan); Otsuru and Ohmori 1960: 33 (bionomics, Japan); Kamimura 1968: 15 (as possible malaria vector. Japan); Reisen et al. 1972: 319
(distribution, Guam); Zhang et al. 1980:
140 (as experimental vector of Vietnam
monkey malarial pamsite. near Plasmodium
cynomolgi); Xu et al. 1981: 265 (scanning
electron micrographs [SEMs 1 of adults. pupae. eggs, China); Takai et al. 1984: 251
(hybridization. Japan); Zhang et al. 1987:
191 (vectorial capacity for malayan filariasis. Sichuan. China); Xu et al. 1988: 247
(control using insecticide-treated bed net.
Guangxi. China); Ma and Wang 1988: 65
(salivary gland chromosome, China); Wang
and Zheng 1989: 175 (blood meals, Guizhou, China); Ye and Zhu 1989: 256 (enzyme
electrophoresis. China); Dapeng et al. 1996:
100 (as vector of P. jalciparum. and chemical vector control. Xinyang. China); Li et
al. 1991: 8 (DNA-restriction fragment
length differences. China); Liu et al. 1991:
147 (as vector of malayan filariasis. Fujian.
China); Niu et al. 1992: 267 (DNA probe);
Chen et al. 1994: 27-30 (trace and macro
elements in hemolymph); Cheng et al.
1995: 321. (control using insecticide-treated
bed net. Henan, China); Gu et al. 1966: 34
(distribution, China); Shahjehan 1996: 205
(DNA probes. China); Song and Peng
1996: 161 (control using mermithid nematodes. Sichuan. China); Xu et al. 1997: 807
(as vector of filariasis, Henan. China);
Sleigh et al. 1998: 265 (as P. vivax vector.
Henan. China); Xu et al. 1998: 135-136 (as
vector of P. v;vax; control using insecticidetreated bed net and residual spraying, Hubei, China); Kim et al. 1999: 181 (seasonal
prevalence. South Korea); Zizhao et al.
1999: 240-242 (as vector of P. jalciparum
malaria, China); Lee et al. 2000: 77 (PCR.
presence of P. vivax circumsporozoite protein. South Korea); Ma et al. 2000a: 325
(PCR assay and rDNA-ITS2 sequencing.
China); Ma et al. 2000b: 36 (rDNA-ITS2,
Korea); Burkett et al. 200 I: 196, 2002: 45
(trap attractants, South Korea); Huang et al.
200 I: 340 (habitat and distribution, Hubei,
China); Coleman et al. 2002: 244 (presence
of P. vivax circumsporozoite protein, South
Korea); Min et al. 2002: 198; Shin et al.
2002: 41 (vector competence to P. vivax.
Korea); Toma 2002: 7 (distribution review,
Ryukyu Archipelago, Japan); Wilkerson et
al. 2003: I (species molecular confirmation.
rDNA-ITS2, China. Philippines. South Korea; note on geographical range. China).
ORIGINAL DESCRIPTION
In support of previous and present interpretations of the name lesteri, the original
description given by Baisas and Hu (1936)
is as follows. "Dark and pale scales of
wings well contrasted. Costa-Jet black excepting for the preapical and subcostal pale
spots. The subcostal spot is composed of
from IOta 22 pale scales. No scattered pale
scales elsewhere on the dark portions of the
costa. Subcosta-Invariably dark excepting
for I to 4 pale scales at the upex. which
form a part of the subcostal spot. Vein 1Preapical pale spot distinct and complete.
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VOLUME 107, NUMBER 3
Subcostal spot usually incomplete, seldom
complete, and more rarely absent. Mid pale
spot usually small and incomplete, sometimes absent. Sector pale spot usually small
and incomplete. A few scattered pale scales
are sometimes present on the dark area between the preapical and subcostal spots but
these are not as many as those found in nigerrimus. A few pale scales towards the
base below the presector dark spot. which
do not, however, form definite spots. Vein
2-Stem mainly pale with some greyish or
dark scales on lateral borders. Anterior fork
dark with complete preapical pale spot.
Posterior fork dark with a pale spot at
about, or a little below the middle. Vein 3A definite dark area, at base, and another at
apex; apical half of intervening area with
median squames mainly pale; lateral
squames mainly dark; basal half mainly
pale with a few scattered dark scales. Vein
4--Stem dark or mainly dark towards base
below cross viens, mainly pale towards
apex. Forks dark at bases and apices, the
intervening portion mainly pale hut lateral
squames with fair distribution of dark
scales. Vein 5-Apical half or more of stem
pale. sometimes with a few scattered dark
scales. A definite dark area a little below
middle, followed by a mixture of dark and
pale scales, the pale ones sometimes predominating or occupying the whole area excepting the extreme base where a few dark
scales are located. Anterior fork mainly
dark with the usual dark spots iII-demarcated. Sometimes the basal and sub-basal
dark areas are formed on the pale portion
towards the apex. Posterior fork pale excepting for the apical dark spot. Vein 6Pale with a dark area at middle, and another
one at the apex. In some specimens. a few
dark scales are sometimes scattered on the
pale portion towards the base. Fringe-Pale
spot at apex of wing involves variable portion of area opposite vein I and vein 4.1.
Pale spot opposite vein 5.2 absent in all
males and females examined. Humeral
vein-Usually with 2 or 3 dark scales. Re-
migium-Mainly dark-scaled with a few
pale scales on the anterior border."
SUPPLEMENTAL DESCRIPTION FROM
THIS STUDY
Female (Fig. I ).-Integument dark
brown with silvery or grayish pollinosity.
The following measurements and counts. II
= 6, except when indicated. Head: lnterocular space with 10-11 (n = 3) long, pale
setae intermixed with long and small, narrow, appressed white scales; vertex, occiput
and upper portion of postgena with numerous erect, truncate scales; patch of white
scales on the middorsal portion of vertex;
patch of dark brown to black scales on lateral portion of vertex, occiput and upper
portion of postgena; long dark brown to
black setae on ventral portion of postgena.
Clypeus bare dorsally, with dark scales laterally. Pedicel of antenna with 6-11 (11 =
4) small, dorsolateral, narrow to broad,
grayish white spatulate scales, and 3 short,
dark ventrolateral setae; Ragellomere I with
numerous narrow to broad white scales; remaining flagellomeres with a few scattered
narrow to broad white scales. Scales of
maxillary palpus slender, spatulate, mostly
dark brown to black with intermixed dark
brown setae; narrow band of white scales
at base of palpomeres 3 and 4, and at base
and apex of palpomere 5; apical white band
of palpomere 5 slightly shorter than other
basal palpomere white bands; base of maxillary palpus dorsally with single long, erect
dark seta; length of maxillary palpus 1.751.98 mm (mean = 1.86 mm); ralio of length
of each of palpomeres 2-5 to total length
of palpus. 2 = 0.33-0.34 (mean = 0.34), 3
= 0.32-0.37 (mean = 0.35),4 = 0.17-0.23
(mean = 0.20), 5 = 0.1-0.12 (mean =
0.1 I); ratio of combined palpomeres 2-5 to
total length of palpus. 0.99-1.0 (mean =
0.99); ratio of combined palpomeres 4, 5 to
total length of pal pus, 0.27-0.35 (mean =
0.31); palpus 1.15-1.26 (mean = 1.22) forefemur length. Proboscis dark-scaled, except labellum light brown; base of proboscis with long, erect dark setae and scales;
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
608
I
1G
1
Fig. I.
Anopheles lesteri. adult female. habitus.
proboscis length 1.75-1.95 mm (mean =
1.86 mm. If = 3); proboscis 0.99-1.01
(mean = 1.0." = 3) palpus lenglh. Thorax:
Scutum dark brown. with gray pollinosity;
2 submedian longitudinal lines on anterior
area; a pair of indistinct black spots near
antealar area; median anterior promontory
with patch of intermixed narrow. short and
long pale yellow scales. Darker lateral areas
of scutum with longer dark setae. Scutal
fossa. antealar area and supraalar area
slightly pale pollinose. Scutellum dark.
slightly pale dusted. with 16-22 shorter and
14-19 long setae. short setae i ntennixed
pale yellow and dark brown. long setae
dark brown. Antepronotum with 12-14
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VOLUME 107. NUMBER 3
Table I. Anophele.f lester;: descriptive statistics for ratios of veins C and R-R. wing spot lengths/length of
wing measured from humeral crossvein*.
wゥョセ
spot
Costa (C)
Basal dark to sector dark (BD+PHD+HD+PD+SD)
Subcostal pale (SCP)
Preapical dark (PO)
Preapical pale (PP)
Apical dark (AD)
Mean:!: SO
Range
0.69-0.74
0.03-0.07
0.27-0.31
0.03-0.05
0.04-0.06
0.72
0.05
0.29
0.04
0.05
:t 0.02
:!: 0.01
:t 0.01
:t 0.01
:t 0.01
[0.73]
[0.05]
[0.29]
[0.03]
[0.04]
0.38-0.41
0.04....0.13
0.21-0.26
0.05-0.06
0.24-0.29
0.04-0.06
0.03-0.05
0.40
0.08
0.24
0.05
0.26
0.05
0.04
:t 0.01 [0.38J
:t 0.04 [0.12]
:!: 0.02 [0.21]
:t 0.00 [0.06]
:!: 0.02 [0.25]
:t 0.01 [0.06]
:!: 0.01 [0.03]
Vein R-R.
Basal dark to presector dark (BD+PHD+HD+PSD)
Sector pale (SP)
Sector dark (SO)
Subcostal pale (SCP)
Preapical dark (PD)
Preapical pale (PP)
Apical dark (AD)
* Six
wings. from the ncotypc and alloneotype. and 3 individuals: [ ]. neotypc female.
dark brown setae. Pleuron brown to dark
brown; upper proepistemum with 3 or 4 setae. without scales; prespiracular area with
2 or 3 setae. without scales; prealar area
with 4 or 5 setae. without scales; upper mesokatepistemum with 3 or 4 setae. without
scales; lower mesokatepistemum with 4 or
5 setae. without scales; upper mesepimeron
with 4 or 5 setae. without scales. Legs:
Fore- and midlegs dark-scaled except white
scales dorsally on apex of tibia; complete
narrow apical pale bands on tarsomeres 13. and very narrow apical dorsal pale patch
on tarsomere 4; apical bands on tarsomeres
2 and 3 longest. complete ventrally. about
O. I length of tarsomere; pale scales on tarsomere 4 dorsally and laterally only. not
connected ventrally; basal pale band on tarsomere 5 absent. Hindlegs dark-scaled as
fore- and midlegs. except white scales on
tarsomeres 1-3 dorsally and ]aterally only.
not ventrally. Forefemur length 1.40-1.72
mm (mean = 1.53 mm. II = 6). ratio of
forefemur length to proboscis length 0.790.88 (mean = 0.82). Midfemur length
1.74-2.12 mm (mean = 1.94 mm). ratio of
midfemur length to proboscis length 0.931.12 (mean = 1.04). Hindfemur length
2.01-2.2 mm (mean = 2.11 mm). ratio of
hindfemur length to proboscis length 1.071.21 (mean = 1.\4).
Willg (Table 1): Length (measured from
humeral cross vein to wing tip. excluding
fringe) 2.8-3.18 mm (mean = 3.05 mm).
Dark scales brown to black. pale wing
scales white and pale yellow. Costa (C)
dark-scaled with small subcostal pale spot
(SCP' mean = 0.15) and preapical pale spot
(PP, mean = 0.12); remigium dark scaled;
humeral crossvein and arculus without scale
patch. Subcosta (Sc) dark-scaled with few
scattered spatulate white scales from base
to sector dark (SO). Radius (R) to R) darkscaled except 3 pale spots (SP' SCP and
PP). scattered white spatulate scales from
base to presector dark (PSO). and a stripe
of white scales on SO before SCP; length
of white stripe 0.3-0.38 mm (mean =
0.33); base of R,. dark-scaled; bifurcation of
R! and R) dark-scaled; tips of R I • R!. R)
and R4+ 5 with pale fringe. Media (M) darkscaled with pale area before fork; bifurcation of M 1... 2 and M3+4 dark scaled; tips of
M I ... 2 and M 3 + 4 with dark fringe. Cubitus
(Cu) with basal dark spot. length 0.18-0.28
(mean = 0.24); CUI with 4 dark spots,
length of first basal spot 0.1-0. I8 mm
(mean = O. 14). second spot 0.23-0.25 mm
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PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHTNGTON
(mean
0.24), third spot 0.25-0.45 mm
(mean = 0.33), distal fourth spot 0.13-0.3
mm (mean = 0.21); CU 2 with distal dark
spot only, length 0.18-0.25 mm (mean =
0.21); tips of CUI and CU2 with dark fringe.
Anal vein (I A) with 2 dark spots, basal spot
length 0.23-0.25 mm (mean = 0.24), distal
spot length 0.25-0.35 mm (mean = 0.31),
tip of IA with dark fringe. Haller: Scabcllum, pedicel and capitellum dark brown
with grayish pollinosity. Abdome,,: Terga
and sterna dark brown to black with grayish
pollinosity, covered with pale brown to
golden brown setae. For neotype female,
descriptive statistics for ratios of costal and
R-R J wing spot lengths/length of wing measured from humeral crossvein are shown in
brackets in Table I.
Male (Figs. 2C, D).-As in female except for following sexual differences. Maxillary palpus 0.94-0.98 length of proboscis
(mean = 0.97; " = 4 for this and following
measurements except where indicated),
apex of palpomere 3 and all of palpomeres
4 and 5 enlarged. Maxillary pal pus with
dark brown and white scales, dorsal surface
of all segments with white scales; palpomere 2 with slightly erect dark brown scales
at base and light gray scales from middle
to apex; palpomere 3 dark-scaled with long
yellowish to light brown setae at apex; palpomere 4 pale yellow to dark brown-scaled
with narrow basal white band, inner surface
with long yellowish-brown to light brown
setae; palpomere 5 pale brown-scaled with
narrow basal white band, lateral surface
with white scales and numerous dark brown
short setae. Proboscis length 2.65-2.9 mm
(mean = 2.69 mm), dark brown-scaled.
Anal vein with single dark spot. Tergum IX
(width, 2.92 mm) bare, with pair of elongate caudally directed capitate lobes; length
of lobe from base to tip 0.82 distance between 2 lobes; middle of lobe narrower,
0.45 width of lobe tip and 0.42 width of
lobe base. Gonocoxite 1.91-2.13x as long
as wide at widest point, widest about 0.08
from base; dorsal (postrotational) surface
with many long setae distally, slender fu-
sifonn and spatulate scales and numerous
small spicules proximally; ventral surface
as dorsal but with lateral scales and numerous longer spicules; mesal parabasal spine
(parabasal I) stout, borne on slightly raised
base; parabasal 2 stout with slender tip; parabasal I base 0.07-0.15 from base of gonocoxite; parabasal 2 base O. 14-0.15 from
base of gonocoxite; internal seta slender,
about as long as parabasal 2, base 0.410.46 distance from base of gonocoxite.
Gonostylus widened at base and narrowed
toward middle and tip, base 2.27x wider
than middle or tip; gonostylus 0.47 length
of gonocoxite; gonostylus 8.89X longer
than gonostylar claw. Claspelle. Dorsal
lobe of c1aspette with 3 closely appressed
setae of about equal length; tips of 2 lateral
setae curved and bluntly rounded: tip of
middle seta slightly curved and round; tip
of composite structure club-shaped. Ventral
lobe of c1aspette with 2 long subapical setae, most apical much longer than other.
Both ventral and dorsal lobes, and areas in
between them, with numerous spicules. Aedeagal leaflets 4 per side; 2 most mesal leaflets broadest, with broad, thin, nearly transparent inner margins; other leaflets with
narrow, thin, nearly transparent inner margins.
Pupa (Figs. 2A, B).-Position and development of setae as figured; range and
modal number of branches, and number of
branches of neotype female as in Table 2.
Integument darkly pigmented. Exuviae colorless to dark brown. Cephalolhorax: Mesothoracic wing with checkered dark
stripes; metathoracic wing pigmented on
middle, ventral spiracular sensilla distinct.
Trumpet with dark thickend areas bearing
saw-toothed or serrate edge, meatus with
simple cleft, and its subbasal part with numerous spinules: trumpet length 0.30-0.45
mm (mean = 0.35 mm, n = IO for this and
following measurements and counts except
where indicated), width 0.16-0.23 mm
(mean = 0.13 mm, measured at base of pinna), index 1.36-2.5 (mean = 1.73); tracheoid area 0.45 length of trumpet. Abc/omen:
VOLUME 107. NUMBER 3
611
Fig. 2. Allopheles lesteri. (A) Pupa. ccphalolhorltx. (B) Pupa. melalhornx and abdomen. left side dorsal. right
side ventral. IC) Male genitalia. (D) Tergum IX. Abbreviations used include: AAOM = anterior accessory dark
mark. ADM = anterior dark mark, CT = cephalothorax, GL = genital lobe, Pa = paddle. PADM = posterior
accessory dark mark. POM = posterior dark m,uk.
Table 2. Pupal setal bmnching for Allopheil'S lesteri: mnge (mode) based on counts made on 5-10 setae of the neotype, alloneotypc. and 3 specimens collected
with them: [ ], neOlype female.
Sc,.
No.
Ahdominal s」セュョl
Ccphalothordx
IT
II
III
tV
v
15-23(21) 11K, 16)
4-7(5) (4. 5)
3-7(3) [6. 71
2-7(5) (5)
2-5(2) (21
1-3(2) l21
2-3(3) 13)
2-4(3) 141
3-9(6) 18. 61
4-10(8) [5. 7]
1-3(2) (2. 3)
1-4(3) 131
2-4(3) 13.41
1-2(1) [I)
2-4(3) 131
2-4(3) 13)
13-19(13) [13.141
5-9(8) [8. 9)
4-7(5) 17. 61
1-4(1) [11
10-16(13) 112.131
1-4(2) (2)
1-4(1) (3)
2-3(2) [2-3)
3-4(4) 14)
7-13(12) [12. 9)
3-7(5) 17)
34)(5) 15. 41
1-4(2) (3)
11-21(13) [12. IS)
1-2(2) (2)
2-3(2) [3]
2-3(3) [3]
1-2(2) (21
I (II
2-4(3) [31
3-6(6) (6. 3)
4-7(5) 151
1-2(2) 121
1-3(3) (3)
13-24(13) [18.191
1-2(1) [I. 21
2-3(2) (3]
1-3(3) [3]
I [I]
I II)
1-2(1) I II
o
I
2
3
4
5
6
7
8
9
10
II
12
13
14
2-3(3) 12. 3)
2-3(2) 12)
1-3(1) II. 2}
1-3(2) (2)
1-5(4) (4)
1-2(2) [2. I)
1-2( I) [I)
1-2(2) 12)
1-2(2) [2]
2 [21
1-4(4) 14. 3)
1-3(3) (31
I (I)
I II)
1-2(2) [2]
1-2(1) (I)
1-2( I) II]
1-2(1) II. 2)
1(-]
Abdomina)
セZ」s
s」セュ ョエ
Vt
VII
VlII
o
2-4(3) (4)
2-5(3) [4]
3-7(5) (7. 6)
1-2(1) [I. 2)
1-3(1) [II
8-17(10) [8. tOl
1-2(1) (I I
1-3(1) II)
1-3(2) 131
I [I]
I [I)
1-2(1) [I I
2-4(3) (2-31
2-3(2) [2]
2-4(4) [41
1-3(2) [11
1-2(1) (I. 2J
3-7(4) ((}-7)
1-2(1) (I)
1-3(1) [I)
2-3(2) 12-3]
1 [I]
1-4(1) [I)
1-2(1) (II
I
2
3
4
5
6
7
8
9
10
II
12
13
14
...
....uldl<
No.
IX
I [I)
1-2(2) [21
I [I]
1-3(\) [1-21
I [I]
I [I]
I [I]
VOLUME 107. NUMBER 3
Abdominal tergum 1 with 2 anterior dark
marks (ADM), and 2 elongate posterior
dark marks (PDM); each PDM narrows
mesally at base, with maximum width
(0.016-0.02 mm, mean = 0.019) towards
distal tip, length 0.14-0.31 mm (mean =
0.26, n = 10), about 0.12-0.28 (mean =
0.24, n = 10) width of abdominal segment
I, and longer than the longest branch of seta
I-I. Abdominal terga II-VII with I ADM,
2 anterior accessory dark marks (AADM),
and 2 posterior accessory dark marks
(PADM); VIII with I ADM and no AADM
and PADM; cuticular reticulations distinct
on II-IV; spinules scattered mostly on anterior 0.37 of dorsal and lateral sides of VII
and VIII. Seta I-I fan-like with 15-23 aciculate dendritic branches; 6-1 with 1-3
branches; 7-1 with 2 or 3 branches; 9-1 with
I or 2 branches. Setae I, 5-II-VII well developed; I-V 1.04-1.31 (mean = 1.16, n =
4) length of 5-V; I-VI 0.83-1.09 (mean =
0.97, " = 4); I-VII 1.84-2.22 (mean =
1.09, n = 6); 3-VI aligned with and mesal
of I-VI unlike on other segments; 8-1-11 absent; 9-1 simple, single or forked; 9-11 very
short, simple, spine-like; 9-111 short, with
slightly rounded tip. 1.50-4.00 (mean =
2.51) length of 9-11; 9-IV strong, lightly
pigmented and slightly pointed, 01.00-2.80
(mean = 1.83) length of 9-11I; 9-V-VIII
long, lightly pigmented and slightly pointed; 9-V 1.00-1.50 (mean = 1.68) length of
9-IV: 9-VI 0.81-1.33 (mean = 1.02) length
of 9-V: 9-Vn 1.00-1.34 (mean = I.J I)
length of 9-VI; 9-VIII 0.80-1.06 (mean =
0.96) length of 9-VII; 9-VI 1.11-1.39
(mean = 1.27, n = 5) length of segment
VI; 9-VII 1.05-1.18 (mean = 1.13, " = 5)
length of segment VII: 9-VIII 0.85-0.92
(mean = 0.90, " = 5) length of segment
VIU. Seta 9-1 positioned near anterolateral
edge of tergum: 9-11 on the posterolateral
edge of tergum; 9-III-VIII near posterolateral edge of tergum. Segment VII 1.05-1. 19
(mean = 1.12, " = 5) length of segment
VI: segment VIII 1.11-1.39 (mean = 1.26,
n = 5) length of segment VI; segment VIII
1.05-1.18 (mean = 1.13, 11 = 5) length of
613
segment VII. Segment VII 0.85-0.92 (mean
= 0.90, n = 5) width of segment VI (width
at posterior margins); segment VIII 0.690.92 (mean = 0.84. " = 5) width of segment VI; segment vrn 0.75-1.01 (mean =
0.93, II = 5) width of segment VII. Width!
length of segment VI 2.29-2.88 (mean =
2.48, n = 5), VII 1.84-2.21 (mean = 1.99,
" = 5), VIII 1.23-1.85 (mean = 1.65, n =
5). Paddle length 0.70-0.75 mm (mean =
0.72 mm, 11 = 5). width 0.15-0.34 mm
(mean = 0.22 mm, n = 5), length/width
ratio 2.24-4.86 (mean = 3.73, n = 5): refractile index 0.34-0.67 (mean = 0.54, 11 =
5); paddle seta I-Pa simple or forked (2-4
apical branches), length 0.14-0.18 mm
(mean = 0.16, II = 9); 2-Pa simple or
forked (2 apical branches), length 0.400.14 mm (mean = 0.11); length of I-Pa
1.11-3.80 (mean = 1.66, n = 9) length of
2-Pa. Widthllength of genital lobe of female
1.33-1.43 (mean = 1.37, n = 3), male
0.87-0.89 (mean = 0.88, " = 2): numerous
spicules present on subapical and apical
margins of genital lobe of female, absent in
male.
Larva (Fig. 3).-Position and development of setae as figured; range and modal
number of branches and number of branches of neotype female as shown in Table 3.
Head: Length 0.68-0.71 mm (mean =
0.70, " = 4), width 0.66-0.76 mm (mean
0.71, " = 4). Antennal length 0.25-0.29
mm (mean = 0.26,11 = 8), slightly tapered
toward apex, 4.57-6.00 (mean = 5.27,11 =
8) longer than wide; with spicules longer
and more numerous ventrally and in vicinity of seta I A; spicule length 0.01-0.02 mm
(mean = 0.02, If = 12). Seta I-A with 913 (mode = 10, n = 8) branches, length
0.16-0.24 mm (mean = 0.19, If = 8), inserted 0.11-0.22 mm (mean = 0.14, If = 8)
from base of antenna; 2-A single. pointed.
length 0.10-0.20 mm (mean = 0.13, " =
8); 3-A single, pointed. length 0.05-0.17
mm (mean = 0.08, 11 = 7); 4-A with 6-8
branches (mode = 8, 11 = 7); 5-A short,
spine-like, 0.06-0.17 (mean = 0.13, tl = 8)
length of seta I-A; 6-A spine-like about
614
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
1---0.5 mm -----I
t.
VII
1
1.0mm
1
Fig. 3. Allopilele.r lesteri, larva. (A) Head. left side dorsal. right side ventral. (B) Dorsomentum (Om), (C)
Thorax and abdominal segments I-VI. left side dorsal. right side ventral. (D) Pecten plate (PP) and pecten spines.
(E) Abdominal segments vnr-x, side view.
616
PROCEEDINGS OF TI-iE ENTOMOLOGICAL SOCIETY
1.6x longer than seta 5-A. Seta 2-C single
1.69-2.37 (mean = 1.92, n = 3) length of
3-C; seta 2-C close to mate of opposite side
0.002-0.006 nun (mean = 0.004, n = 4);
3-C densely dendritic with 32-68 main
branches (mode = 42),0.10-0.14 (mean =
0.12. n = 8) length of 2-C, clypeal index
(distance between bases 2-C and 3-C on I
side/distance between bases of 2-C) 11.3340.00 (mean = 27.83. " = 4). Thorax: Seta
I-P with 1-3 branches (mode = I. n = 8);
9-IO,12-P single; 9- I 2-P setal support
plate spine length 0.04 mm. Setae 9-12-M
single; 9-M 3.12-12.67 (mean = 9.48, 11 =
3) length IO-M; 9-12-M setal support plate
spine length 0.01-0.02 mm (mean = 0.02,
n = 4). Setae 9-IO-T single; 9-T I.l8-1.31
(mean = 1.24, n = 4) length of IO-T; seta
12-T with 1-3 branches; 9-12-T setal support plate spine length 0.02 mm (n = 2);
13-T with 3 branches. Abdomen. Seta I-I
with 9-14 branches (mode = 12,,, = 8);
I-II 10-15 branches (mode = 13,1l = 8).
Seta l-m-VlI palmate with well-developed
leaflets, each leaflet with short filament; 0U-VIII and 14-Ul-VIII weakly developed;
0,8,14-1, 14-11 absent or rare; 3-1l-IIl.V-VI
single; 3-1, rv,VIT branched. Seta I-X single, 1.30-2.40 (mean = 1.81." = 5) length
of saddle; I-X inserted on saddle. Saddle
with minute, sparse spicules on lateral surface. Integument of posterior margin of segment X with strongly developed dark
brown to black spicules. Spiracular apparatus. Pecten plate with 12-18 spines; arrangement of spines alternating long and
short. with 7 or 8 (mode = 7. Il = 6) long
spines and 5-11 (mode = 9, n = 6) short
spines; long spines 1.27-11.67 (mean =
3.86, II = 37) length of short spines. Two
posterolateral spiracular lobe plates present.
each plate with elongate, slender, sclerotized projection from inner caudal margin.
Type material.-Neotype female with associated slide-mounted larval and pupal exuviae and DNA of a midleg of female;
reared from a larva cotlected from a ditch
on hill with slow flowing clear water.
28.8°C, pH 6.79. salinity 0.07 ppt. conduc-
or WASHINGTON
tivity 0.15 mS, data as follows: "Tanque",
Calauan. Laguna. Luzon. Philippines, L. M.
Rueda Coli. 29 July 2002, 14°08'44n N,
121°18'54n E. collection and specimen no.
PH 9-7. Deposited in the National Museum
of Natural History, Smithsonian Institution,
Washington, DC. (WRBU ACC No. 1729).
Alloneotype male with slide-mounted genitalia. extracted DNA of combined head.
thorax and abdomen, and associated slidemounted pupal exuviae with collection no.
PH 9-6 and same collection data as neotype female. The morphological descriptions of the head, thorax and abdomen of
the alloneotype male were recorded before
being processed for DNA. We were unable
to cotlect any specimens from the original
type locality of Santa Mesa, Manila, Luzon.
because it is now a highly urban area totally
lacking typical larval habitats. Baisas and
Hu (1936) noted that many cotypes of An.
le.fiferi were cotlected from Calauan, Laguna, Luzon. about 50 km south of Santa
Mesa. This locality remains rural and we
were able to collect specimens from Calauan for the present study. Morphological
data in Tables 1-3 are based on measurements of the neotype, alloneotype and associated specimens collected from Calauan
in 2002.
Other material examined.-247 specimens in the National Museum of Natural
History. Smithsonian Institution, Washington DC, consisting of 54 2,29 d, 76 pupal
exuviae, 85 larval exuviae, and 2 d genitalia. PHILIPPINES. LUZON: Province of
LAGUNA. Calauan, same collection data
as female neotype, PH 9-3, I F PeLe; PH
9-8, I <2 PeLe; PH 9-11, I d G PeLe; 17
April 1930. Lot 77-19, I 0; 20 Apr. 1930.
Lot 77-19, I 0; 19 Jan. 1931, Lot 122. 2
d; 9 July 1931, Lot 247, 3 9; 17 Sept.
1931, 9 9. 3 d; ; I Sept. 1932, W. V. King
coli.. Lot K317i. 1 d. Province of MINDORO ORIENTAL, OrdoviIla, 0.5 km W.
Victoria. seepage spring, B. Harrison and
Kol coil .. 17 July 1969. P58-127, I Pe; B.
Harrison coil.• 19 July 1969. P61-37, I
LePe; 1969, B. Harrison and Kol coli.,
617
VOLUME 107. NUMBER 3
1
61
121
181
241
301
361
421
aattagaagt
tcgtgcataa
aaggccgcga
gttctaggca
ttttatttta
caagagactg
tccgtgcaga
aggccactcg
ttggcaaaca
aggtgtaaga
agacagacaa
ggcgcgccct
aaaattgagg
gaatcggaag
acgactggaa
gtggtcaa
gaaaactacg
gagatctcgt
gtagtaaaca
gacgctgtgt
taagacttcc
ttgaacaacg
gatgcaagtt
cagtgattgg
cgatcgcttg
gcagcagatg
gtagatggag
aacgtttctt
gaacactcta
ctacctgaga
tgctggtcac
catctcggaa
tgttcccgcg
caggtgtctt
cgagatagtg
ttagcaaaca
atcattatca
cacgtcacgg
cttgtgttga
attggcggaa
cctcatctat
gaatgggctg
ctacccagaa
cttacgagtg
Fig. 4. Imernal transcribed spacer 2 (ITS2) sequence of the ribosomal DNA for A,l. lester; from the type
locality. Calauun. Laguna. Philippines. GenBank accession number AY375469.
SEAMP Acc. 233. P44-1. I <}; P58-36. I
0; P58-37. I セ[ P58-11I, I 0; P58-116L,
I 0; P58-127, 1 0; Caminawit Pt.. 30 Dec.
1944, G. H. Pena coil., No. C-41, I M.
MOUNTAIN PROVINCE. Baguio. 121une
1945, 32MSU#140, I 2; S. E. Sheilds coil .•
lO Aug. 1945, 24MSU#432. 12 Nセ 4 0;
Trinidad, May 1945, 1. G. Franclemont,4
セN
I O. Province of NUEVA ECIJA. Munoz, Rozeboom. Knight and Laffoon coli.•
8 Aug. 1945, #1153, 3 LePe. Province of
PAMPANGA, Olongapo (Santa Rita). Rozeboom, Knight and Laffoon coil .• 10 Aug.
1945, #1115, 4 PeLe; #1116, I PeLe. Province of RIZAL. Camp Nichols. PVT Carraway coli., 14 Dec. 1924. I 2; 18 Dec.
1945, P469, salty fishpond with algae. I 0;
Mandaluyong, 17 Dec. 1945. P468. rice
field. I セN VISAYAS: Province of LEYTE,
Tacloban. Rozeboom. Knighl and Laffoon
coli., 16 Aug. 1945, No. 1713. I LePe. I
2, I 0; Southern Leyte, 2 Dec. 1944. O.
H. Graham. 4 2, I 0; Palo Alto, 1945, I
2; 8 lan. 1945, 32MSU#384a, I 2; I Feb.
1945, 32MSU#P430, I 0; 11 Mar. 1945.
32MSU#474. I 2. Province of NEGROS
OCCIDENTAL. La Carlola, 31 lan. 1930.
W. V. King coli., Lot 124-16, 1 0.; Silay,
3 Feb. 1930. Lot 137. I O. Province of SAMAR. Osmena, Rozeboom, Knight and
Laffoon coli.. 8 Sept. 1945. #1334.30, progeny brood, 9 Le, 29 PeLe; # 1334.34. 2 Le,
12 PeLe; #1348.3, 13 PeLe; #1348.6. 4
PeLe; 1945. Rozeboon, Knight, Laffoon
coil., No. 1348.5. 3 M; San Antonio. 29
Nov. 1944, J. H. Paullos coil.• No. 506. I
2; I Dec. 1944.1. H. Paullos coil., No. 507.
1 2; Dec. 1944,2 9,3 O.
Molecular characterization.-DNA was
extracted from a midleg of the neotype female (PH9-7), the whole body (excluding
genitalia) of the alloneotype male (PH9-6),
a midleg of a female (PH9-3), and the
whole body (excluding genitalia) of a male
(PH9-ll). Ribosomal DNA ITS2 sequences are the same for all 4 (GenBank accession number AY375469 (PH9-6; Fig. 4).
Other sequences in GenBank that match
these sequences are as follow: under the
name An. amhropophagus, Acc. Nos.
AF384172, Al004941. AF543860; An. lester; Korea. Acc. No. AY375468); An. lester; China (locally identified as An. anthropophaglls), Acc. No. AY375467.
Distribution.-China (Hong Kong, south
and central areas of the mainland. extending
west to 105°E longitude and north to 43°N),
Guam, Japan (including Ryukyu Islands).
Korea, Philippines (Luzon: Laguna, Manila,
Mindoro Oriental, Mountain Province,
Nueva Ecija. Pampanga, Rizal; Visayas:
Leyte, Negros Occidental. Samar).
Medical importance.-Anopheles lester;
is a human biter and is considered a principal vector of malaria in southern China
(Beales 1984, Chow 1991, Ho et al. 1962,
Ma 1981) and other areas of the country (as
anthrophophaglls, Tang et al. 1991). It is
suspected of being a primary vector in lapan and Korea (Kamimura 1968, Otsuru
1949, Tanaka et al. 1979). Natural infection
rates of An. lesteri in the 1960s were 1.9 to
14.4 times greater than An. sinensis in China (Gu et al. 1966). In the Philippines and
Guam. its biting habits are unknown, and it
is not known to transmit malarial parasites.
618
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Il (as amhropophagus) has a strong prefer- tions of the species in many parts of its geoence for human blood, and plays an impor- graphical range in Asia. particularly Korea,
tant role in the transmission of filariasis and Japan and China. The morphological informalaria in central and south China (Xu and mation in this paper, coupled with rONA
Feng 1975). Harrison (1973) suggested a ITS2 sequence (Wilkerson et al. 2003), will
need to determine the distribution, behavior, help in solving those problems.
Atlophleles lesteri has the following diand malarial and filarial parasite susceptibilities of All. lesteri throughout its range. agnostic features. Adult female. Maxillary
This species, instead of All. sillellsis. may pal pus with palpomere 3 having narrow
be the more significant vector in Taiwan, basal pale band about as wide as pale bands
Okinawa. Japan, Korea, and central and of other palpomeres: vein Cu2 without apical pale fringe spot (unlike sinensis. sillenorthern China.
Bionomics.-The larvae of All. lesteri roides. pullus); subcostal pale (SCP) spot
are found in a variety of habitats including narrow; humeral crossveins without scales
freshwater pools, margins of ponds, lakes, (unlike pseudosillellsis); midcoxa without
areas preferably not affected by salt tides upper patch of pale scules (unlike s;lIensis);
(Baisas 1974). and ditches with slowly hindtarsomeres 2-4 with narrow apical pale
flowing clear water in hilly areas in the bands. hindtarsomere 4 without basal pale
Philippines. In Japan. the larvae occur in band (unlike pediwelliatus). Adult male.
marshes. sluggish streams, ground pools. Male genitalia with dorsal lobe of cIaspette
ponds. rice fields, and other impounded wa- having 3 closely appressed setae of about
ters (Tanaka et al. 1979). Unlike All. sitlell- equal length. Aedeagus with 4 leaflets on
sis, All. leste"; prefers places that are cool each side; 2 most mesal leaflets with broadand shaded. Adult populations of Atl. lesteri er transparent inner margins than other leafreach their peaks during the summer season lets. Tergum IX bare, with pair of caudally
in Hokkaido (Kamimura 1976). and during directed elongate capitate lobes. Pupa.
June and October in Honshu and Kyushu. Trumpet with thick and serrate rim. Wing
Japan (Otsuru and Ohmori 1960). The spe- with checkered dark stripes. Setae 9-III-VII
cies is more frequemly found in coastal single, with nan'owly rounded apex; seta 5areus than inland. In Hong Kong, it com- V with 13-24 branches. Larva. Setae 2-C,
monly occurs in hilly areas and grassy 3-P, 3, 5-M single; 3-C with 32-68 branchfields (Chau 1982). In Guam. All. lesteri es; 4-M with 3-5 erect branches; 9-M more
larvae were found in a carabao wallow than 3 times the length of IO-M; 9-M about
(Basio and Reisen 1971). Adults of An. les- 1.5 times longer than lO-T; 5-111 with 5-8
teri. An. sinellsis and other anophelines branches; 9-111 with 6-11 branches; 13-IV
were collected in cow sheds and living with 3-5 branches; I-X strong. single about
rooms of houses in villages during malaria 2 times or more length of saddle; pecten
surveys in Korea (Whang 1962). Anopheles with 7 or 8 long spines and 5-11 short
lesteri has been confused with Atl. sinensis spines. Reid (1953) and Harrison (1973)
and other members of the Hyrcanus Group, provided useful diagnostic pupal and adult
l\"d some published records of its distribu- morphological characters to separate An.
tion and bionomics are not accurate. partic- lester; from All. sinellsis. Harrison and
Scanlon (1975) also listed several characularly in Japan, Korea and China.
ters of all life stages of lO species of the
DISCUSSION
Hyrcanus Group found in Thailand. They
Although Baisas and Hu (1936) provided also discussed extensively the morphologithe original description of Atl. lesteri. it was cal taxonomy of the Lesteri Subgroup.
not sufficient for accurate identification of Comparisons of pupal and larval characters
the species. This resulted in misidentifica- of All. lester; and related species of the Hyr-
619
VOLUME 107. NUMBER 3
canus Group from China and other areas of
Asia are needed to further clarify species
differences. Other morphological features
described in this paper for larvae. pupae
and adults may be helpful for separating
An. lesteri from related species.
Morphological similarities in all stages.
along with intraspecific variation of many
species in the Hyrcanus Group. have lead
to much confusion in Korea. Japan and China (Tanaka et al. 1979). and possibly in other areas of Asia where they occur. Wilkerson et al. (2003) suggested that the best way
to infer conspecificity of populations across
large geographic areas is to compare specimens from type localities. Based on the
combination of published and their newly
generated rONA ITS2 sequences. Wilkerson et al. (2003) found that All. lester; from
South Korea and An. anrhropophagus from
Jiangsu Province. China. are the same as
An. lester; from near its type locality in the
Philippines (Calauan. Laguna. Luzon).
Anopheles antlrropohagus, considered a
major malaria vector in central and north
China. is actually An. lester;, not a separate
species. With that finding. they placed An.
anrhropophagus in synonymy with its senior synonym, An. lesteri. Any morphological features previously thought to differentiate An. anthropophagus and All. lester;
are evidently variable characters of a single
species. What is called An. lester; in China
(as reported by Gao et al. 2004: 7. 9) is
actually an unknown species when compared with the work of Wilkerson et al.
(2003). Several molecular studies (e.g.. Li
et al. 1991; Ma et al. 2000a. b; Gao et al.
2004) were conducted but were unable to
clarify the taxonomic identity of An. lesteri
found in China and Korea. With the collection of An. lester; specimens from near the
type locality in Luzon, Philippines. Wilkerson et al. (2003) were able to compare the
specimens with those from Korea and China, and they concluded that the so called
An. anrhropophagus from China. and the so
called An. lester; from Korea are conspecific with An. lester; from the Philippines.
With the designation of the neotype and
detailed descriptions of various life stages
of An. lester; based on specimens collected
near the type locality. future systematic
studies may be conducted using various
methods. including morphological. molecular or biochemical. With the identity of All.
lester; resolved. the effectiveness of malaria
vector control practices could be further improved. As suggested by Harrison (1973).
additional information is needed on the distribution. behavior. and malarial and filarial
parasite susceptibilities of vector species
throughout their ranges. A vector species
may be a more significant parasite vector in
one geographical area than in others. For
example. An. lester; is considered an important malaria vector in China and Korea.
but not in the Philippines. Furthermore. because several species of the Hyrcanus
Group are involved in the transmission of
malaria and filariasis in Asia, there is a need
to revise the taxonomy of the whole group
and to further clarify the identities of the
cryptic species. particularly the vectors, in
the group.
ACKNOWLEDGMENTS
We thank Bruce A. Harrison. Yiau-Min
Huang and Maria Anice M. Sallum for reviewing the manuscript and providing invaluable advice. Special thanks to the following: Taina Litwak for her illustrations;
Bel Rueda, Buddy Buenavista. Lerma Buenavista. Benjie Puma. and Modesta Coronado-Puma for their help in collecting mosquitoes. Jim Pecor provided technical support in curation of specimens and preparation of slide mounts. This research was
performed under a Memorandum of Understanding between the Walter Reed Army Institute of Research and the Smithsonian Institution. with institutional support provided
by both organizations. The opinions and assertions contained herein are those of the
authors and are not to be construed as official or reflecting the views of the Department of the Army or the Department of Defense.
620
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