一目■王

一第23巻第4号平成7年12月
内

容

著

原

コレラ菌の菌体成分に対するコレラ患者の免疫反応（英文）
・中島

一敏，岩永

正明……………・…・・…………

227−231Cηρ螂ρ07観襯窺癬sスポロゾイトに対する家兎抗血清の中和作用に関する研究（英文）
・・宇仁

茂彦，林

Nazem
木俣

真二，福永

昭廣，高橋

研一

M，AbdElmaksoud

勲，井関基弘一……………一・…・………

233−238・

パプアニューギニア，ブーゲンビル島のブユ（英文）・・高岡

宏行……………7…・・…………一……・……一・ 239−252

ソロモン群島のブユ（英文）

一高岡
記

宏行，鈴木

博…………一……・…………

253−272

投稿規定・………………・・………一………・………………・……………・……・…………………

−274

日本熱帯医学会雑誌−275

録
23巻平成7年（1995年）総内容・………………・・…・・…一………・・…

日本熱帯医学会雑誌編集委員名簿……………・・…・……………・・…・…………・……・・………

・・281

圏■

227

Jpn. J. Trop. Med. Hyg., Vol. 23, No. 4 (1995) pp. 227‑231

Immune Response of Cholera Patients
against the componentsof Vibrio cholerae
KAZUTOSHI NAKASHIMAAND MASAAKI IWANAGA ,'Received August 2, 1995/Accepted October 4, 1995

Abstract: There was a sporadic outbreak of cholera in 1994 in Okinawa. Immune response of the cholera
patients against Vibrio cholerae cell components and extracellular proteins were examined. A potent
response to cholera toxin and hemolysin, and generally poor response to purified cellular components wereseen. Although the antigen was not defined, marked elevation of vibriocidal antibody and cell agglutination

antibody was observed. Only little elevation of the agglutination titer against heat‑killed organisms
suggests that the antigen (agglutinin) is heat labile protein on the cell surface.
ldentification and purification of this protein antigen will elucidate a protective antigen of V. cholerae.

INTRODUCTION
Cholera patients acquire potent protective immu‑
nity after the infection (Benenson et al., 1968) , however,sera from 6 healthy individuals without the history ofcholera and traveling abroad were used as the control.
Some informations about the sera were shown in Table
1.

the ideal vaccine for cholera is not developed yet, andAntigens: Antigens to examine the immune

even the protective antigen of cholera vibrio is not
sufficiently elucidated. Protective antigen should be
immunoresponsive, therefore, immune response to Vi‑brio cholerae O1 has been intensively studied in human

response included V. cholerae whole cells, cytosol pro‑teins (complex and purified 40 kDa protein) , innermem‑

brane proteins, outermembrane proteins (complex andpurified OmpS), flagellar core protein, 5 kinds of V.

volunteers, but a few studies in naturally infected chol‑cholerae pili (VCF, VIO, TCP, Mini, V14) , cholera toxin

era patients (Pierce and Sack, 1977; Richardson et al.,(CT) and hemolysin. VIO pili, V14 pili and hemolysin
were obtained from V. cholerae non‑O1, Mini‑pili from1989). In those papers, the antibodies against LPS,
cholera toxin (CT) , whole bacterial cell, and someoutermembrane proteins were studied in relation tovibriocidal activities.

There was a sporadic outbreak of cholera in Okin‑awa in July 1994, and 6 bacteriologically confirmed
patients were officially reported. The organisms iso‑lated from the patients were all V. cholerae O1 El Tor,

V. cholerae 0139. The other antigens were obtained
from V. cholerae O1.
Preparation of the antigens: The organisms cul‑
tured in heart infusion broth at 37 C for over night were

harvested. The heavy suspension of the organisms innormal saline solution was treated with biomixer(Nihon Seiki CO., Tokyo, Japan) to detach pili andserovar Ogawa. This paper described their immune

flagella. The cells without pili and flagella were frozen

response to a variety of cellular components and extra

at ‑80 C for a few days, once made them thaw, refrozen
at ‑20C, and then crushed the cells with X‑press. The
crushed materials were centrifuged at 85,000 X g for 1
hour, and the supernatant was regarded as cytosol. The
pellet was washed 2 times with saline and collected withcentrifugation. The final pellet was suspended in 10 mM‑cellular proteins of V. cholerae.

MATERIALS AND METHODSPatients and sera: The convalescent sera frorn 4
patients with severe diarrhea and a patient with mild
self‑limited diarrhea were obtained to determine theimmune response against the pathogens. In addition, thePBS (pH 7.4) supplemented with 1% sodium lauroyl
sarcosinate, and homogenized in a glass‑hornogenizer
under mild sonication at times. The homogenized sam‑

1 Department of Bacteriology, 2 Research Center of Comprehensive Medicine, Faculty of Medicine, University of the Ryukyus 207
Uehara, Nishihara, Okinawa 903‑01, Japan

228

Table 1

Serum Age
P1
P2
P3
P4
P5

H1
H2
H3
H4
H5
H6

52

80
44

39
51

Informations about the sera

Sex samplmg

diarrheaother symptoms

Mmuscle cramp, acute renal f ailure

9

SevereSevereSevereSevere11

mild (self ‑ Iimited)

F
F

M
F

17
9

10

21

25
24
21

26
28

muscle cramp
acute renal failuremuscle cramp

healthyM
M
M
M
M

he a I thy

healthyhealthyhealthyhealthyDays after onset, P: Patients, H: Controls (healthy subjects)

ple was centrifuged at 85,000 X g for I hour, and thesupernatant was regarded as innermembrane. The
pellet was repeatedly homogenized and sonicated in 1%
SLS, and the final pellet was regarded as outermem‑
brane. Flagellar core was purified from a V. cholerae
O1 El Tor by detaching with biomixer, sedimentation inpolyethylene glycol, and DEAE column chromatogra‑
phy. The other antigens were purified in accordance

zthiazoline sulfonic acid) was used as the substrate.
Antibody titers were expressed as reciprocal of the
maximum dilution of the patient's sera giving the OD415
more than 0.3.

RESULTS

with the references as VCF pili (Ehara et al., 1987;

Agglutination titers: Agglutination titers of thepatient's sera against live organisms were markedly

lwanaga et al., 1989), VIO pili (Yamashiro et al., 1993),

elevated when live organisms were used as the antigen.

TCP (Osek et al., 1994), Mini‑pili (Yamashiro et al.,However, serum of the patient with mild diarrhea

1994) , CT (Nakashima et al., 1995; Spyrides and Feeley,

revealed low elevation of the titer as 256. The highest

1970), hemolysin (Iwanaga and lchinose, 1991) , etc.

titer in the control sera was 32. The agglutination

Detection of antibodies: Presence of antibodies
against various antigens were examined by western

against heat‑killed organisms was very poor (Table 2) .

blotting as described by Towbin et al. (8) using 1:200
dilution of the patient's and control sera as the firstantibodies. Horse radish peroxydase (HRP) conjugatedanti*human lgG goat serum (CAPPEL) diluted to 1:1000
was used as the second antibody.
Titration of the antibodies: Vibriocidal antibody
was titrated by using microplate technique as describedpatient's sera showed the titer more than 20,480 except
the serum of the patient with mild diarrhea. The control

by Bennenson (1968). Agglutination titers were
examined by serial 2 fold test tube dilution method using

live and heat‑killed organisms. Vibriocidal‑ and agglu‑
tination activities of the sera were examined for theorganisms isolated from the patient corresponding toeach serum. Antibody titers were expressed as the recip‑
rocal of the maximum dilution in the positive findings.
Anti‑CT and anti‑hemolysin antibodies were titrated by
ELISA using serially 2 fold dilution of the patient's sera

and the antigen coated plastic plates. HRP conjugatedanti‑human lgG goat serum diluted to 1:1000 was usedas the second antibody, and 2,2'‑azino‑bis (3‑ethylben‑

Vibriocidal titer: Vibriocidal activities of theTable 2Serum

Titration of the antibodies

Cell Agglutination

Vibriocidal

Live

Heat‑killed

P1
P2
P3
P4
P5

> 8,1928> 20,480

4,096

8> 20,480

1,024

8> 8,1928>20,480>20,480256<410,240

H1
H2
H3
H4
H5
H6

8

<4
<4
<4
<4
<4
<4

<4
<4
32
8

<4

80
< 10

40
2,560

320
640

229

Immunoreactivities as examined by Western blot (No. of positive/examined)

Table 3

MW
115
85
77
71

56

OMPS

IMPS

CPs

HC Pts P value MW HC

Pts

3/6
1/6
0/6
2/6
3/6
1/6
0/6
0/6
1/6
1/6

415
3/5
1/5
3/5
2/5
415
2/5
1/5
2/5
2/5

416
0/6
1/6
0/6
1/6

4/5
1/5
1/5
1/5
1/5

0.575

115

0.455

85

0.727

77

0.455

67

0.727

56
45

43 0/6 1/5

28 0/6 1/5
14 2/6 l/5

0.455

43
33

0.455

23

0.576

14

P‑value
0.348

MW HC

Pts

P‑value

45 4/643 41633 1/6

2/5
3/5
3/5

0.392

0.197
0.455
0.392
0.608

0.067
0.182

0.455
0.424

0.652
0.197

0.424

CPs: Cytosol proteins, IMPs: Innermembrane proteins, OMPs: Outermembrane proteins,
molecular weight of reacted protein, HC: sera of healthy controls,

MW
Pts: patients' sera, P‑value: calculated by applying Fisher's exact probability

Table 4

Immunoreaction frequency of the sera against
purified antigens as examined by Western blot
(No. of positive/examined)

Antigens

HC

Pts

P‑value

Flagellin

0/6
0/6
1/6

0/5
1/5
1/5

1.000

40k p

OmpS

0.455
0.727

Cholera toxin

A subunit
B subunit
Hernolysin

0/6
1/6
0/6

2/5
5/5
4/5

0.182

0/6
0/6
0/6
0/6
0/6
0/6

0/5
0/5
0/5
0/5
0/5
2/5

1.000

0.013
0.015

Pili

TCP
VCF/classic

VCF/EI Tor
Mini
V 10

V14
HC:

1.000
1.000

Table 5ELISA titer of sera to extra‑
cellular proteins

Serum

CT

Hemolysin

P1
P2
P3
P4
P5

400

1,600

H1
H2
H3
H4
H5
H6

200

400

800

< 50

800

200

400

800

100

50

< 50

< 50

< 50

< 50

50

50

< 50

< 50

< 50

< 50

1.000
1.000

0.182

Antibbdies against extracellular proteins: Contrary

Control sera, Pts: Patient's sera, 40kp: 40kDa

to the cellular components, the patient's sera potently
reacted to extra‑cellular proteins such as cholera toxin

cytosol protein
P‑value: calculated by applying Fisher's exact prob‑
abi lityand hemolysin, whereas the control sera did not react(Table 4, 5) .

serum with its agglutination titer of 32 revealed vi‑
briocidal titer of 2,560 (Table 2).

Antibodies against cellular components: Some of
the cytosol proteins, innermembrane proteins, and outer‑membrane proteins reacted to the patient's sera and to

DISCUSSION
A Ithough there have been many studies on immune

the control sera with the simillar extent (Table 3).

response to V. cholerae in cholera infection, the atten‑
tion was directed to the response against some limited

Reactions of the sera to the purified components such as

antigens (Hall et al., 1991; Majumdar et al., 1981; Richar‑pili, flagella, OmpS (maltose induced 43kDa outenTlem‑
brane protein) , 40kDa cytosol protein, etc. were almost
negligible, except that 2 sera showed a potent reaction
to V14 pili (Table 4).

dson et al., 1989). The protective antibody for cholera
has not been identified so far. In the present study, theresponse to a variety of antigens such as extracellular

proteins, cytosol proteins, innermembrane proteins,

230

outermembrane proteins, flagellin, and pilins was

willingly cooperated us for collecting samples and infor‑

examined. The organism of V. cholerae has uncountablemation.

number of antigens. However, the present study
revealed that the host immune response was not seen
against all the antigens but a limited number of theantigens. A protective antigen of a certain pathogen is
supposed to induce a potent immune response and to bea pathogenic factor. Cholera toxin is a main pathogenic

factor of V. cholerae and induce potent immune
response. Animals immunized with CT are protectedfrom the illness to some extent, however in humanbeings, vaccination with CT does not protect the vac‑cinees from cholera (Noriki, 1976; Pierce and Sack,
1977). Hemolysin is also a potent immunogen but not
likely a protective antigen. The elevation of the anti‑
bodies against a variety of the purified cellular compo‑

nents including a group of proteins in cytosol andmembranes was not cornmonly seen. Besides, the anti‑
bodies against those proteins were also detected fromsome of the control sera with the simillar frequency. It

suggests that there must be some cross antigens in theintestinal floral organisms. Anti‑pili antibodies were
not detected at all except anti‑V14 pili. TCP has beenrecognized to be involved in colonization and not toelevate the antibody in cholera patients (Hall et al.,1991) . The other pili excluding V14 pili were proved not

to be colonization factors (Iwanaga et al., 1989; Na‑kasone and lwanaga, 1990), therefore, it is no wonderthat anti‑pili antibodies were not elevated in these

REFERENCES
1 ) Benenson, A.S., Saad, A., Mosley, W.H. (1968): Ser‑ological studies in cholera. 2. The vibriocidal antibody
response of cholera patients determined by a microtech‑nique. Bull World Health Organ 38, 277‑2852 ) Ehara, M., Ishibashi, M., Ichinose, Y., Iwanaga, M.,
Shirnotori, S., Naito, T. (1987): Purification and partial

characterization of fimbriae of Vibrio cholerae O1.
Vaccine 5, 283‑288
3 ) Hall,R.H., Losonsky, G., Silveira, A.P.D., Taylor, R.K.,

Mekalanos,J.J., Witham, N.D., and Levine, M.M. (1991)
Immunogenicity of Vibrio cholerae O1 toxin‑coregulat‑
ed pili in experimental and clinical cholera. Infect.
Immun. 59, 2508‑2512

4 ) Iwanaga, M., Ichinose, Y. (199D: An Aberrant
Hemolysin of Vibrio cholerae non‑O1. Microbiol. Im‑
munol. 35, 705‑715
5 ) Iwanaga, M., Nakasone, N., Ehara, M. (1989) : Pili of
Vibrio cholerae O1 biotype E1 Tor: a comparative study
on adhesive and non‑adhesive strains. Microbiol. Im‑
munol. 33, 1‑96 ) Majumdar, A.S.. Dutta, P., Dutta, D., and Ghose, A.C.
(1981): Antibacterial and antitoxin responses in theserum and milk of cholera patients. Infect. Immun. 32, 1‑87 ) Nakashima, K., Eguchi, Y., Nakasone, N. (1995) : Char‑

acterization of an enterotoxin produced by ' Vibrio
cholerae 0139. Microbiol. Immunol. 39, 87‑94

cholera patients. V14 pili have been newly identified in
the recent, and this pilus is likely a colonization factor8 ) Nakasone, N and lwanaga, M. (1990): Pili of Vibrio

according to the studies so far (unpublished data) . In
this viewpoint, the immunological reaction against V14
pili, although 2 cases out of 5, is noteworthy.

9 ) Noriki, H. (1976) : Evaluation of toxoid field trial in the

Outstanding immune response of cholera patients,

except the elevation of antibodies against CT andhemolysin, was seen in marked elevation of vibriocidal
antibody and agglutination antibody against live organ‑
isms. Vibriocidal activity of those sera was elevated inparallel with agglutinating activity against live organ‑
isms but not against heat‑killed organisms. It suggeststhat the agglutinin reacting to the patient's sera is heat

labile substance located at the surface of the organisms.

The antigen for vibriocidal antibody and the heat labile
agglutinin could be identical, therefore, the identifica‑tion of non‑LPS heat labile agglutinin may be a key toelucidate protective antigen of V. cholerae Ol.

ACKNOWLEDGMENT
We are grateful to Dr. T. Kishaba and those whocholerae non‑O1. Infect. Immun. 58, 1640‑1646
Philippines. In. Proc. 12th Joint Conference on cholera(US‑Japan medical Cooperation) . pp302‑310
10) Osek, J., Jonson. G., Svennerholm, A.‑M., Jan, H. (1994):

Role of antibodies against biotype‑specific Vibrio
cholerae pili in protection against experimental classicaland El Tor cholera. Infect. Immun. 62, 2901‑2907
11) Pierce, N.F., and Sack, R.B. (1977) : Immune response of
the intestinal mucosa to cholera toxoid. J. Infect. Dis.
136. S113‑S117
12) Richardson, K., Kaper, J.B., and Levine, M.M. (1989):

Human immune response to Vibrio cholerae O1 wholecells and isolated outermembrane proteins. Infect.
Immun. 57, 495‑501
13) Spyrides, G.J., Feeley. J.C. (1970) : Concentration andpurification of cholera exotoxin by absorption on alu‑
miniurn compound gels. J. Infect. Dis. 121, S96‑S99
14) Towbin, H.. Staehelin, T., Gordon, J. (1979.): Electro‑
phoretic transfer of proteins from polyacrylamide gelsto nitrocellulose sheets:Procedure and some applica‑
tions. Proc. Natl. Acad. Sci. U.S.A. 76, 4350‑4354

15) Yamashiro, T., Nakasone, N., Honma, Y., Albert, M.J.,

231

Iwanaga，M．（1994）：Purification and characterizationof四6n。o ohoJ召耀0139fimbriae．FEMS Microbio1．Lett．
115，247−25216）Yamashiro，T．，Nakasone，N．，Iwanaga，M．（1993）：
Purification

andcharacterization

of

pili

of

a

Viあ7zbohol召名鷹non−01strain．Infect．Immun．61，5398−5400

233

Jpn. J. Trop. Med Hyg Vol 23 No 4 1995, pp. 233‑238

NEUTRALIZATION OF CR YPTOSPORIDIUM
MURIS SPOROZOITES BY RABBITANTI‑C. MURIS SERUMSHIGEHIKO UNll, SHlNJI HAYASH11 AKIHIRO FUKUNAGA2, KENICHI TAKAHASH13

NAZEM M. ABDELMAKSOUD1, ISAO KIMATAI AND MOTOHIRO ISEKI
Received September 20, 1995/Accepted Nobember 5, 1995

Abstract: The neutralization of Cmptosporidium muris sporozoites by rabbit antiserum was examined invitro and in mice. The sporozoites of C. muris could be separated from intact oocysts by repeatedcentrifugation. The antiserum reacted strongly with three bands of sporozoite antigens at > 200, 78, and 47.5

kilodaltons during immunoblotting. Heat‑inactivated antiserum caused morphological changes seen ultra‑
structurally in the sporozoites. The antiserum reduced the viability of the sporozoites as measured withfluorescein diacetate. When antiserum‑treated sporozoites (1 >< 105) were used to inoculate 7‑day‑old mice
per os, endogenously developed parasites were first detected 10 days later by an indirect fluorescence
antibody test. When sporozoites treated with normal rabbit serum were used to inoculate mice, parasites
were first detected 6 days later. The results suggest that the antiserum partially neutralizes the infectivityof C, muris sporozoites.

INTRODUCTION

are limited to studies by Uni et al. (1987), Iseki et al.
(1989) , and McDonald et al. (1992) . The purpose of thisCmptosporidium is a genus of parasitic protozoa

study is to identify the effects of antiserum to C. muristhat infect respiratory and gastrointestinal epithelial
cells of a wide range of vertebrates, causing cryptospo‑
ridiosis in humans and domestic animals (Dubey et al..
1990). C. muris parasitizes the stomachs of cattle andon sporozoite viability and infectivity in an examination

rodents, causing gastric lesions and decreasing weightgains in cattle (Anderson, 1989).

Chemotherapy is not effective in this infectionof the efficacy of immunotherapy for C. muris infection.

MATERIALS AND METHODSThe RN 66 strain of C. muris was maintained bydium. Hyperimmune bovine colostrutn and anti‑para‑subinoculation into specific‑pathogen‑free (SPF) micein our laboratory. Purified oocysts were obtained fromthe feces of infected mice by sucrose flotation andstored in 2.5% K2Cr207 at 4'C for up to I month before

site antibodies are said to be at least partially effica‑use (Iseki et al.. 1989).

(Dubey et al.. 1990) . Mata (1986) suggested that breast‑

fed children are partly protected againSt Cmp,tospori‑cious in preventing and treating infection by C. parvum
in mice (Riggs and Perryman, 1987; Fayer et al.. 1989;
Perryman et al.. 1990; Fayer et al.. 1990; Tilley et al..
1991; Riggs et al.. 1994) or an immunodeficient child(Tzipori et al.. 1986). However, other workers havefound that antibodies and lacteal factors are not usefulfor the prevention or treatment of infection in mice
(Moon et al., 1988; Arrowood et al.. 1989) or humans(Saxon and Weinstein 1987; Sterling et al.. 1991).

Reports on experimental infections with C. muris

Sporozoite preparation: Purified oocysts (4xl07) weresuspended in minimum essential medium (MEM, Flow
Laboratories, Irvine, Scotland) and centrifuged at 600 ><g for 5 min at 4'C to remove light oocysts. This process

was repeated a total of three times. The sedimented
oocysts were suspended in MEM and incubated for 30
min at 37'C. This excystation preparation, in which 75%
of the oocysts were excysted, was centrifuged at 600 X g
for 5 min at 4'C. The supernatant (the upper 3/4 of thelO ml in the plastic tube) was taken out with care not to*Department of Medical Zoology,
2Laboratory of Biology,'Department of Anatomy,
Osaka City University Medical School, 1‑4‑54 Asahimachi, Abeno‑ku, Osaka 545, Japan234

shake the tube and contained both sporozoites (1 x 108)

(1:500), and treated with biotinylated anti‑rabbit lgG.

and oocysts (1xl07). MEM was added to the super‑

The strips were then treated with avidin‑biotin com‑natant and the mixture was centrifuged at 600 >< g. The
supernatant obtained was centrifuged one more time inplexes (ABO bound with horseradish peroxidase

the same way. The supernatant included sporozoites(4xl06) and intact oocysts (1xl03), which were col‑CA) . Antigens were made visible with 4 ‑ chloro ‑ I ‑naphthol. In the control experiment, the nitrocellulose

lected by further centrifugation at 1,500 X g for 15 min

Preparation of antiserum against oocysts and sporozoites:

strips were treated with normal rabbit serum in thesame way. Molecular mass standards for comparison
were rabbit skeletal muscle myosin (200 kDa), Esche‑richia coli p‑galactosidase (116 kDa) , rabbit muscle

Excysted oocysts (4xl07; about 0.5 mg/mD in phos‑

phosphorylase b (97.4 kDa) , bovine serum albumin (66.2

phate‑buffered saline (PBS) were homogenized with anultrasonic homogenizer (Nihonseiki Co., Ltd., Tokyo).
One healthy New Zealand White rabbit without coccidia
infection was immunized by a subcutaneous injection ofan emulsion of excysted oocysts with Freund's completekDa), hen egg white ovalbumin (42.6 kDa), bovinecarbonic anhydrase (31.0 kDa), soybean trypsin in‑hibitor (21.5 kDa), and hen egg white lysozyme (14.4
kDa) (Bio‑Rad Laboratories, Richmond, CA) .

adjuvant, and a booster injection with excysted oocysts

and used as the sporozoite preparation in this experi‑

ment.

(Vectastain; Vector Laboratories, Inc., Burlingame,

Sporozoite viability assay: A portion of the sporozoite(1xl08) in Freund's incomplete adjuvant was made 5

preparation (1x 105 sporozoites in I ml of MEM) was
mixed with 100 pl of heat‑inactivated (56'C, 30 min)weeks later. Blood was collected 8 days after the final
injection of the antigen and the serum was obtained.
The specificity of the antiserum for C. muris sporo‑antiserum, incubated at 37'C for 30 min, and washed in
PBS. A 100‑pl sample of this sporozoite preparation in
PBS was mixed with 900 pl of fluorescein diacetate (0.1

zoites was examined by an indirect fluorescence anti‑

mg/ml, Lambda Probes & Diagnostics. Graz, Austria)
and the mixture was incubated for 15 min at room
temperature. The sporozoites were examined under abody test (IFAT) . Air‑dried sporozoites fixed in
acetone were incubated with diluted rabbit antiserumfor 30 min, washed with PBS, and incubated for 30 min
with goat anti‑rabbit immunoglobulin G (IgG) Iabeledwith fluorescein isothiocyanate (Organon Teknika,
Malver, PA) .

Electrophoresis and western blotting: A sample of thefluorescence microscope. In the control experiment, aportion of the sporozoite preparation was incubatedwith MEM or heat‑inactivated normal serum and treat‑
ed with fluorescein diacetate. Sporozoites boiled for 5
min also were treated with fluorescein diacetate in the

sporozoite preparation (5xl07 sporozoites) wassuspended in 200 pl of buffer (150 mM NaC1, 5 mM

same way.

ethylenediaminetetraacetic acid, 50 mM Tris, and 0.02%
sodium azide, pH 7.4) containing 0.5% nonionic deter‑gent Nonidet P‑40 (Mead et al.. 1988). The preparation

the sporozoite preparation (1 x 105 sporozoites in I ml of

was freeze‑thawed twice and homogenized byparaformaldehyde‑3% glutaraldehyde in 0.1 M

ultrasonication. A 50‑pl sample was diluted with anequal volume of sample buffer CIOO mM Tris buffer, 4%
sodium dodecyl sulfate (SDS) , lO% 2‑mercaptoethanol,

0.2% bromophenol blue, and 15% glycerin] and the
mixture was boiled for 4 min. SDS‑polyacrylamide gelelectrophoresis (PAGE) of the sporozoite sample wasdone with 5% stacking gels and 10% separating gels at
a constant current of 20 mA in a slab‑electrophoresis
cell (Rapidas AE‑6450, Atto Corp., Tokyo).

Transmission electron microscopy ( TEM): A portion of

MEM) was mixed with 100 pl of heat‑inactivatedantiserum, incubated at 37'C for 30 min, and fixed in I %cacodylate buffer '(pH 7.4). For the control specimen,some of the sporozoite preparation was incubated withheat‑inactivated normal serum and fixed in the sarneway. The fixed specimens were dehydrated, embedded,
and sectioned as described previously (Uni et al., 1987) .

The specimens were examined with a JEOL 1200‑EX
electron microscope at 80 kV.
sporozoite infectivity assay: A portion of the sporozoite

After SDS‑PAGE, the antigens were transferred

preparation (1xl06 sporozoites in I ml of MEM) was
mixed with 100 pl of heat‑inactivated antiserum andfrom the gels to nitrocellulose membranes ("Clear blot

incubated at 37'C for 30 min. In the control experiment,

membrane‑P", Atto) with a device for semi‑dry west
ern blotting (Horize‑Blot AE‑6675P, Atto). Electro‑

some of the sporozoite preparation was incubated withphoretic transfer was done with a constant current of 1.5
mA/cm2 for I hr. The nitrocellulose strips were treatedthe incubation, a sample of the sporozoite preparation(1 x 105 sporozoites in 0.1 ml of the incubation medium

with goat serum, incubated with the rabbit antiserum

per mouse) was given orally to 7‑day‑old BALB/C

MEM or heat‑inactivated normal rabbit serum. After235

CrSlc mice (Japan SLC Inc.. Shizuoka) through a poly‑

A B

ethylene catheter (20 gauge, Terumo Corp., Tokyo).
fror evaluation of the infectivity of the contaminatingM' " ;' =" /"'/ '""'2 OO ‑ i' ;oocysts in the sporozoite preparation, a portion of the
sporoz;oite preparation was suspended in distilled water''and used to inocuiate mice; the step of incubation
mentioned above was omitted. ( Sporozoites are de‑stroyed by osmosis, but intact oocysts remain viable.)
In all experiments, one or two mice from each treatmentgroup were killed and examined daily for parasites ondays 6 to 10 and on day 13 postinoculation (PI) .

Impression smears of the gastric wall (the entire
posterior glandular part was used) were prepared from
inoculated mice, and IFAT was used to find the para‑sites. The posterior glandular part of the gastric wall is

the parastic location af C. muris.

RESULTS/.'xcyslation ard sporozoile praparatian: A high percent‑age (70‑90%) of excystation was achieved by the incu‑
bation of C. muris oocysts for 30 min at 37'C in MEM
Figure 1.

without treatment with sodium hypochlorite, sodium
taurocholate, or trypsin when the oocysts were usedwith rabbit antiC. muris serum (lane A) andnormal rabbit serum (1ane B). M, molecularwithin 2 weeks after isolation from experimentally in‑weight marker proteins (l03) .

fected mice. The oocysts were sticky, and usuallyadhered to the surface of the plastic tube. Aftercentrifugation (600 Xg) for 5 min, sporozoites and a few

diffuse mass of antigens had moved. Use of normal

intact oocysts were found in the medium, and almost alloocysts were found at the bottom of the plastic tube. By

rabbit serum resulted in no bands (lane B) .
The antiserum had a titer of l:8,000 for sporozoites

repeated centrifugation of the supernatants, theby IFAT. The serum reacted also with merozoites,

contaminating oocysts in the sporozoite preparation

meronts, macrogametes, and oocyst walls in impression
smears, but did not react with microgametes.

could be reduced to 0.03% of the number of sporozoites.
Characterization of spor;ozoite antigens recognized hy
rabbit anti.serum: Twelve bands (>200, 200, 78, 47.5, 40,

Viabilily of sporozoites trealed with antiserum: In vitro,

23% of sporozoites treated with heat‑inactivated33.5, 31.5, 30, 29, 27.5, 26.5, and 26 kDa) of antigens fromantiserum did not fluoresce (were not viable) after

C. muris sporozoites were detected with the antiserum

treatment with fluorescein diacetate. In contrast, l% of(Fig. l, Iane A) . There were strong bands at > 200, 78,

sporozoites treated with normai serum or medium alone
did not fluoresce (Table D . Boiled sporozoites did not

and 47.5 kDa. Three bands (27.5, 26.5, and 26 kDa) wereseen faintly in the low‑molecular‑weight range, where af luoresce.

Table I. Viability of C. muris sporozoites assayed with fluorescein diacetate

Treatmentl x lO' sporozoites (sp)Sporozoites

FluorescentNot f.1uorescent

Total

278

84

362

11 9

1

l 20

297

3

300

+ immune rabbit serum
diluted (1:1.0) in medium

Sp+nonTlal rabbit serum
diluted (1:10) in medium

S p + medium
P<0.01, ;i2 test.

236

=

'"

;::(:

.‑=

## ‑

'"**" .*:. +* +*

:*

={*

* ･ ';*;?

Figures 2‑3. Transmissien electron micragraphs of sporazoites treated with immune and normal sera.
Cytoplasmic vacuolization ( * ) and agglutination are seen in the sporozoites treated with theal"Itiserun,1 (Fig. 2) . x ll,OOO. Sporozoites treated with normal serum (Fig. 3) . x ll,OOO. AP, apical

camplex; RO, rhoptry; AM, amylopectin body; S, sporo oite.

Morph.r)!(,S.fical changes in sporOzc,)ites treat<ed with per smear was found in the four control mice, but there

antiserur/1: Morphak)gical changes such as swelling, were fewer than 10 parasites in the one mouse infected

cytoplasmic vacuolizatian, and agglutination of the when inoculated with sporozoites treated with the
sporozaites were seen by TEM in the sporozoites treat‑ antiser, um. The parasite was not detected on day 13 PI

ed with heat‑･inactivated antiserum (Fig. 2). Normal in the mice inoculated with the sporozoite preparation
serum caused virtually no morphological changes (Fig. treated with distilled water.
3) .

Jnfectivily of sporozoites trealed with antiserum' Para‑sites were first detected on day 10 PI in the mice when

' DISCU̲ SION

It was essential for detection of neutrali2;ation afheatinactivated antiserum was used (Table II). In

centrol mice, parasites̲ had already appeared on day 6 the sporozoites to have infectious sporozoites without
Pl. In preliminary experiments, the parasite was not intact oocysts. SodiruTl hypochlorite, sodium tauro‑
detect･,ed ti;1 day 5 l･)1. On day IS, a mean of 615 parasites chaiate, and trypsin have been used in the excystation of

Table rL Neutralization of C. muris spor02;oites by antiserum

Number of infected mice/number of mice examinedon pastinoculation days

Inoculun l
6

7

8

9

10

13

014

014

0/8

014

5/8

1/3

2/3

314

5/6

3/3

5/5

414

medium

0/2

ll2

214

2/2

3/3

3/3

Sp+distilled water

0/3

0/l

014

0/1

014

014

l x lO* sporozoites (sp)+ immune rabbit serum(1:lO) *Sp narmal rabbit serum(1:lO)

Sp

Dilution was with the mediurn.

237

oocysts of C. parvum parasitizing the intestine (Dubeyet al.. 1990), but in C. muris parasitizing the stomach,weights have been found (Nina et al.. 1992), unlike ourthese treatments were not necessary for excystation.

finding of sporozoite antigens with bands at 78, 47.5, andOocyst contamination (2.2%) of the sporozoites isolated
by isopycnic Percoll gradients (Arrowood and Sterling,

27.5 kDa. C. parvum sporozoites have surface antigensof 55, 28, 20, and 15 kDa that may be responsible for1987 ) and 0.06% oocyst contamination of the sporo‑zoites separated by DEAE‑cellulose anion‑exchange

induction of the neutralization antibodies (Mead et al..

chromatography (Riggs and Perryman, 1987) were
found in experiments with C. parvum. We collectedsporozoites of C. muris with 0.03% oocyst contamina‑tion by repeated centrifugation. The omission of the useof sodium hypochlorite probably allowed oocysts toadhere to the tube, accounting for the low contamina‑tion rate. The experiment with the sporozoite prepara‑tion treated with distilled water showed that the few
oocysts remaining had virtually no effect on results ofthis sporozoite infectivity assay.

Treatment with antiserum did not completely pre‑vent C. muris infection of mice. Nevertheless, theprepatent period was longer and parasites were fewer inmice inoculated with sporozoites treated with anti‑
serum. The result in mice seems to be correlated with

kDa, 54‑51 kDa, and in the region of low molecular

1988; Riggs et al.. 1989; Tilley et al.. 1991). The C.
muris sporozoite antigen of 27.5 kDa that was recog‑
nized by this antiserum seems to be similar to the C.

parvum sporozoite antigen of 28 kDa. Antibodies thatneutralize sporozoites reduce the infectivity of mer‑ozoites also, because C. parvum sporozoites and mer‑ozoites share epitopes (Bjorneby et al.. 1990; Tilley etal.. 1991). However, the 15‑kDa antigen of C. parvum
was not detected in C. muris from cattle (Tilley et al..
1991) .

The results from our experiments show that anti‑
bodies in anti‑C. muris serum may neutralize the viabil‑

ity and infectivity of C. muris sporozoites. Secretory
immunoglobulin of animals (e.g., cattle) resists diges‑
tion in the human gastrointestinal tract; when colostrum

was administered to patients, Saxon and Weinstein

the data on the viability of sporozoites in the in vitro(1987) neutralized gastric acidity to help increase suchassay. In infections by C. parvum, a high titer orresistance. Thus, immunotherapy with oral administra‑tion of monoclonal antibodies or of immune serum or
immune colostrum from suitable animals may be usefulconcentration of antibodies in the passively administer‑
ed bovine colostrum is said to be effective in reducing
the infection (Taghi‑Kilani et al.. 1990, Doyle et al..
1993) . Arrowood et al. (1989) found that neonatal micefor the control of gastric cryptosporidiosis, which might

be caused by C. muris in immunocornpromised patients.

receiving an oral dose of antisporozoite monoclonal
antibodies before oocyst inoculation are as susceptible
to infection as the control mice, but that daily oral
treatment with the same monoclonal antibodies resultsin fewer parasites in the infected mice.

The antiserum may include antibodies that can
agglutinate sporozoites. A monoclonal antibody that
agglutinates Eimeria tenella sporozoites has been prepared (Speer et al.. 1985). Another monoclonal anti‑
body causes circumsporozoite precipitation ( CSP )and abolishes the infectivity of malaria sporozoites(Yoshida et al.. 1980). CSP is accompanied with cyto‑
plasmic vacuolization of the sporozoites (Cochrane etal.. 1976). Riggs et al. (1994) have reported that bovine

antibodies against C. parvum have an immunother

ACKNOWLEDGEMENTS
We thank Ms. C. Latta and Mr. Y. Shin for editing
this manuscript.

REFEREN CES1 ) Anderson, B.C. (1989): Cmptosporidium spp. in cattle, 55‑

63. In Angus, K.W. and Blewett, D.A., (eds.) Cryptospo‑
ridiosis. Proceedings of the first international workshop.

Anirnal Disease Research Association, Edinburgh.
2 ) Arrowood, M.J. and Sterling, C.R. (1987) : Isolation of
Cmptosporidium oocysts and sporozoites using discontin‑uous sucrose and isopycnic Percoll gradients, J. Par‑
asitol., 73, 314‑319.

apeutic effect against cryptosporidiosis in severe com‑bined immune‑deficient (SCID) mice, and elicit a CSP‑

3 ) Arrowood, M.J., Mead, J.R., Mahrt, J.L., and Sterling, C.

like reaction to the sporozoites. Here also, someadministered antisporozoite monoclonal antibodies on
the outcome of Cmptosporidium parvum infections inmorphological changes observed in C. muris sporozoites
treated with heat‑inactivated antiserum suggested that
there was a CSP‑1ike reaction.
When antigens obtained from C. muris oocysts andsporozoites are used in immunoblotting, bands at 110

R. (1989): Effects of immune colostrum and orally

neonatal mice, Infect. Immun., 57, 2283‑2288.

4 ) Bjorneby, J.M., Riggs, M.W., and Perrymann, L.E.
(1990): Cmptosporidium parvum merozoites share neu‑
tralization‑sensitive epitopes with sporozoites, J. Im‑

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5 ) Cochrane, A.H., Aikawa, M., Jeng, M., and Nussen‑zweig, R.S. (1976): Antibody‑induced ultrastructuralchanges of malarial sporozoites, J. Immunol., 116, 859‑
867.

bovine serum, and immune bovine colostrum, Infect.
Immun., 58, 257‑259.
17) Riggs, M.W. and Perryman, L.E. (1987): Infectivity andneutralization of Cmptosporidium parvum sporozoites,
Infect. Immun., 55, 2081‑2087.

6 ) Doyle, P.S., Crabb, J., and Petersen, C. (1993): Anti‑
Cmptosporidium parvum antibodies inhibit infectivity invitro and in vivo, Infect. Immun., 61, 4079‑4084.
7 ) Dubey, J.P., Speer, C.A., and Fayer, R. (eds.) (1990):
Cryptosporidiosis of Man and Animals, CRC Press, Boca
Raton, Florida.

18) Riggs, M.W., McGuire, T.C., Mason, P.H., and Per‑8 ) Fayer, R., Perryman, L.E., and Riggs, M.W. (1989) :Hyperimmune bovine colostrum neutralizes Cmptospo‑
ridium sporozoites and protects mice against oocyst

parvum elicits a circumsporozoite precipitate‑1ike reac‑tion and has immunotherapeutic effect against persist‑ent cryptosporidiosis in SCID mice, Infect. Immun., 62,challenge, J. Parasitol., 75, 151‑153.

9 ) Fayer, R., Guidry, A., and Blagburn, B.L. (1990): Im‑munotherapeutic efficacy of bovine colostral immuno‑

globulins from a hyperirnmunized cow against
cryptosporidiosis in neonatal mice, Infect. Immun., 58,
2962‑2965.
10) Iseki, M., Maekawa, T., Moriya, K., Uni, S., and Takada,S. (1989): Infectivity of Cmptosporidium muris (strain
RN 66) in various laboratory animals, Parasitol. Res.,75, 218‑222.

11) Mata, L. (1986) : Cmptosporidium and other protozoa indiarrheal disease in less developed countries, Pediatr.

ryman, L.E. (1989) : Neutralization‑sensitive epitopes
are exposed on the surface of infectious Cmptosporidium
parvum sporozoites, J. Immunol., 143, 1340‑1345.
19) Riggs, M.W., Cama, V.A., Lee Leary, H., Jr., Sterling, C.

R. (1994): Bovine antibody against Cryptosporidium

1927‑1939. '20) Saxon, A. nd Weinstein, W. (1987) : Oral adrninistra‑tion of bovine colostrum anti‑cryptosporidia antibody
fails to alter the course of human cryptosporidiosis, J.
Parasitol., 73, 413‑415.

21) Speer, C.A., Wong, R.B., Blixt, J.A., and Schenkel, R.H.
(1985) : Capping of irnmune complexes by sporozoites ofEimeria tenella. J. Parasitol., 71, 33‑42.

22) Sterling, C.R., Gilman, R.H., Sinclair, N.A., Cama, V.,
Castillo, R., and Diaz, F. (1991) : The role of breast milkin protecting urban Peruvian children against
cryptosporidiosis. J. Protozool., 38, 23S‑25S.

Infect. Dis., 5. S117‑S130.
12) McDonald, V., Deer, R., Uni, S., Iseki, M., and Bancroft,23) Taghi‑Kilani, R., Sekla, L., and Hayglass, K.T. (1990):

G.J. (1992) : Immune responses to Cmptosporidium muris

infection: studies with B cell‑depleted mice, J. Immunol.,and Cryptosporidium parvum in adult immunocompetentor immunocompromised (nude and SCID) mice, Infect.

145, 1571‑1576.
24) Tilley, M., Upton, S.J., Fayer, R., Barta, J.R., Chrisp, C.

Immun., 60, 3325‑3331.
13) Mead, J.R., Arrowood, M.J., and Sterling, C.R. (1988):
Antigens of Cmptosporidium sporozoites recognized by

E., Freed, P.S., Blagburn, B.L., Anderson, B.C., andimmune sera of infected animals and humans, J. Par‑
asitol., 74, 135‑143.

14) Moon, H.W., Woodmansee, D.B., Harp, J.A., Abel, S.,
and Ungar, B.L.P. (1988): Lacteal immunity to enteric
cryptosporidiosis in mice: immune dams do not protect
their suckling pups, Infect. Immun., 56, 649‑653.
15) Nina, J.M.S., McDonald, V., Dyson, D.A., Catchpole, J.,
Uni, S., Iseki, M., Chiodini. P.L., and McAdarn, K.P.W.J.
(1992) : Analysis of oocyst wall and sporozoite antigens
from three CIOiptosporidium species, Infect. Immun., 60,

1509‑1513.

16) Perryman, L.E., Riggs, M.W., Mason, P.H., and Fayer,
R. ( 1990): Kinetics of Cmptosporidium parvum sporo‑zoite neutralization by monoclonal antibodies, immune

The role of humoral immunity in Cmptosporidium spp.

Barnard, S.M. (1991): Identification of a 15‑kilodaltonsurface glycoprotein on sporozoites of Cryptosporidium
parvum. Infect. Immun., 59, 1002‑1007.

25) Tzipori, S., Roberton, D., and Chapman, C. (1986) :Remission of diarrhoea due to cryptosporidiosis in an
immunodeficient child treated with hyperimmune bovine
colostrum. Br. Med. J., 293, 1276‑1277.
26) Uni, S., Iseki, M., Maekawa, T., Moriya, K., and Takada,S. (1987): Ultrastructure of Cmptosporidium muris(strain RN 66) parasitizing the murine stomach, Par‑
asitol. Res., 74, 123‑132.

27) Yoshida, N., Nussenzweig, R.S., Potocnjak, P., Nussen‑zweig, V., and Aikawa, M. (1980): Hybridoma produces
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J pn. J. Trop.

239

Med Hyg., Vol. 23, No. 4 1995, pp. 239‑252

THE SIMULIIDAE (DIPTERA) FROMBOUGAlNVILLE ISLAND,PAPUA NEW GUlNEAHIROYUKI TAKAOKA

Received September 10, 1995/Accepted October 13, 1995

Abstract: Seven black fly species were identified, chiefly on the basis of reared adults, pupae and maturelarvae collected frorn Bougainville Inland, Papua New Guinea. A11 were assigned to the genus Simulium
Latreille s. l. and were further classified into two subgenera, i.e., six species in Simulium (Morops) and one
in Simulium (Gomphostilbia) . S. (M.) noroense and S. (G.) hiroshii were newly recorded from BougainvilleIsland, and five other species were described as new species. Under the subgenus Morops, a new species‑
grotip was proposed.

Smart and Clifford (1965) reported 52 species of

1. Simulrnm ( Gomphostllbra) hlroshll Takaoka 1994

Simulium Latreille s. l. from Papua New Guinea.
Crosskey (1967) classified these species, and those ofAustralia and the Western Pacific, into subgenera andSimulium (Gomphostilbia) hiroshii Takaoka, 1994: 97‑
101.

species‑groups. Stone and Maffi (1971) recorded two

black fly species from Guadalcanal, the Solomon

SPECIMEN EXAMINED. I

Islands. Seven other species were also collected fromcaught inside the house, 10/39 Arawa, Bougainville(ANlO , in alcohol,

the Solomon Islands (Takaoka, 1994; Takaoka andIsland, Papua New Guinea, 1. X. 1987, by C. Yule.

Suzuki, 1994 & 1995) . However, no study has previously
been made on black flies of Bougainville Island.
In this study seven species of the genus Simulium s.

DISTRIBUTION. Bougainville Island (new record)l. were recognized. They were reared or light‑trappedcanal Island) .

and Solom6n Islands (New Georgia Island and Guadal‑

adults, pupae and mature larvae collected from
Bougainville Island in 1965 by Dr. R.W. Crosskey and in

REMARKS. The female specimen well agreed with the

1987‑1989 by Miss Cathy Yule.
Descriptions of five new species are given, and aoriginal description of S. ( G. ) hiroshii reported from the

new species‑group within the subgenus Morops is
proposed.

Solomon Islands (Takaoka, 1994). The female of thisspecies is easily distinguished from the other Morops

History Museum (BMNH), London, U.K., and in thespecies by the large basal tooth of the tarsal claws aswell as the bare pleural membrane. It is interesting thatthe female genitalia of S. ( G. ) hiroshii including theAustralian National Insect Collection (ANIO , Canber‑spermatheca with internal setae are very similar tora, Australia.

those of the shermoodi‑group of the subgenus Morops, as

Type specimens will be deposited in the Natural

noted later.

Genus Simulium Latreille s. l.

Subgenus Morops Enderlein

Subgenus Gomphostilbia Enderlein This subgenus was well defined by Crosskey (1967)
This subgenus was first defined by Crosskey (1967) and was divided into several species‑groups (Crosskey,
and recently revised by Takaoka and Davies (1995). 1967; Colbo, 1976). The subgenus Morops is character‑
This is characterized by the bare pleural membrane and ized in the adults by the presence of hairs on bothhaired katepisternum in adults of both sexes. pleural membrane and katepisternum. In this study, all
Division of Medical Zoology, Oita Medical University, Hasama, Oita 879‑55, Japan240

the seven, but one in the subgenus Gomphostilbia Ender‑
lein, were assigned to this subgenus; two species were
further classified into the clathrinum‑group and four
were into the newly proposed sherwoodi‑group.

with a large, transpareht pouch (Fig. 2a) , which
expands laterally, thus partially covering the pit‑likeorgan; tubular filaments subequal in length and thick‑ness to each other, with several annular constrictions

along their length, and, on their surface, with many(A) clathrinum ‑group
This species‑group is easily distinguished from

10ngitudinal ridges, covered with somewhat larger tuber‑cles on ridges and with smaller ones on interspaces (Fig.
2b) ; slender filaments subequal in length and thickness

other species‑groups in having the pit‑like cuticularorgan at the base of pupal gill filaments (Crosskey,

to each other, with well‑defined annular ridges and1967). The identification of species within this species‑

furrows on their surface, and covered with minute tuber‑group is usually difficult in the adult and larval stages

shape of gill filaments. Takaoka and Suzuki (1995)
proposed plo ratio ( = A/B as shown in Fig. 1) tocles; all filaments pale yellow to dark yellow except
basal portion of tubular filaments dark brown. Abdo ‑men. Terga I and 2 almost transparent, without tuber‑cles; tergum I with I Iong seta on each side, tergum 2
with 5 simple spinous setae and I much longer seta on

compare relative sizes of the pit‑1ike organ.

each side; terga 3 and 4 each with 4 hooked spines

The clathrinum‑group species were recorded from
Australia (one species) (Mackerras and Mackerras,
1948), Papua New Guinea (four species) , New Britain(one species) (Smart and Clifford, 1965) and Solomon

directed forward on each side; terga 7 and 8 each with abut is most reliably made in the pupal stage by compar‑
ing the size and feature of pit‑like organ, as well as the

Islands (four species) (Takaoka and Suzuki, 1995). Our
recent investigation in Indonesia indicates the presenceof species of this species‑group in Halmahera (twospecies) , Ambon (two species) , Seram (three species) ,Biak (one species) and lrian Jaya (two species) , extend‑

ing its distribution up to Halmahera Island in the west
(unpublished data) . Interestingly, more specialized pit‑

like organs, such as that of S. (M.) aropaense. are atpresent recognized only in four species, each on Papua
New Guinea, Bougainville Island, the Solomon Islands
and Australia.

transverse row of spine‑combs and comb‑like groups ofminute spines directed caudad on each side; tergum 9with comb‑like groups of minute spines and a pair ofsmall, terminal hooks but lacking spine‑combs. Sternum
4 with I distinct simple hook and a few minute setae oneach side; sternum 5 with a pair of bifid hooks on eachside; sterna 6 and 7 each with a pair of inner bifid andouter simple hooks on each side; Iast segment without
grapnel‑like hooklets ventrolaterally. Cocoon (Fig. 3).

Shoe‑shaped, moderately woven, somewhat extending
ventrolaterally; anterior margin thickly woven; individ‑
ual threads visible; interspaces thinnly walled but with‑out perforations.

Mature larva. Body length 4.0‑4.6 mm. Body greyish.

2. Simulium (Morops) aropaense sp. nov.

Cephalic apotome pale yellow, with faint, positive headspots in 24 Iarvae, but markedly darkened medially and

DESCRIPTION. Female and Male. Unknown.

posteriorly on posterior l/2 (Fig. 4) in 30 Iarvae.
Antenna with 3 segments and apical sensillum, Ionger

Pupa. Body length (excluding gill filaments) ca. 2.6mm. Head. Integument dark yellow, bare, with 4 pairs

than stem of labral fan; proportional lengths of 3 seg‑ments from base to tip 1.0 : 1.0 : 0.64. Labral fan with ca.

of trichomes, all long and simple. Antennal sheath with

35 main rays. Mandible (Fig. 6) with comb‑teeth

smooth surface. Thorax. Integument dark yellow, bareon anterior 1/2 except area along middle longitudinal
suture with round tubercles, and moderately covered
with cone‑shaped tubercles on posterior 1/2; pit‑likedecreasing in size from Ist to 3rd; mandibular serration

organ large, circular, thickly fringed with distinct, comb‑

like processes, and its diameter (A in Fig'. 1) nearly as

long as the distance (B in Fig. 1) from its innermost
margin to middle longitudinal suture of thorax, i.e., plo
ratio ( = A/B) 1; thoracic trichomes 6 pairs (5 antero‑dorsally and I posterolaterally) , all long and simple.
Gill (Figs. I & 2) composed of 2 inflated, tubular fila‑

ments each with 2 slender filaments at its apex (total
length ca. 1.8 mm) ; basal stalk of moderate length andcomposed of 2 teeth (1 Iarge and I smalD, with orwithout supernumerary serrations. Hypostomium (Fig.
7) with a row of 9 apical teeth; median tooth as long as
each corner tooth, Ionger than 3 intermediate teeth oneach side; Iateral margin serrate; hypostomal bristles 6
or 7 in number per side, Iying slightly divergent poste‑
riorly from lateral margin. Postgenal cleft (Fig. 5)nearly triangular, wide basally, gradually narrowed
anteriorly with a narrow apex, which is approaching
posterior border of hypostornium. Thoracic cuticle

bare. Abdominal cuticle bare except last segment
moderately covered with numerous colorless setae on

241

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a

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Figs. 1‑8.

7

Pupal and larval characters of Simulium (Morops) aropaense sp. nov. 1, pupal gill andanterior portion of thoracic integument (dorsal view) showing pit‑like organ, trichomes andtubercles along middle longitudinal suture (A, diameter of pit‑like organ; B, distance frominnermost margin to middle longitudinal suture) ; 2, pupal gill filaments (side view) showing
a transparent pouch (a) and an enlargement of surface pattern of inflated filament (b) ; 3,pupa and cocoon; 4 & 5, Iarval head capsule (4, dorsal view; 5 ventral view) ; 6, apex of larval
mandible; 7, Iarval hypostomium; 8, Iarval rectal papilla. Scale bar 0.02 mm for Fig. 2b.

242

(

C¥‑

r¥
C n Io

"

.

n

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oO

ooooOO o

A,

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12

14
a e

OQ

c o eG,tc e4""

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ef/v/:/1/ D
o

13

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11
FlgS. 9‑14.

Pupal characters of Simulium (Morops) kietaense sp. nov. 9 & 10, tubercles on frontal
integument (9, dorsal view; 10, side view) ; 11, base of gill and anterior part of thoracic
integument showing a pit‑like organ, trichomes and distribution of tubercles (dorsal view) ;12, pit‑1ike organ (anterodorsal view) ; 13, basal portion of gill filaments (side view) ; 14,
terminal hooks. Scale bar 0.02 mm for Figs. 9, 10 & 12.

each side of anal sclerite. Rectal papilla (Fig. 8) com‑filaments. Among nine known species, S. (M. ) Ialokien ‑pound, each of 3 Iobes with ca. 4 finger‑like, secondary

se from central Papua New Guinea (Smart and Clifford,1965), S. (M.) clathrinum from Australia (Mackerras

lobules. Anal sclerite of usual X‑form, with anteriorarms ca. 0.7 x as long as posterior ones, broadly sclero‑

tized at base. Ventral papillae absent, though last
abdominal segment expanded ventrolaterally to formand Mackerras, 1948), and S. (M.) solomonense from
New Georgia Island. Solomon Islands (Takaoka anddouble bulges on each side. Posterior circlet with ca. 94

Suzuki, 1995) have a pit‑1ike organ fringed with comb‑
like processes. The pit‑like organ of this species is as

rows of up to 16 hooklets per row.

large as those of S. (M.) Ialokiense and S. (M.)

TYPE SPECIMENS. Holotype pupa (BMNH), slide

somewhat larger than that of S. (M.) clathrinum (plomounted, together with its cocoon, collected from astream near Aropa, 13 Km S of Kieta, BougainvilleIsland, 8. VII. 1965, by R.W. Crosskey. Paratypes: 19

ratio ca. 0.7). However, S. (M.) aropaense differs fromsolomonense (plo ratio 1.1 and 1.0 respectively) but isthese three known species in the arrangement of thepupal gill filaments.

pupae and 53 mature larvae (BMNH), same data as
holoty pe.

3. Simulium (Morops) kietaense sp nov

DISTRIBUTION. Bougainville Island.

DESCRIPTION. Female and Male. Unknown.
Pupa. Body length (excluding gill filaments) ca. 2.4

REMARKS. The species aropaense refers to the locality
where this species was collected.

mm. Head. Integument dark yellow, moderately cov‑

This new species belongs to the clathrinum‑groupof the subgenus Morops, defined by Crosskey (1967) bywith 4 pairs of trichornes, all long and simple. Antennal

having the pit‑like organ (Fig. 1), as well as four gillered with round tubercles (Figs. 9 & 10) on frons, andsheath with smooth surface. Thorax. Integurnent dark
yellow, sparsely covered with round tubercles on ante‑

243rior 1/2, densely along middle longitudinal suture, and(M.) kietaense (which had been identified at that time as

moderately covered with cone‑shaped tubercles on pos‑terior 1/2; pit‑like organ (Fig. 11) small, circular,
moderately fringed with distinct, comb‑like processes

S. (M.) raunsimnae by Dr. Crosskey according to theattached labeD , Bougainville Island should be omitted atpresent from the range of S. (M.) raunsimnae.

(Fig. 12) , plo ratio 0.33; thorax with 5 pairs of trichomes

anterodorsally and I pair posterolaterally, all long andsimple; 5 anterior pairs are much longer than postero‑lateral one. Gill (Figs. 11 & 13) composed of 4 slenderfilaments in 2 pairs (outer and inner) , subequal in thick‑ness and length (ca. 1.5 mm) to each other, though inner

pair slightly shorter than outer pair, and graduallytapered toward apical tip; basal stalk somewhat elon‑
gate, with a transparent pouch (Fig. 13) ; inner and outerpairs of filaments sessile or very short‑stalked (Fig. 1D ;surface of filaments with well‑defined ridges (forming
reticulate pattern on basal 1/4) , covered with somewhat

larger tubercles on ridges and with smaller ones on(B) sherwoodi‑group
The following three new species are distinct among
species of the subgenus Morops in having the spermathe‑ca with internal setae and the parameres of the male

genitalia with distinct hooks. The three ungrouped
species, S. (M.) sherwoodi Stone and Maffi, S. (M.)
herei Takaoka and Suzuki, and S. (M.) Iioroen:se Ta‑
kaoka and Suzuki, all from the Solomon Islands, sharethese characters (except S. (M. ) kerei of which thefemale is still unknown) . The new species‑group is here

proposed to relate the six species which possess thefollowing combination of characters.

interspaces; filaments dark yellow to brown, paler dis‑

DIAGNOSIS. Female. Mandible with teeth on

tally. Abdomen and Cocoon. As in S. (M.) aropaense.
Mature larva. Almost indistinguishable from S. (M.)

inner margin but no or a few teeth on outer margin; fore
basitarsus somewhat dilated, without dorsal hair crest;

aropaense except following features: Body length 4.3‑4.6

tarsal claw with a minute basal tooth; abdomen lacking

mm. Body greyish. Cephalic apotome pale yellow, with
faint, positive head spots in I Iarva, but markedly

dense covering of pubescence on dorsal and lateraldarkened medially and posteriorly on posterior 1/2 in 3

theca with internal setae. Male. Fore basitarsus some‑what dilated, without dorsal hair crest; hind basitarsus
slender, not inflated; abdomen lacking dense covering ofpubescence, and with a pair of shiny patches on terga 2
and 5‑7; ventral plate setose ventrally, posteriorly anddorsally; paramere with distinct hooks. Pupa. Frontal

larvae. Mandibular serration composed of 2 teeth (1large and I smalD, with I or 2 supernumerary serra‑
tions. Rectal papilla compound, each of 3 Iobes with 4 or
5 finger‑like, secondary lobules. Anal sclerite of usual X‑

form, with anterior arms ca. 0.8 x as long as posteriorones, broadly sclerotized at base.

TYPE SPECIMENS. Holotype pupa (BMNH), slide
mounted, together with its cocoon, collected from astream near Aropa, 13 km S of Kieta, BougainvilleIsland, 8. VII. 1965, by R.W. Crosskey. Paratypes: 4pupae and 4 mature larvae (BMNH), same data as
holotype.

DISTRIBUTION. Bougainville Island.
REMARKS. This new species is also assigned to theclathrinum‑group by having the pit‑like, cuticular organ(Fig. 12) in the pupa. As in S. (M.) aropaense this organ

is furnished with comb‑like processes along the inner
margin, but is much smaller than that of the former
species, and also those of S. (M.) clathrinum. S. (M.)
lalokiense and S. (M. ) solomonense.

Crosskey (1989) added Bougainville Island as arange of S. (M.) raunsimnae in his list of Simuliidae in

the Australasian and Oceanian Regions. Since this wasmade on the basis of the present pupal specimens of S.

surface, and shiny on terga 2 and 5‑8 (or 6‑8) ; sperma‑trichomes 2 or 3 pairs, simple or branched; gill filaments

5 or 6 in number per side, all slender or I or 2 of them
inflated; terminal hooks of usual cone‑shape or plate‑like; cocoon simple, wall‑pocket‑shaped. Larva.
Hypostomium with smooth lateral margins; mandible
without supernumerary serrations; ventral papillae welldeveloped.

SPECIES INCLUDED. S. (M.) kerei. S. (M.)
noroense. S. (M.) pangunaense sp. nov., S. (M.) peggyae
sp. nov., S. (M.) sherwoodi. S. (M.) yuleae sp. nov.
S. (M.) kerei, though the adult female is still un‑

known, apparently is assignable to this species‑group
because its adult male has the paramere with distinct
hooks. This species had been tentatively placed in the
farciminis‑group mainly because of the similar arrange‑
ment of the pupal gill filaments (Takaoka and Suzuki,1994). As shown later, S. (M.) kerei is very similar toS. (M.) pangunaense.

This species‑group is distributed only in the
Solomon Islands and Bougainville Island. This fact,
together with the close morphological similarities of all

the six species, suggests that speciation within this244

species‑group might have occurred after an ancestor
species had arrived on these islands. It is intriguing thatamong four species of this species‑group found inspecies were not found in other species‑groups ofMorops, but found in the subgenus Gomphostilbia, aspreviously pointed out by Takaoka (1983). Future

Bougainville Island, successive changes appear to havetaken place in morphology of the pupal gill filaments, inparticular, of lower paired filaments, as illustrated in
Figs. 30‑35. If the decreased number and inflation offilaments are considered to be apomorphic, Iower two
filaments must have changed as follows: first, Iower two
filaments which were slender (Fig. 30) became inflated(Fig. 31), then, stalk of lower filaments elongated, as
shown in Figs. 32 and 33, resulting in the loss of one oftwo branches (Fig. 34) and, finally, the remaining fila‑
ment have lost its apical slender portion (Fig. 35).

studies of these Philippine, Solomon and Bougainville

Taking these successive changes into consideration, it iswith a few dark hairs; frontal ratio 2.1 : 1.0 : 3.0. Frons‑

presumed that S. (M.) noroense and S. (M.) sherwoodi
are most primitive, S. (M.) pangunaense and S. (M.)
kerei are most derivative, while S. (M.) yuleae and S.
(M.) peggyae are intermediate in specialization. How‑

head ratio 1.0 : 4.6. Fronto‑ocular area (Fig. 15) wellever, this is not conclusive since there are other charac‑

ish black except scape, pedicel and base of Ist flagellarters which suggest the opposite direction of specializa‑
tion. S. (M.) noroense has a smaller basal tooth of thefemale claw and a wider fore basitarsus (both of which

segment yellowish when viewed from above (scape,are generally regarded as being a more derived charac‑
ter than a large claw tooth and a narrow basitarsus,
respectively) than those of the other three Bougainvillespecies (Takaoka and Suzuki, 1995); further, pupal
terminal hooks are of cone‑shape (derived form) in S.

(M.) noroense and S. (M.) sherwoodi, whereas thesehooks are plate‑1ike (supposedly primitive form) in theother species, as shown in Figs. 38‑40.

This species‑group appears closely related to theeight Morops species (all so far ungrouped) describedfrom the Philippines (Takaoka, 1983) because its mem‑bers share many characters, such as the paramere withspecies, though far apart geographically from eachother, may clarify the phylogenetic relationship between

the two subgenera.

4. Simulium (Morops) yuleae sp. nov.

DESCRIPTION. Female. Body length ca. 2.2 mm.
Head. Slightly narrower than thorax. Frons darkbrown, whitish grey pruinose, dull, densely covered withyellowish white, recumbent pubescence interspersed

developed. Clypeus dark brown, densely covered withyellowish white pubescence, interspersed with several
dark hairs. Antenna composed of 2 + 9 segments, brown‑

pedicel and basal few flagellar segments yellow, apical 5

flagellar segments brownish black when viewed fromfront). Maxillary palp with 5 segments; proportional
lengths of 3rd, 4th and 5th segments 1.0 : 1.2:2.3; 3rdsegment somewhat enlarged, sensory vesicle (Fig. 16)
much elongate, 2.3 x as long as wide, 0.55 x as long as
3rd segment, with very large opening distally. Maxilla‑ry lacinia with 10 inner and ll outer teeth. Mandiblewith ca. 26 small inner teeth and lacking outer ones,
though outer margin weakly erose near apex. Cibariumsmooth (nearly as in Fig. 27). Thorax. Scutum blackish
brown in ground color, whitish grey pruinose, with 3
dark, rather broad, Iongitudinal vittae, densely covered

with whitish yellow, recumbent pubescence, also with

distinct hooks, a pair of dorsolateral shiny spots each on

dark brown pubescence on longitudinal vittae, andthe male abdominal segrnents 2, 5, 6 and 7, pupal gillwith six filaments (four filaments in one Philippine
species) , plate‑like terminal hooks on the pupal abdo‑

sparsely with dark upright hairs on prescutellar area.
Scutellum blackish brown with whitish yellow pubescen‑ce and long dark upright hairs along posterior margin.

men (of usual cone‑shape in one Philippine and two
Solomon species) , wall‑pocket‑shaped cocoon, Iarval
hypostomium with smooth lateral margins, Iarval man‑
dible without supernumerary serrations and the pres‑ence of the larval ventral papillae. However, these

Postscutellum blackish brown, whitish grey pruinose

Philippine species differ frorn the sherwoodi‑group in
having no internal setae of the spermatheca, a few hairson the pleural membrane, a large tooth on the female
tarsal claw (small tooth in one species) and grapnel‑likehooklets on the last segment of the pupal abdomen. Itshould be noted that most characters shared by mem‑bers of this species‑group and eight Philippine Morops(shining in light) , bare. Pleural membrane covered withca. 80 yellowish white pubescence and several darkhairs; several hairs also on sclerotized portion justanterior to pleural membrane. Katepisternum longer
than deep, dark brown, whitish grey pruinose (shining inlight) , with numerous dark hairs and yellowish whitepubescence; sulcus distinct. Legs. Foreleg: coxa andtrochanter yellow; femur dark yellow to pale brown;
tibia yellow with distal 1/4 black, outer surface widely
white shining when illuminated; tarsus black; basitarsus
somewhat dilated, ca. 6.2 x as long as its greatest width.

245

Midleg: coxa dark brown; trochanter yellow; femur

Postscutellum brownish black, white pruinose (white

yellow with apical cap somewhat dark; tibia yellow onshining in light) , bare. Pleural membrane with ca. 75
yellow pubescence; 16‑20 additional pubescence present
just before it. Katepisternum longer than deep, withyellow and dark hairs. Legs. Foreleg: coxa pale yellow;

basal 1/3 (posterior surface) or on basal 1/2‑2/3 (ante‑rior surface) , brownish black on rest; posterior surfaceof tibia widely white shining when illuminated; tarsus
black. Hind leg: coxa dark yellow; trochanter yellow;
femur yellow with apical cap brown; tibia yellow onbasal 2/3, brownish black on apical 1/3; posterior sur‑face of tibia widely white shining when illuminated;tarsus brown with basal 3/5 of basitarsus and basal 1/2

of 2nd tarsomere white; basitarsus slender, parallel‑
sided, ca. 7.0 X as long as wide. Calcipala (Fig. 17) welldeveloped, ca. 1.5 x as long as wide. Pedisulcus welldeveloped. Hind femur and tibia ca. 2.4 x and 1.8 x as
wide as hind basitarsus, respectively. All femora andtibiae densely covered with scale‑like hairs on outersurface. Claws (Fig. 18) each with a round, minutebasal tooth. Wing. Length 2.0 mm. Costa with spinules
as well as hairs. Subcosta haired except distal 1/4 bare.

trochanter yellow; femur dark yellow to pale brown;
tibia yellow with distal 1/3 brownish black, and white
shinihg in light on outer surface; tarsus brownish black;basitarsus somewhat dilated, ca. 6.4 x as long as its
greatest width. Midleg: coxa dark brown; trochanter
yellow; femur yellow with distal cap sornewhat dark‑
ened; tibia yellow on basal 1/2, brown on distal 1/2
though its border not well defined; tarsus brownish
black. Hind leg: coxa dark yellow; trochanter yellow;

femur yellow with distal cap brownish black; tibia
yellow on basal 2/3, brownish black, on rest; tarsus
brownish black except basal 3/5 of basitarsus and basal
l/2 of 2nd tarsomere white; basitarsus slender, parallel‑

Hair tuft on stem vein dark brown. Basal portion ofradius fully haired. Abdomen. Basal scale brown withsided, ca. 5.8 x as long as wide. Hind femur and tibia
ca. 2.1 x and 1.8 x as wide as hind basitarsus, respec‑tively. Calcipala well developed (nearly as inbutfringe of yellowish white hairs. Dorsal surface of abdo‑slightly shorter) , ca. 1.3 x as long as wide. Pedisulcus

men black, with dark hairs; tergite of 2nd segment

well developed. All femora and tibiae densely covered

whitish pruinose, tergites of segments 5‑8 shiny. Genita‑

with scale‑like hairs on outer surface. Wing. Length 1.9

lia (Figs. 19 & 20). Sternal plate undeveloped on 7th
abdominal segment. Sternite 8 bare medially, with ca.

mm. Other features as in

35 dark macrosetae on each side. Anterior gonapo‑

fringe of yellow hairs. Dorsal surface of abdorninal
segments brownish black, covered with dark hairs; seg‑ments 2, 5‑7 each with a dorsolateral pair of shiny,

physis triangular, thin, membraneous, covered denselywith microsetae; inner margin well sclerotized. Genital
fork of usual inverted‑Y form, with arms folded medi‑
ally; each arm with a small projection directed for‑wards. Paraproct not produced ventrally, with ventral
margin somewhat depressed in middle. Cercus 0.57 x as
long as wide, with posterior border gently rounded whenviewed laterally. Spermatheca (depressed) well sclero‑
tized except small area near tubal juncture unsclero‑
tized; internal setae present.

Male. Body length ca. 2,2i mm. Head. Wider thanthorax. Upper eye consisting of 12 vertical colurnns and13 horizontal rows of large facets. Clypeus brownish
black, whitish pruinose, covered densely with yellow

pubescence, interspersed with dark hairs. Antennacomposed of 2 + 9 segments, yellow, but 4 or 5 apicalsegments brown; Ist flagellomere somewhat elongate,ca. 1.6 x as long as 2nd one. Maxillary palp with 5
segments; proportional lengths of 3rd, 4th and 5th seg‑

except subcosta with a few

hairs. Abdomen. Basal scale brownish black, withwhitish pruinose patches; those on segment 2 widelyconnected in middle. Genitalia (Figs. 21‑26). Coxite
nearly rectangular in ventral view, much longer thanwide. Style shorter than coxite, tapered toward apicaltip, gently curved inwards, with an apical spine. Ventral

plate transverse, ca. 0.4 x as long as wide, with bothsides moderately angulated, widely produced ventrally
along posterior margin, with microsetae on ventral,posterior and dorsal surface; basal arms converging.
Paramere with 3 Iong hooks and several indistict ones.
Median sclerite weakly sclerotized, plate‑like, slightly
widened toward tip, with round tip.
Pupa. Body length (excluding gill filaments) ca. 2.5mm. Head. Integument yellow, moderately covered
with tubercles; a pair of facial trichomes long, simple orments 1.0 : 1.2:2.8; sensory vesicle oblong, 0.2 x as longbifid, and 3 pairs of frontal trichomes all long andsirnple. Antennal sheath with smooth surface. Thorax.
Integument yellow, sparsely covered with tubercles; 5

as 3rd segment, with opening near distal end. Thorax.

anterodorsal pairs of trichomes all long, simple or bifid,

Scutum brownish black, densely covered with golden
yellow pubescence. Scutellum brownish black withgolden yellow pubescence and dark upright hairs.

1 posterolateral pair long and simple. Pit‑like organ
absent. Gill (Fig. 31) composed of 2 inflated, tubular
filaments and 4 slender filaments arising dorsally from

246

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Figs. 15‑29.

Adult characters of the sherwoodi‑group of Simulium (Morops). 15, fronto‑ocular area;
16, 3rd maxillary palpal segment showing a sensory vesicle; 17, distal tip of hindbasitarsus and 2nd tarsomere showing calcipala and pedisulcus; 18, claw with a minutetooth; 19, genitalia in situ (spermatheca omitted; ventral view) showing 8th sternite (a),gonapophysis (b) , genital fork (c) , paraproct (d) and cercus (e) ; 20, paraproct and cercus(side view) ; 21, coxite (a) and style (b) (ventral view) ; 22‑24, ventral plate (22, ventral view;23, side view; 24, end view) ; 25, paramere; 26, median sclerite; 27,

cibarium; 28, genital fork;
29, spermatheca. 15‑25, S. (M.) yuleae sp. nov.; 27, S. (M.) peggyae sp. nov.; 28 & 29, S. (M.)

pangunaense sp. nov.

basal stalk of tubular filaments; tubular filaments sub‑equal in length (ca. 1.0 mm) and thickness to each other(although somewhat varied individually) , Iying horizon‑

apical filament which was cut off, not allowing thetally, parallel‑sided or somewhat divergent, directing

tubular filaments with many ridges forming irregularpatterns, covered with somewhat larger tubercles on

forwards and each apex tapered, bearing a slendermeasurement of its length; outer tubular filament withseveral annular constrictions along its length; surface of

247

31

30

Figs. 30‑35.

32

:

33

34

35

Pupal gill filaments of the sherwoodi‑group of Simulium (Morops) . 30, S. (M.) noroense; 31,
S. (M.) yuleae sp. nov.; 32‑34, S. (M.) peggyae sp. nov.; 35, S. (M.) pangunaense sp. nov. Scalebar 0.5 mm for Figs. 30‑35.

ridges and with smaller ones on interspaces; slenderwoven; individual threads visible but without perfora‑

filaments subequal in length (0.6‑0.8 mm) and thickness
to each other, with annular furrows on their surface, andcovered with minute tubercles; all filaments pale yellow.

tions.

Mature larva. Body length 5.4 mm. Body greyish.

Abdomen. Terga I and 2 almost transparent, without

Cephalic apotome pale yellow with distinct, positive
head spots. Antenna with 3 segments and apical sensil‑

tubercles; tergum I with I Iong seta on each side; tergum

lum, Ionger than stem of labral fan; proportional lengths

2 with 5 simple spinous setae and I much longer seta onof 3 segments from base to tip 1.0 : 1.2: 1.1. Labral fan

each side; terga 3 and 4 each with 4 hooked spines
directed forward on each side; terga 6‑9 each with awith 32 main rays. Mandible (Fig. 42) with comb‑teeth

transverse row of spine‑combs and comb‑1ike groups ofminute spines directed caudad on each side, thoughspine‑combs on tergum 9 are much smaller than thosecomposed of 2 teeth (1 Iarge and I smalD , without
supernumerary serrations. Hypostomium (Fig. 43) with
9 apical teeth in row; each corner tooth longer thanon other terga; tergum 9 with a wide, plate‑1ike, termi‑

median tooth, which is longer than 3 intermediate teeth

decreasing in size from Ist to 3rd; mandibular serration

nal hook, which is moderately serrate along outeron each side; Iateral margin not serrate; hypostomal

margin (Fig. 38). Sternum 4 with I distinct bifid hook

bristles 5‑7 ,in number per side lying slightly divergent

and I spinous seta on each side; sternum 5 with a pair ofposteriorly from lateral margin. Postgenal cleft (Fig.

bifid hooks on each side; sterna 6 and 7 each with a pair44) nearly triangular, gradually narrowed anteriorly, ca.

of inner bifid and outer simple hooks on each side; Iast

2.9 x as long as postgenal bridge. Thoracic cuticle bare.

segment without grapnel‑like hooklets ventrolaterally.

Abdominal cuticle bare except last segment which ismoderately covered with numerous colorless setae oneach side of anal sclerite. Rectal papilla compound,

Cocoon. Wall‑pocket‑shaped, moderately woven, wide‑
ly extending ventrolaterally; anterior margin thickly

248

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41

Figs. 36‑44.

c) c:) oJ

37

42

Pupal and larval characters of the sherwoodi‑group of Simulium (Morops). 36 & 37,
tubercles on frontal integurnent (36, top view; 37, side view) ; 38‑40, terminal hooks; 41, 9th

abdominal segment showing a pair of terminal hooks, a transverse row of spine‑combs andscattered tubercles (dorsal view) ; 42, apex of larval mandible; 43, Iarval hypostomium; 44,
larval head capsule (ventral view) . 38, 42‑44, S. (M.) yuleae sp. nov.; 39, S. (M.) pe :yae sp.
nov.; 36, 37, 40 & 41, S. (M.) pangunaense sp. nov. Scale bar 0.01 mm for Figs. 36 & 37.

each of 3 Iobes with 3‑5 finger‑like, secondary lobules.

VIII. 1988, I (ANlO , in alcohol together with pupal
skin and cocoon, same data as holotype except date: 29.

Anal sclerite of usual X‑form, with anterior arms ca.
0.9 x as long as posterior ones, broadly sclerotized at

VIII. 1988, 6 pupae and I mature larva (ANlO , database. Ventral papillae present. Posterior circlet with 98

same as holotype except date: 18.X.1987; I mature larva

rows of up to 18 hooklets per row.

(ANlO , collected from a stream flowing from Lake

TYPE SPECIMENS. Holotype c71 (ANlO, reared from

Loloru to Mivo River (1515 m in altitude) , Mt. Loloru,
Bougainville Island, Papua New Guinea, 8.X.1988, by C.
Yule.

pupa, pinned together with pupal skin and cocoon, col‑lected from Konaiano Creek (720 m in altitude) , Pan‑guna, Bougainville Island, Papua New Guinea, 24. IX.

1988, by C. Yule. Allotype

DISTRIBUTION. Bougainville Island.

(ANlO , reared frompupa, same data as holotype except date: 12. 111. 1989.
Paratypes: I c71 , I(BMNH), slide mounted, emerged
from pupa, same data as holotype except date: 26. 111.
1989, 2 (ANlO , pinned together with pupal skin andcocoon, same data as holotype except date: 14 and 29.

REMARKS. The species name yuleae acknowledges
Miss C. Yule who kindly gave simuliid specimens col‑lected from Bougainville Island.
The adults of this species, as well as the following

two new species, differ from those of S. (M.) shermoodi

249

and S. (M.) noroense from the Solomon Islands in lack‑ing the brownish pubescence on the median dorsal sur‑face of the scutum, and also from S. (M.) sherwoodi by
their frons densely covered with pubescence, and from S.

(M.) noroense by their femora which are much paler
than that of S. (M.) noroense. The pupa of S. (M.)
yuleae is easily separated from the other related species
by the fused, plate‑like, terminal hook (Fig. 38) , as wellas the arrangement of the six gill filaments of which two
are inflated (Fig. 3D.

basitarsus, respectively. Wing. Length 2.0 mm.
Pupa. Body length (excluding gill filaments) ca. 2.4mm. Head. Integument pale yellow to dark yellow,
bare; a pair of facial trichomes long, simple or bifid, and3 pairs of frontal trichomes all long and simple (1 or 2

pairs bifid in some pupae) . Antennal sheath withsmooth surface. Thorax. Integument pale yellow todark yellow, bare except posterodorsal area moderately
covered with tubercles; 5 anterodorsal pairs of tri‑chomes all long, simple or bifid, I posterolateral pairlong and simple. Pit‑like organ absent. Gill (Figs. 32‑

5. Simulium (Morops) peggyae sp. nov.

34) composed of I inflated, tubular filament and 4

DESCRIPTION. Female. As in

slender filaments arising dorsally near base of inflated
filament; inflated filament directing forwards, graduallyof S. (M.) yuleae

except following features: Head. Frontal ratio 1.6 : 1.0:

1.7. Frons‑head ratio 1.0 : 4.4. Maxillary palp with 5
segments; proportional lengths of 3rd, 4th and 5th seg‑tapered toward apex, bearing I or 2 slender apicalfilaments (when 2 filaments are present, bifurcation
occurs at different levels in some specimens) ; total

ments 1.0 : 1.0 : 1.9; 3rd segment somewhat enlarged, sen‑length, i.e., inflated portion (up to 0.8‑1.0 mm long fromsory vesicle much elongate, 2.5 x as long as wide,base) plus apical slender filament probably around 1.5

0.56 x as long as 3rd segment, with very large opening
somewhat distally from middle. Maxillary lacinia with
8 or 9 inner and 12 or 13 outer teeth. Mandible with ca.
26 small inner teeth and a few weak outer ones at somemm; inflated tubular filament with several annular
constrictions along its length; surface of tubular fila‑

ment smooth with no patterns, covered with minutetubercles; dorsal 4 slender filaments subequal in length

distance from apex. Cibarium smooth (Fig. 27).

(0.8‑1.0 mm) and thickness to each other, with indis‑

Thorax. Pleural membrane covered with ca. 62 yellow‑
ish white pubescence; a few pubescence also on sclero‑
tized portion just anterior to pleural membrane. Legs.

tinct, annular furrows on their surface, and covered withminute tubercles; all filaments pale yellow. Abdomen.
As in S. (M. ) yuleae except following features: Terga

Fore basitarsus slender, ca. 6.4 x as long as its greatest

7‑9 each with a transverse row of spine‑combs andwidth. Mid tibia yellow on basal l/2 and a little more,brownish black on rest. Hind basitarsus slender, paral‑

comb‑like groups of minute spines directed caudad oneach side, though pine‑combs on tergum 9 much small‑lel‑sided, ca. 6.7 x as long as wide. Hind femur andtibia ca. 2.2 x and 1.7 x as wide as hind basitarsus,

er than those on other terga; tergum 6 with no or a few
spine‑combs; tergum 9 with a pair of wide, plate‑like,

respectively. Wing. Length 2.0 mm. Abdomen. Ter‑terminal hooks which are widely separated from eachother; each terminal hook with moderate serrations
along outer margin (Fig. 39). Cocoon. As in S. (M.)

gites of segments 6‑8 shiny. Genitalia. Sternite 8 withca. 20 dark macrosetae on each side. Anterior gonapo‑
physis triangular, thin, membraneous, covered denselywith microsetae, interspersed with a few short setae.

Male. As in c71 of S. (M.) yuleae except following
features: Head. Wider than thorax. Upper eye consist‑ing of 12 vertical columns and 12 horizontal rows ofyuleae.

Mature larva. Body length 5.3‑5.8 mm. Body greyish.
Cephalic apotome yellow, with distinct, positive headspots. Antenna with 3 segments and apical sensillum,
longer than stem of labral fan; proportional lengths of 3

large facets. Antenna composed of 2 + 9 segments,

segments from base to tip 1.0 : 0.8: 0.7. Labral fan withyellow, but 3 apical segments brown; Ist flagellomere

ca. 40 main rays. Mandible and hypostomium as in S.
(M.) yuleae except hypostomal bristles 10 in number.

somewhat elongate, ca. 1.5 x as long as 2nd one.
Maxillary palp (5th segment lost) with sensory vesicle
oblong, 0.28 x as long as 3rd segment, with opening near

distal end. Thorax. Pleural membrane with ca. 60
yellow pubescence; 8‑10 additional pubescence present
just before it. Legs. Fore basitarsus somewhat dilated,
ca. 6.5 x as long as its greatest width. Hind basitarsus
slender, parallel‑sided, ca. 6.8 x as long as wide. Hind

femur and tibia ca. 2.2 x and 1.8 >< as wide as hind

Postgenal cleft similar to that of S. (M.) yuleae, ca. 2.8

x as long as postgenal bridge. Thoracic cuticle bare.
Abdominal cuticle bare except last segment which ismoderately covered with numerous colorless setae oneach side. Rectal papilla compound, each of 3 Iobes with
3‑5 finger‑1ike, secondary lobules. Anal sclerite of usual

X‑form, with anterior arms ca. 0.7 x as long as poste‑rior ones, broadly sclerotized at base. Ventral papillae

250

present. Posterior circlet with 148 rows of up to 18

basitarsus, respectively. Genitalia. Sternite 8 with ca.

hooklets per row.

27 dark macrosetae on each side. Anterior gonapo‑

TYPE SPECIMENS. Holotype (fl (ANlO , reared fromphysis triangular, thin, membraneous, covered denselywith microsetae, interspersed with a few setae. Genital

pupa, slide mounted, with its pupal skin and cocoon

pinned, collected from Konaiano Creek, Panguna,Bougainville Island, Papua New Guinea, 24. IX. 1988, by
C. Yule. Allotype,(ANlO , reared from pupa, pinned
together with its pupal skin and cocoon, same data as

holotype except date: 3. 111. 1989. Paratypes: 2(ANlO , reared from pupa, pinned, same data as
holotype except date: 16. X. 1988 and 30. 111. 1989; I(BMNH), reared from pupa, slide mounted, with its
pupal skin and cocoon on pin; 9 pupae, 4 pupal skins andfork with a more distinct projection directed forward oneach arm (Fig. 28). Spermatheca ovoid, well sclerotized

except small area near tubal juncture unsclerotized(Fig. 29); internal setae present.

Male. As in c71 of S. (M.) yuleae except following
features: Head. Upper eye consisting of 13 verticalcolumns and 13 horizontal rows of large facets.
Antenna composed of 2 + 9 segments, yellow with 4
apical segments brown; Ist flagellomere somewhat elon‑
gate, ca. 1.7 x as long as 2nd one. Maxillary palp with

1 mature larva (ANIO , same data as holotype except

5 segments; proportional lengths of 3rd, 4th and 5th

date: 31.X.1987; I mature larva (ANlO , collected fromsegments 1.0 : 1.2:2.3; sensory vesicle oblong, 0.29 x as

a stream flowing from Lake Loloru to Mivo River, Mt.

long as 3rd segment, with opening near distal end.

Loloru, Bougainville Island, 8. X. 1988, by C. Yule.

Thorax. Pleural membrane with ca. 70 yellow pubescen‑ce, and a few additional pubescence also present just

DISTRIBUTION. Bougainville Island.

before it. Legs. Fore femur brown. Fore basitarsus
slightly dilated, ca. 7.0 X as long as its greatest width.

REMARKS. The species pe :yae is named after the

Hind tibia yellow on basal 1/2 and a little more, brown‑

wife of Dr. R.W. Crosskey.

ish black on rest. Hind basitarsus slender, parallel‑
sided, ca. 8.4 x as long as wide. Hind femur and tibia
ca. 2.7 x and 2.1 x as wide as hind basitarsus, respec‑tively. Wing. Length 2.0 mm. Other features as inexcept subcosta with a few hairs.
Pupa. Body length (excluding gill filaments) ca. 2.5This species is separable from all the other related

species of the sherwoodi‑group by the female abdomenwith the tergites 2, 6, 7 and 8 shiny (tergite 5 not shiny) ,and the pupal gill composed of 4 slender filaments and 1

inflated filament tapering toward apex and ending as
the one or two slender filaments (Figs. 32‑34). The

mm. Head. Integument dark yellow, densely covered

female and male of this species are also differentiatedfrom those of S. (M.) sherwoodi and S. (M.) , noroense
by the color pattern of the scutal pubescence, as well aswith tubercles of salient polygonal forms (Figs. 36 &the number of pubescence on the pleural membrane.

Antennal sheath moderately covered with cone‑shaped
tubercles. Thorax. Integument dark yellow, densely
covered with cone‑shaped tubercles, intermixed with

6. Simulium (Morops) pangunaense sp. nov.

37) ; a pair of facial trichomes long, simple or bifid, and3 pairs of frontal trichomes all long, simple or bifid.

somewhat larger tubercles of salient polygonal forms; 5

DESCRIPTION. Female. As inof S. (M.) yuleae

anterodorsal pairs of trichomes all long, simple or bifid,

except following features: Head. Frontal ratio 1.9: 1.0 :1 posterolateral pair long and simple. Pit‑like organ

2.0. Frons‑head ratio 1.0:4.4. Maxillary palp with 5absent. Gill (Fig. 35) composed of I inflated, horn‑likefilament and 4 slender, individual filaments, of which 3
are arising dorsally and I is arising inwardly near basesegrnents; proportional lengths of 3rd, 4th and 5th seg‑ments 1.0 : 1.1 : 1.9; 3rd segment somewhat enlarged, sen‑sory vesicle much elongate, 2.5 x as long as wide, 0.57 x

as long as 3rd segment, with very large opening mediallyor slightly distally. Maxillary lacinia with 9 or 10 innerand 12 or 13 outer teeth. Mandible with ca. 27 small
inner teeth and a few weak outer teeth at some distancefrom apex. Legs. Fore basitarsus somewhat dilated, ca.
5.9 x as long as its greatest width. Mid tibia yellow onbasal 1/2, brownish black on rest. Hind basitarsus
slender, parallel‑sided, ca. 7.9 x as long as wide. Hind

femur and tibia ca. 2.4 x and 2.0 X as wide as hindof horn‑like filament, and arising at some distanceanterior to the bases of 3 other slender filaments; horn‑like filament (ca. 0.9 mm long) directing forward anddownward, thick‑walled, thickest at base when viewed
dorsally, gradually tapered toward apex, with no secon‑
dary apical filament; horn‑like filament with several
annular constrictions along its length; surface of horn‑like filament smooth with no patterns, densely covered
with minute tubercles; 4 slender filaments subequal in
thickness to each other, though I innermost filament is251much thicker at base than 3 dorsal filaments, with

7. Simulium(Morops) noroense Takaoka and Suzuki, 1995

indefinite, annular furrows on their surface, and covered

with minute tubercles; 4 slender filaments probably as
long as or a little shorter than the inflated filament; all

Simulium (Morops) noroense Takaoka and Suzuki,
1995: 243‑245.

filaments brown (or pale yellow except basal portion of
inflated filament which is dark yellow in alcoholic speci‑

SPECIMENS EXAMlNED. I pupa, together with itsmens) . Abdomen. As in S. (M.) yuleae except following
features: Terga I and 2 pale yellow, sparsely or moder‑ately covered with minute tubercles, respectively; ter‑gum 9 with a pair of wide, plate‑like, terminal hooks
which are very widely separated from each other; eachterminal hook with weak serrations along outer margin(Fig. 40); several tubercles present around hooks (Fig.
41). Sterna 6 and 7 each with a pair of inner bifid andouter simple spinous seta or hooks (much less developed
than inner hook) on each side. Cocoon. As in S. (M.)

cocoon, and 2 mature larvae (BMNH), in alcohol, col‑lected from a stream near Aropa, 13 Km S of Kieta,

yul eae.

Mature larva. Unknown.
TYPE SPECIMENS. Holotype c71 (ANlO , reared frompupa, slide mounted together with pupal skin andcocoon, collected from Konaiano Creek, Panguna,Bougainville Island, 8. VII. 1965, by R.W. Crosskey.

DISTRIBUTION. Bougainville Island (new record)
and Solomon Islands (New Georgia Island) .

REMARKS. This species was described from New
Georgia Island, Western Province, Solomon Islands,
based on the adult female, male, pupal and larval speci‑

mens (Takaoka and Suzuki, 1995). This species is very
similar to S. (M.) sherwoodi, described from Guadal‑
canal Island (Stone and Maffi, 1971). The pupa of S.
(M.) noroense is distinguished from that of the latter
species by the thoracic integument moderately covered
with tubercles (the thoracic integument is usually bare

Bougainville Island, Papua New Guinea, 28. 111. 1989, byor sparsely covered with tubercles, especially on the

C. Yule. A110type(ANlO , reared from pupa, pinhed,same data as holotype except date: 29. X. 1988. Para‑
types: I c71 (BMNH), reared from pupa, pinned, samedata as allotype, I(BMNH), reared from pupa, slide

dorsomedial portion in the latter species) .

mounted, with its pupal skin and cocoon in alcohol, samedata as holotype except date: 28. II. 1989, 2 pupae

I am grateful to Dr. R.W. Crosskey, Natural His‑
tory Museum, London, U.K., for forwarding material(ANlO , same as holotype except date: 28. II. 1989.

collected by him and by Miss C. Yule. My appreciation

DISTRIBUTION. Bougainville Island.

goes to Dr. A. Shelley, Ms C. Lowry and Miss T.
Howard, Department of Entomology, Natural History

REMARKS. The species pangunaense refers to the
locality where this species was collected.

This new species seems to be very closely related toS. (M.) kerei, described from the Solomon Islands(Takaoka and Suzuki, 1994) in having the similar

ACKNOWLEDGEMENTS

Museum, London, U.K., who kindly arranged the loan of
the type specimens of several Papua New Guinean andAustralian species of the clathrinum‑gfoup. Thanks are
due to Miss C. Aoki, Division of Medical Zoology, Oita
Medical University, for her help.

arrangement of the pupal gill filaments and the similar

REFERENCES

form of the pupal terminal hooks. However, S. (M. )pangunaense is separable in the pupa from the latter by
its frontal and thoracic integuments which are densely
covered with tubercles. Moreover, in S. (M.) kerei, four

slender filaments are subequal in thickness to eachother, but, in S. (M. ) pangunaense, the innermost fila‑

ment is basally much thicker than the three dorsal
filaments. This new species is readily separated fromthe other species of the sherwoodi‑group by the arrange‑
ment of the pupal gill filaments (Fig. 35).

1 ) Colbo, M. H. (1976): Four new species of Simulium
Latreille (Diptera: Simuliidae) from Australia. J. Aust.
Ent. Soc., 15, 253‑269.

2 ) Crosskey, R. W. (1967): The classification of Simulium

Latreille (Diptera: Simuliidae) from Australia, New
Guinea and the Western Pacific, J. Nat. Hist. 1, 23‑51.

3 ) Crosskey. R. W. (1989): Family SIMULIIDAE. In:
Evenhuis, N. L. (ed) , Catalog of the Diptera of theAustralasian and Oceanian Regions, p. 221‑225. Bishop
Museum Special Publication 86, Bishop Museum Press& E. J. Brill.

4 ) Mackerras, M. J. and Mackerras, I. M. (1948):

252

Simuliidae (Diptera) from Queensland, Aust. J. Scient.
Res., Ser. B, 1, 231‑270.

5) Smart, J. and Clifford, E. A. (1965) : Simuliidae (Dipter‑a) of the territory of Papua and New Guinea, PacificInsects, 7, 505‑619.

6) Stone, A. and Maffi, M. (1971): A new species ofSimulium from Guadalcanal, Solomon Islands (Diptera:
Simuliidae) , J. Med. Entomol., 3, 299‑300.

7) Takaoka, H. (1994): A new blackfly species ofSimulium ( Gomphostilbia) from Solomon Islands, South
Pacific (Diptera: Simuliidae) , Jpn. J. Trop. Med. Hyg.,22, 103‑108.

8 ) Takaoka, H. and Davies, D. M. (1995): The black flies(Diptera: Simuliidae) of Java, Indonesia, Bishop
Museum Entomology Monograph, in press.
9 ) Takaoka, H. and Suzuki, H. (1994) : A new blackfly
species of Simulium (Morops) from Solomon Islands,
South Pacific (Diptera: Simuliidae) , Jpn. J. Trop. Med.
Hyg., 22, 207‑210.

10) Takaoka, H. and Suzuki, H. (1995) : The Simuliidae(Diptera) from the Solomon Islands, South Pacific, Jpn.
J. Trop. Med. Hyg., 23, 227‑246.

253

Jpn. J. Tro p. Med. Hyg., Vol. 23, No. 4 (1995) pp. 253‑272

THE SIMULIIDAE (DIPTERA) FROM THE SOLOMONISLANDS, SOUTH PACIFICHIROYUKI TAKAOKA

AND HIROSHI SUZUKI'

Received Septernber 14, 1995/Accepted October 15, 1995

Abstract: Nine black fly species including five new species were taxonomically studied, chiefly on the basisof reared adults, pupae and mature larvae collected from the Solomon Islands, South Pacific. All species, but

one in Simulium (Gbmphostilbia) , were placed in Simulium (Morops). Within the subgenus Morops four
were further classified into the clathrinum‑group, one into the papuense‑group and three remained un‑grouped. S. (M.) papuense was newly recorded from the Solomon Islands. Descriptions of all new species
and redescription of S. (M.) sherwoodi were given; mature larvae of S. (M.) sherwoodi and S. (M.) kerei
were described for the first time. The pupal pit‑like organs of all five known species of the clathrinum‑group from Papua New Guinea and Australia were also illustrated.

Black flies (Diptera: Simuliidae) of the Solomon
Islands, South Pacific were little studied. Stone andMaffi (1971) described Simulium (Morops) sherwoodi,

species, S. (G.) hiroshii, was known (Takaoka, 1994) . It

is noteworthy that this species is very unique in having

several characters which depart from the diagnosis ofand recorded the other black fly species (as S. (M.) sp.

Gomphostilbia but agree with those of Morops or

nr avilae) from Guadalcanal Island. Recently we de‑scribed two new species: S. ( Gomphostilbia) hiroshii
from New Georgia and Guadalcanal Islands, and S. (M.)
kerei from New Georgia Island (Takaoka, 1994; Takao‑

Simulium s. str., as mentioned by Takaoka (1994).

ka and Suzuki, 1994).

Simulium ( Gomphostilbia) hiroshii Takaoka, 1994: 97‑

In this study nine species including five new and one

newly recorded species were treated on the basis ofreared adults, pupae and mature larvae collected from
Guadalcanal and New Georgia Islands in 1992 and 1993
by one of us (HS).

1. Simulium ( Gomphostilbia) hiroshii Takaoka, 1994

101.

SPECIMENS EXAMlNED. I pupa, I pupal exuvia and22 mature larvae (all in alcohoD, collected from astream, Noro, New Georgia Island, Western Province,

Descriptions of five new species and redescriptionof S. (M.) sherwoodi are given, and mature larva of S.
(M.) sherwoodi and S. (M. ) kerei is described for theSolomon Islands, 21. II. 1993, by H. Suzuki; I pupa and1 mature larva, tollected from a stream, Selwyn College,

first time.

H. Suzuki.

Guadalcanal Island, Solomon Islands, 12. VIII. 1993, by

Type specimens will be deposited in the Natural

History Museum (BMNH) , London, U.K.

DISTRIBUTION. Solomon Islands (New Georgia
Island and Guadalcanal Island) .

Genus Simulium Latreille s. l.
Subgenus Gomphostilbia Enderlein
The subgenus Gomphostilbia was well defined by
Crosskey (1967) and recently revised by Takaoka andDavies (1995). This subgenus is characterized by the

REMARKS. This species was originally reported fromthe Solomon Islands (Takaoka, 1994). The pupa andlarva were collected for the first time from the Guadal‑

canal Island where only the adult female and male had
been previously known.

bare pleural membrane and haired katepisternum ofadults of both sexes. From the Solomon Islands only one
1 Division of Medical Zoology, Oita Medical University, Hasama, Oita 879‑55, Japan
2 Department of Virology, Institute of Tropical Medicine, Nagasaki University, Sakamoto, Nagasaki 852 Japan

254

Subgenus Morops Enderlein
This subgenus was well defined by Crosskey (1967)
and was divided into several species‑groups (Crosskey,
1967; Colbo, 1976). The subgenus Morops is character‑ized by the haired pleural membrane and katepisternumof adults of both sexes. In this study, all the nine, but

one in subgenus Gomphostilbia Enderlein, are assigned inthis subgenus; four species are further classified into theclathrinum‑group and one is into the papuense‑groupand three remain ungrouped.

(A) clathrinum‑group
This species‑group is easily distinguished by having

the pit‑like cuticular organ at the base of pupal gill
filaments (Crosskey, 1967) . To express relative size of

Cibarium (Fig. 3) with upper margin widely bent for‑wardly, Iacking any processes. Thorax. Scutum darkbrown in ground color, densely covered with yellowish
white, recumbent pubescence except median, Iarge areawith dark brown pubescence; 3 Iongitudinal vittae ofyellowish white, recumbent pubescence (1 median vitta
narrow and 2 submedian ones rather wide) , which areconnected anteriorly and posteriorly to areas of theconcolored pubescence; Iong, erect, dark hairs present
on prescutellar area. Scutellum blackish brown withyellowish white pubescence and many long, erect, darkhairs. Postscutellum blackish brown, bare. Pleural mem‑brane with ca. 120 yellowish white pubescence and a few

dark hairs. Katepisternum longer than deep, dark

the clathrinum‑group are illustrated and measured forbrown, with numerous, yellowish white pubescence anddark hairs; sulcus distinct. Legs. Foreleg: coxa andtrochanter pale yellow; femur pale yellow with apicalcap brown; tibia pale yellow with basal and distal 114
brown; tarsus brown; basitarsus somewhat dilated, ca.
5.8 x as long as its greatest width. Midleg: coxa darkbrown; trochanter pale yellow; femur pale yellow withapical cap brown; tibia brownish with median large

plo ratio.

portion pale; tarsus brown. Hind leg: coxa pale yellow‑

the pit‑like organ needed for species identification, pit‑

like organ ratio (=A/B), abbreviated as plo ratio, ishere proposed, where A is a diameter of the pit‑likeorgan and B is a distance from its innermost margin to
the middle longitudinal suture of the thorax (see Fig.
12) . The pit‑like organs of all the five known species ofStone and Maffi (1971) recorded S. (M.) sp. nr
avilae Smart & Clifford, on the basis of the pupal andlarval specimens from Guadalcanal Island, which was
the only species of the clathrinum ‑group from theSolomon Islands. This unidentified species may be S.
(M.) pohaense sp. nov. or S. (M.) selwynense sp. nov.,
both of which are very similar to S. (M. ) avilae, as noted
later.

ish brown; trochanter pale yellow; femur pale yellow
with pical cap brown; tibia brownish with base andmedian large portion pale yellow; tarsus pale brownwith basal 3 / 5 of basitarsus and basal I / 2 of 2nd
tarsomere white; basitarsus slender, parallel‑sided, ca.
6.7 x as long as wide. Calcipala (Fig. 4a) well devel‑
oped, as long as wide. Pedisulcus (Fig. 4b) well devel‑

Frons dark brown, densely covered with yellowish

oped. Hind femur and tibia 2.1 x and 1.6 x as wide as
hind basitarsus, respectively. All femora and tibiae
densely covered with scale‑like hairs on outer surface.
Claws (Fig. 5) each with a small basal tooth. Wing.
Length 1.7 mm. Costa with spinules as well as hairs.
Subcosta haired. Hair tuft on stem vein dark brown.
Basal portion of radius fully haired. Abdomen. Basal

white, scale‑like, recumbent pubescence, interspersedwith several dark hairs along both lateral margins;scale brown with fringe of yellowish white hairs. Dorsal
surface of segments 2‑5 very densely covered with scale‑

frontal ratio 1.57: 1.00 : 1.87. Frons‑head ratio 1.00 : 4.75.

2. Simulium (Morops) solomonense sp. nov.
DESCRIPTION. Female (coloration incomplete) . Body
length ca. 2.5 mm. Head. Slightly narrower than thorax.

3rd, 4th and 5th segments 1.0:1.0:1.7; 3rd segment some‑like, recumbent pubescence, which is yellowish white
on segment 2 but brown dorsally and yellowish whitelaterally on segments 3‑5; that of segments 6 and 7
moderately covered with yellowish white, recumbent
pubescence and dark hairs; that of segments 8 and 9
moderately covered with yellowish white, short hairs
and dark, Ionger hairs; tergite of 2nd segment whitish

what enlarged, sensory vesicle (Fig. 2) ellipsoidal, 1.6 x

pruinose, tergites of segments 6‑8 shiny. Genitalia. Ster‑as long as wide, 0.3 x as long as 3rd segrnent, with a

nal plate undeveloped on abdominal segment 7. Sternite

moderate opening near distal end. Maxillary lacinia

8 (Fig. 6a) bare medially, with 32‑36 macrosetae on

with 7 or 8 inner and 10 outer teeth. Mandible with ca.
20 small inner teeth and I or 2 outer teeth near apex.

each side. Anterior gonapophysis (Fig. 6b) triangular,

Fronto‑ocular area (Fig. D well developed. Clypeus
dark brown, densely covered with yellowish white, scale‑like, recumbent pubescence interspersed with several
dark hairs. Antenna composed of 2 + 9 egments, pale.
Maxillary palp with 5 segments; proportional lengths ofthin, membraneous, covered densely with microsetae

255

interspersed with a few setae; inner margin well sclero‑
tized. Genital fork (Fig. 6c) of usual inverted‑Y form,
with arms rather broad, folded medially, without projec‑

Male. Body length ca. 2.8 mm. Head. Wider than thorax.

Upper eye consisting of 14 vertical columns and 15
horizontal rows of large facets. Clypeus brownish black,tion directed forward; arms furnished with microsetae
posteriorly. Paraproct (Figs. 6d & 7a) of usual form.
Cercus (Figs. 6e & 7b ) 0.5 x as long as wide, withposterior border nearly straight when viewed laterally.
Spermatheca ovoid, well sclerotized and with reticulate
patterns on surface; tube and small area near tubal

covered densely with yellow pubescence, interspersedwith several dark hairs. Antenna composed of 2 + 9juncture unsclerotized; internal setae absent.

ment, with a small opening. Thorax. As insegments, yellowish; Ist flagellomere elongate, 2.0 X aslong as 2nd one. Maxillary palp with 5 segments;
proportional lengths of 3rd, 4th and 5th segments 1.0:1.1:

2.0; sensory vesicle globular, 0.2 x as long as 3rd seg‑i&": :
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',t,aEfl I *L 11
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Figs. 1‑11.

11

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L1 .ili' 'F;: 4f4 """ "'

10

,

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b 4

Adult characters of Simulium (Morops) solomonense sp. nov. 1, fronto‑ocular area; 2, 3rd maxillarypalpal segment showing a sensory vesicle; 3, cibarium; 4, distal tip of basitarsus and 2nd tarsal
segment of hind leg showing calcipala (a) and pedisulcus (b) ; 5, claw with a small tooth; 6, genitalia
in situ (ventral view) (a, 8th sternite; b, gonapophysis; c, genital fork; d, paraproct; e, cercus; f,spermatheca); 7, paraproct (a) and cercus (b) (side view) ; 8, coxite (a) and style (b) (ventralview) ; 9‑11, ventral plate (9, ventral view; 10, side view; 11, end view) . 1‑7,

; 8‑11, c?1.

256

pleural membrane with ca. 130 yellowish white pubes‑

combs on tergum 9 somewhat smaller than those on

cence interspersed with several dark hairs. Legs.

terga 7 and 8. Tergum 9 with distinct, cone‑shaped,
terminal hooks (Fig. 14). Sternum 4 with I distinct,

Coloration as in. Fore basitarsus somewhat dilated,

6.3 x as long as its greatest width. Hind basitarsus
slender, almost parallel‑sided, 6.5 x as long as wide.
Hind femur and tibia ca. 2.3 x and 1.8 x width of hindbasitarsus, respectively. Calcipala and pedisulcus well

simple hook and a few minute setae on each side; ster‑num 5 with a pair of bifid hooks on each side; sterna 6
and 7 each with a pair of inner bifid and outer simple
hooks on each side; Iast segment without grapnel‑likedeveloped, as in. All femora and tibiae densely

hooklets ventrolaterally on each side. Cocoon. Shoe‑covered with scale‑like hairs on outer surface. Wing.
Length 1.8 mm. Other features as in except subcostabare. Abdomen. Basal scale brown, with fringe of darkhairs. Dorsal surface of abdominal segments brownish,covered with dark hairs; segments 2, 5‑7 each with adorsolateral pair of shiny patches. Genitalia. Coxite

shaped, moderately woven, somewhat extending(Fig. 8a) much longer than wide. Style (Fig. 8b) shorter

than coxite, tapered toward apex, and with an apicalventrolaterally; anterior margin not thickly woven; indi‑vidual threads visible; interspaces thinnly walled, but
without perforations.

Mature larva. Body length 4.7‑5.2 mm. Body grey togreyish black. Cephalic apotome pale yellow with alarge dark brown area medially on posterior 1/2; headspots distinct, often merged in surrounding dark brownspine. Ventral plate (Figs. 9‑1D transverse, much short‑
er than wide, concave posteriorly (in ventral view) , fully

area. Antenna with 3 segments and apical sensillum,

setose on ventral surface but bare on posterior andsegments from base to tip 1.0 : 1.0 : 0.7. Labral fan withdorsal surfaces; basal arms of moderate length, subpar‑

ca. 44 main rays. Mandible (Fig. 15) with comb‑teeth

longer than stem of labral fan; proportional lengths of 3

allel to each other. Paramere without hooks. Mediandecreasing in size from Ist to 3rd; mandibular serrationsclerite tansparent, probably wide.
Pupa. Body length (excluding gill filaments) ca. 2.8 mm.

tubercles; near base of gill there is on each side a large,

composed of 2 teeth (1 Iarge and I smalD, with a few
supernumerary serrations. Hypostomium (Fig. 16) witha row of 9 apical teeth; median tooth as long as eachcorner tooth, Ionger than 3 intermediate teeth on eachside; Iateral margin serrate; hypostomal bristles 7 innumber per side, Iying slightly divergent posteriorly
from lateral margin. Postgenal cleft (Fig. 16) withcircular pit‑Iike organ (Fig. 12a) thickly fringed withlateral margins subparallel to each other on basal 1/2,

distinct, comb‑li e processes (as in Fig. 20) ; plo ratio 1;

with apex pointed, which often has a narrow extension

Head. Integument dark yellow, bare with 4 pairs oftrichomes, all long and simple. Antennal sheath withsmooth surface. Thorax. Integument dark yellow, bare
except posterodorsal surface very sparsely covered withthoracic trichomes 6 pairs (5 anterodorsally and I pos‑
terolaterally) , all long and simple. Gill (Fig. 13) com‑

tomium. Thoracic cuticle bare. Abdominal cuticle bareanteriorly approaching posterior border of hypos‑

posed of 4 slender filaments, arising from very longexcept each side of anal sclerite of last segment moder‑basal stalk, subequal in length and thickness (total gilllength ca. 2.0 mm) ; basal stalk with a large, transparent

papilla compound, each of 3 Iobes with 4 or 5 finger‑like,pouch, which expands laterally and ventrally (Figs.
12b & 13a); surface of distal 1/2 of basal stalk withmany longitudinal and transverse ridges, covered withsomewhat larger tubercles on ridges and with smaller

secondary lobules on posterior surface. Anal sclerite(Fig. 17) of usual X‑form, with anterior arms ca. 0.7 x
as long as posterior ones, broadly sclerotized at base.
Last abdominal segment expanded ventrolaterally form‑

ones on interspaces (Fig. 13b) ; slerider filaments sub‑ing double bulges on each side, visible as a small ventral

equal in length and thickness, with well‑defined annular

papilla when viewed from side. Posterior circlet with ca.

ridges and furrows on their surface, and covered withrather larger tubercles on ridges and smaller ones oninterspaces; all filaments dark yellow to yellowish

l04 rows of up to 16 hooklets per row.

TYPE SPECIMENS. Holotype

brown. Abdomen. Terga I and 2 pale brown, without

slide‑mounted, collected from a stream, Noro, New

tubercles; tergum I with I Iong seta on each side, tergum

Georgia Island, Western Province, Solomon Islands, 5.

ately covered with numerous colorless setae. Rectal, reared from pupa,

2 with 5 simple spinous setae and I much longer seta on

IX. 1992, by H. Suzuki. Allotype c71 , reared from pupa,

each side. Terga 3 and 4 each with 4 hooked spines
directed forward on each side. Terga 7‑9 each with atransverse row of spine‑combs and comb‑like groups ofslide‑mounted, same data as holotype. Paratypes: 5
pupae and 4 pupal exuviae, same data as holotype; 1pupa and 2 pupal exuviae and 1 1 mature larvae, same

minute spines directed caudad on each side, though spine‑

data as holotype except date: 21. II. 1993.

257

DISTRIBUTION. Solomon Islands (New Georgia

ratio ca. 0.7). However, there are clear differences

Island) .

between this new species and the latter two known
species. In the latter two species, the basal stalk is

REMARKS. This new species belongs to the clathrinum‑

short, and there are tubercles along the middle longitudi‑group of the subgenus Morops, defined by Crosskey

nal suture of the anterodorsal surface of thorax (Figs.
18 & 19).

(1967) by having the pit‑like organ (Fig. 12a) , as well asfour gill filaments. Among five known species, S. (M. )clathrinum from Australia (Mackerras and Mackerras,
1948) and S. (M.) Ialokiense from central Papua New

3. Simulium (Morops) pohaense sp. nov.

Guinea (Smart and Clifford, 1965) have a large, pit‑likeDESCRIPTION. Female. Body length ca. 2.5 mm. Head.
Slightly narrower than thorax. Frons dark brown,organ with comb‑1ike processes along the inner margin(Figs. 18 & 19). The pit‑like organ of this species is aslarge as that of S. (M.) Ialokiense (plo ratio ca. 1.D but

white pruinose, densely covered with white, scale‑like,somewhat larger than that of S. (M.) clathrinum (plohairs along both lateral margins; frontal ratio 1.51:1.00:

recumbent pubescence, interspersed with several dark

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Figs. 12‑17. Pupal and larval characters of Simulium (Morops) solomonense sp. nov. 12‑14, pupa; 15‑17, Iarva.

12, anterior part of thorax (right half) showing a pit‑like organ (a) and a transparent pouch (b)at the base of gill filaments (dorsal view) ; 13, gill filaments showing a transparent pouch (a) andsurface sculpture on the basal stalk (b) ; 14, terminal hooks; 15, apical tip of mandible; 16, headcapsule showing hypostomium and postgenal cleft; 17, anal sclerite. Scale bar 0.02 mm for Fig. 13b.

258

1.54. Frons‑head ratio 1.0: 4.3. Fronto‑ocular area (Fig.

22) well developed. Clypeus dark brown, densely cov‑
ered with white, scale‑like, recumbent pubescence inter‑

spersed with several dark hairs. Antenna composed of
2 + 9 segments, dark brown except scape, pedicel andbase of Ist flagellomere yellow when viewd from top.
Maxillary palp with 5 segments; proportional lengths of3rd, 4th and 5th segments 1.0:1.1:2.1; 3rd segment
somewhat enlarged, sensory vesicle (Fig. 23) ellipsoidal,

1.7 x as long as wide, 0.3x as long as 3rd segment, witha medium opening near distal end. Maxillary laciniawith 7 inner and 11 outer teeth. Mandible with 21 small
inner teeth and I outer tooth near apex; outer marginnear apex very slightly erose (Fig. 25). Cibarium (Fig.
26) with smooth posterior margin, which is covered withtransparent, elongated projections extending from botharms. Thorax. Scutum dark brown in ground color,
densely covered with white, recumbent pubescence
except median, Iarge area with dark brown pubescence;
3 Iongitudinal vittae of white recumbent pubescence (lmedian vitta narrow and 2 submedian ones rather wide) ,which are connected anteriorly and posteriorly to areas' 'L.
'lbb IL

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Figs. 18‑21. Pupal characters of two species of the clathrinum‑group. 18 & 19, anterior part of thorax showing
a pit‑like organ (18, dorsal view; 19, side view, slide‑mounted) ; 20, comb‑like processes along inner

margin of pit‑like organ; 21, gill filaments. 18. S. (M.) clathrinum (alcoholic specimen collectedfrom North Queensland, Australia, 15. IX. 1949, by I. M. & M. J. Mackerras, BMNH) ; 19‑21, S.
(M.) Ialokiense (allotype slide‑mounted specimen collected from Central District, Papua New
Guinea, 17. IX. 1957, by J. Smart, BMNH). Scale bar 0.02 mm for Fig. 20.

259

of the concolored pubescence; Iong, erect, dark hairspresent on prescutellar area. Scutellum blackish brownwith white pubescence and many, Iong, erect, dark hairs.

rior surface silvery iridescent in certain angles of light;

tarsus dark brown with basal 3/5 of basitarsus and basal
1/2 of 2nd tarsomere white; basitarsus slender, parallel‑

Postscutellum blackish brown, bare. Pleural membranewith ca. 120 white pubescence and a few dark hairs.

sided, ca. 7.2 x as long as wide. Calcipala (Fig. 27) wellKatepisternum longer than deep, dark brown, withwell developed. Hind femur and tibia 2.3 x and 1.7 >< asnumerous white pubescence and dark hairs; sulcus dis‑tinct. Legs. Foreleg: coxa and trochanter pale greyish

wide as hind basitarsus, respectively. All femora andtibiae densely covered with scale‑like hairs on outerbrown; femur pale brown; tibia brown with median,large portion somewhat lighter, white pubescence on

surface. Claws each with a small basal tooth, similar toouter surface silvery iridescent in certain angles of light;

Costa with spinules as well as hairs. Subcosta haired on

tarsus blackish brown; basitarsus dilated, ca. 5.3x as

basal 3/5. Hair tuft on stem vein dark brown. Basal
portion of radius fully haired. Abdomen. Basal scale
brown with fringe of white hairs. Dorsal surface ofsegments 2‑5 pale except tergites brown, very densely

long as its greatest width; tarsal segments 1‑3 moderate‑

ly furnished with dorsal crest of dark brown, scale‑likehairs. Midleg: coxa dark brown; trochanter pale greyish

brown; femur greyish brown with apical cap brown;
tibia brown with base yellow, white pubescence onposterior surface silvery iridescent in certain angles oflight; tarsus brown to blackish brown. Hind leg: coxa
and trochanter pale greyish brown; femur greyish brownwith apical cap brown; tibia brownish with base yellow
and apical 1/4 dark brown, white pubescence on poste‑

developed, ca. 1.2 x as long as wide. Pedisulcus (Fig. 27)

that ofS. (M.) solomonense. Wing. Length 1.7 mm.

covered with scale‑like, recumbent pubescence which isentirely shiny white on segment 2, and mostly darkbrown (intermixed with white ones along posterior
margin) on segments 3‑5; that of segments 6‑8 eachwith a large, dark brown tergite, moderately covered
with white, recumbent pubescence, interspersed withdark hairs; tergites 2, 6‑8 shiny; abdomen moderately

.a.

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26

27

29

Figs. 22‑29. Adult characters of Simulium (Morops) pohaense sp. nov. 22, fronto‑ocular area; 23 & 24, 3rd

maxillary palpal segment showing sensory vesicle; 25, mandible; 26, cibarium; 27, calcipala andpedisulcus; 28, genitalia in situ (ventral view) ; 29, paraproct and cercus (side view) . 24, c71 ; others,

.

260

Male. Body length ca. 2.5 mm. Head. Wider thancovered with white, scale‑like pubescence also laterally.
Genitalia (Figs. 28 & 29). Sternal plate well developed

thorax. Upper eye consisting of 14 or 15 vertical col‑umns and 13‑15 horizontal rows of large facets. Clypeus

on abdominal segment 7 and with ca. 50 hairs. Sternite
8 bare medially, with 24‑26 macrosetae on each side.
Anterior gonapophysis triangular, thin, membraneous,brownish black, covered densely with whitish yellow
pubescence, intermixed with many dark hairs. Antenna

covered densely with microsetae interspersed with a few
setae; inner margin moderately sclerotized. Genital forkof usual inverted‑Y form, with arms rather broad, fold‑ed medially, with a short projection directed forward;
arms furnished with microsetae posteriorly. Paraproct

composed of 2 + 9 segments, pale brown, becoming dark‑of usual form. Cercus 0.5 x as long as wide, with

2.00; sensory vesicle (Fig. 24) globular, 0.2 x as long as

er apically except scape, pedicel and base of Ist flagel‑lomere yellow; Ist flagellomere elongate, 1.7 x as longas 2nd one. Maxillary palp with 5 segments; propor‑
tional lengths of 3rd, 4th and 5th segments 1.00 : 1.03 :3rd segment, with a medium opening. Thorax. As in

posterior border rounded when viewed laterally. Sper‑matheca nearly globular, well sclerotized, with reticu‑

except pubescence on scutum and scutellum yellowish; 3
longitudinal vittae of yellow pubescence on scutum notwell defined. Legs. As in
except following features:late patterns on surface; tube and small area near tubal
juncture unsclerotized; internal setae absent.

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32

31

Figs. 30‑34. Pupal characters of Simulium (Morops) pohaense sp. nov. 30, frons (right half) showing 4 trichomes

and scattered tubercles; 31, anterior part of thorax (right half) showing a pit‑like organ, tubercles
along middle longitudinal suture, five trichomes and basal portion of gill (dorsal view) ; 32, pit‑likeorgan showing minute, spine‑like processes on floor; 33, dorsal eaves‑like wall of pit‑like organ
showing minute, spine‑like processes on under surface and along inner margin; 34, gill filaments(side view) (a, interspiracular trunk) .

261

fore basitarsus somewhat dilated, 6.3 x as long as its
sided, 7.0 X as long as wide; hind femur and tibia ca.
2.1 x and 1.8 x width of hind basitarsus, respectively.

ment dark yellow, bare or very sparsely covered withround tubercles except posterodorsal surface moderate‑
ly covered with smaller tubercles and both sides of
middle longitudinal suture narrowly covered with roundWing. Length 1.6 mm. Other features as inexcepttubercles (Fig. 31) ; near base of gill there is on each side

subcosta bare. Abdomen and Genitalia. As in c7, of S.
(M. ) solomonense.

a medium‑sized, pit‑like organ sparsely covered withminute, spine‑like processes on floor (Fig. 32) and onunder surface and along inner margin of dorsal, eaves‑
like wall (Fig. 33); plo ratio variable from 0.31‑0.53(median = 0.43, n = 83); thoracic trichomes 6 pairs (5anterodorsally and I posterolaterally) , all long andsimple. Gill (Fig. 34) composed of 4 slender filaments,

greatest width; hind basitarsus slender, almost parallel‑

Pupa. Body length (excluding gill filaments) ca. 2.5 mm.

Head. Integument dark yellow, bare or sparsely (moder‑ately in some pupae. Fig. 30) covered with round tuber‑cles, and with 4 pairs of trichomes, all long and simple.

Antennal sheath with smooth surface. Thorax. Integu‑r'
Oo ,pppoo o IIOGSO I

pt

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Figs. 35‑38. Pupal characters of Simulium (Morops) avilae. 35, frons (right half) showing four trichomes andscattered tubercles; 36, anterior part of thorax showing a pit‑like organ, tubercles along middle
longitudinal suture, five trichomes and basal portion of gill (side view, slide‑mounted) ; 37, pit‑likeorgan showing minute, spine‑like processes on under surface and along inner margin; 38, gill
filaments (side view) (a, interspiracular trunk). (Examined was the paratype slide‑mounted
specimen collected in Northern District, Papua New Guinea, 19. I . 1958, by J. Smart, BMNH)

262

arising from long basal stalk, sessile or short‑stalked,

rinum‑group by having the pit‑like organ (Fig. 31) , assubequal in length and thickness (total gill length 1.5‑

well as four gill filaments. The pit‑like organ of this

2.3 mm, average ca. 1.8 mm); basal stalk with a large,transparent pouch, which expands laterally; each fila‑
ment rather thick basally, nearly as thick as interspir‑

species differs in size from one pupa to another, then plo

ratio also varies from 0.31 to 0.53 with a median of 0.43.

acular trunk (Fig. 34a) , tapered gradually to mid point,

Among five known species of the clathrinum‑group, S.
(M.) avilae from Northern District, Papua New Guinea

then nearly of the same diameter up to apical tip; sur‑face of each filament with well‑defined, annular ridgesof which plo ratio is 0.45 (measured from two slide‑forming polygonal patterns, covered with somewhat
larger tubercles on ridges and with smaller ones oninterspaces; all filaments dark yellow to yellowish
brown. Abdomen and Cocoon. As in S. (M.) solomonense
except tergum 9 Iacking spine‑combs.
Mature larva. As in S. (M.) solomonense except follow‑
ing features: Body length 4.3‑5.0 mm. Cephalic apotome
pale yellow with a large, dark brown area medially on
posterior 1/2 with head spots distinct, often merged in(Smart and Clifford, 1965) has a similar pit‑like organ

mounted specimens loaned from BMNH) . However, S.
(M.) pohaense is separated from S. (M.) avilae by
having the somewhat inflated base of each gill filament(Fig. 34) which is as thick as interspiracular trunk(much thinner in the latter species, see Fig. 38) , and by

the presence of spinous processes on the floor of the pit‑

like organ (Fig. 32) (absent in the latter species
although there are several processes on the dorsal wall,as seen in Fig. 37) .

surrounding dark brown area, or cephalic apotome
entirely pale yellow with positive head spots, at least

4. Simulium (Morops) selwynense sp. nov.

middle longitudinal spot. Proportional lengths of 3antennal segments from base to tip 1.0 : 1.0 : 0.6. Labral

DESCRIPTION. Female. Unknown.

fan with 36‑42 main rays. Hypostomal bristles 6 innumber per side. Rectal papilla compound, each of 3
lobes with 3 or 4 finger‑Iike, secondary lobules on

Male. As in c l of S. (M.) pohaense except following
features: Body length ca. 2.2 mm. Head. Wider thanthorax. Upper eye consisting of 12 vertical columns andposterior surface. Posterior circlet with 92‑100 rows of

13 horizontal rows of large facets. Sensory vesicle

up to 15 hooklets per row.

nearly globular, 0.21 x as long as 3rd segment, with asmall opening medially. Thorex. Pleural membrane

TYPE SPECIMENS. Holotype, reared from pupa,

slide‑mounted, collected from Poha River, GuadalcanalIsland, Solomon Islands, 15. VIII. 1993, by H. Suzuki.
Allotype c , reared from pupa, slide‑mounted, samewith ca. 100 yellowish white pubescence interspersedwith a few dark hairs. Legs. Fore basitarsus somewhat
dilated, 5.8 x as long as its greatest width. Hind basitar‑sus slender, almost parallel‑sided, 7.0 X as long as wide.

alcohol except 2 pupal exuviae and 3 mature larvae slide‑

Hind femur and tibia ca. 2.1 x and 1.7 >< width of hindbasitarsus, respectively. Wing. Length 1.5 mm.
Pupa. Body length (excluding gill filaments) 2.5‑2.8

mounted, same data as holotype. Other specimens: 5,

mm. Head. Integument dark yellow, densely covered

7 crl , all reared from pupa, 15 pupae, 33 pupal exuviae

with round tubercles, and with 4 pairs of trichomes, all

and 17 mature larvae, collected from Tenaru River,
all reared from pupa, 18 pupae and 13 pupal exuviae,
collected from Tenaru River, Guadalcanal Island, 16.

long and simple (Fig. 39) . Antennal sheath with smoothsurface. Thorax. Integument dark yellow, densely cov‑
ered with round tubercles except anterior surface bareor sparsely covered with tubercles (Fig. 40) ; near base

VIII. 1992, by H. Suzuki; 26, all reared fromof gill there is on each side a medium‑sized, pit‑1ike

pupa, 49 pupae, 13 pupal exuviae and 3 mature larvae,
collected from Tanatita River, Guadalcanal Island, 16.
VIII. 1992, by H. Suzuki; I , reared from pupa, and 4
pupae, collected at Gold Ridge, Guadalcanal Island, 30.

organ (Fig. 40), similar to that of S. (M.) pohoense; plo
ratio 0.45; thoracic trichomes 6 pairs (5 anterodorsallyand I posterolaterally) , all long and simple. Gill (Fig.

data as holotype. Paratypes: 2, 5 c71, all reared frompupa, 23 pupae, 10 pupal exuviae, 5 mature larvae, all in

Guadalcanal Island, 10. VIII. 1992, by H. Suzuki; 8, 13 c, 3 c !,

VIII. 1992, by H. Suzuki.

DISTRIBUTION. Solomon Islands (GuadalcanalIsland) .

41) nearly as in S. (M.) pohaense. Abdomen and Cocoon.
As in S. (M. ) solomonense except terga 6‑8 each with atransverse row of spine‑combs though I or 2 in number

on tergum 6.
Mature larva. As in S. (M.) solomonense except follow‑

ing features: Body length 4.0‑4.2 mm. Proportional
REMARKS. This new species also belongs to the clath‑lengths of 3 antennal segments from base to tip 1.0 : 1.1 :263

0.7. Labral fan with ca. 37 main rays. Hypostomal

Suzuki.

bristles 6 in number per side. Rectal papilla compound,each of 3 Iobes with 3 or 4 finger‑like, secondary lobules

DISTRIBUTION. Solomon

on posterior surface. Posterior circlet with 85‑88 rowsof up to 14 hooklets per row.

Island) .

Islands(Guadalcanal

REMARKS. This new species resembles S. (M. ) pohaen‑

TYPE SPECIMENS. Holotype c ,, reared from pupa,
slide‑mounted, collected from a stream at Selwyn Col‑lege, Guadalcanal Island. Solomon Islands, 12. VIII.

se in many features including the pit‑like organ and thegill filaments of the pupa. However this species differsfrom the latter species by the head and thoracic integu‑

1993, by H. Suzuki. Paratypes: 8 pupae and 7 pupal

ment densely covered with tubercles (Fig. 40), which

exuviae and 5 mature larvae, all in alcohol, same data as

also separates it from S. (M. ) avilae.

holotype. Other specimens: 3 pupal exuviae, collectedfrom Tenaru River, Guadalcanal Island, 10. VIII. 1992,

5. Simulium (Morops) kawagishii sp. nov.

by H. Suzuki; 3 pupae and 2 pupal exuviae, in alcohol,
collected Tanatita River, Guadalcanal Island, 16. VIII.

DESCRIPTION. Female and Male. Unknown.

1992, by H. Suzuki; I pupal exuvia, in alcohol, collected

Pupa. Body length (excluding gill filaments) ca. 2.5 mm.

at Gold Ridge, Guadalcanal Island, 30. VIII. 1992, by H.

Head. Integument dark yellow, densely covered with

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Figs. 39‑41. Pupal characters of Simulium (Morops) selwynense sp. nov. 39, frons (right half) showing fourtrichomes and dense tubercles; 40, anterior part of thorax (right half) showing a pit‑like organ, fivetrichomes, dense tubercles and basal portion of gill filaments (dorsal view) ; 41, gill filaments
showing surface sculpture near base of each filament (a) . Scale bar 0.02 mm for Fig. 4la.

264

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Figs. 42‑47. Pupal characters of Simulium (Morops) kawagishii sp. nov. and two related species. 42, 44 & 46,

anterior part of thorax showing a pit‑like organ, five trichomes, distribution of tubercles and basal
portion of gill filaments (42, dorsal view; 44 & 46, side view, slide‑mounted) ; 43, gill filaments (sideview) ; 45 & 47, pit‑like organ. 42 & 43, S. (M.) kawagishii; 44 & 45; S. (M.) raunsimnae (paratypeslide‑mounted specimen collected from New Britain, 14. XI. 1957, by J. Smart, BMNH); 46 & 47,
S. (M.) gagiduense (paratype slide‑mounted specimen collected from Morobe District, Papua New
Guinea, 11‑13. I. 1958, by J. Smart, BMNH).

265

round tubercles, and with 4 pairs of trichomes, all longand simple. Antennal sheath with smooth surface.

closer value of plo ratio, 0.2 and 0.26‑0.29, respectively(see Figs. 44 & 46) . However, S. (M.) hawagishii differs

Thorax. Integument dark yellow, densely covered withfrom both species by the thoracic integument densely

round tubercles (Fig. 42); near base of gill there is on

covered with tubercles (almost bare or sparsely covered

each side a small, circular, pit‑like organ (Fig. 42) ; plowith tubercles in the latter species, as seen in Figs. 44 &ratio 0.25; thoracic trichomes 6 pairs (5 anterodorsallyand I posterolaterally) , all long and simple. Gill (Fig.
43) composed of 4 slender filaments, arising from long46) .

(B) papuense‑groupbasal stalk, sessile or short‑stalked, subequal in lengthand thickness (total gill length 1.8‑2.0 mm) ; basal stalk

6. Simulium (Morops) papuense Wharton, 1948

with a transparent pouch, which expands laterally tosome extent; each filament not so thick basally, much

Simulium papuensis Wharton, 1948: 357‑66.

thinner than interspiracular trunk, tapered gradually to
mid point, then nearly of the same diameter up to apicaltip; surface of each filament with well‑defined, annularridges forming polygonal patterns near base, covered

with somewhat larger tubercles on ridges and withsmaller ones on interspaces; all filaments yellow.
Abdomen and Cocoon. As in S. (M.) pohaense.

Simulium papuense: Mackerras and Mackerras, 1949:
372‑405; Mackerras and Mackerras, 1950: 167‑87;
Smart and Clifford, 1965: 591.
Simulium (Morops) papuense: Crosskey, 1967: 44; Cross‑key, 1988: 457; Crosskey, 1989: 225.

This species was originally described from Papua

Mature larva. As in S. (M.) solomonense except follow‑

New Guinea, based on the female, male, pupal anding features: Body length 4.0‑4.5 mm. Body greyish

mature larval specimens (Wharton, 1948). The pupa isyellow. Cephalic apotome pale yellow with a large darkbrown area medially on posterior l/2 with head spots
distinct, often merged in surrounding dark brown areaor cephalic apotome entirely pale yellow with indistinct

distinctive among Morops species by the gill of dendroid
type (i.e., with ca. 30 slender filaments) , and the larva,

by the deep postgenal cleft reaching the posterior borderof the hypostomium. Crosskey (1967) erected theor distinct head spots. Labral fan with ca. 36 main rays.

papuense‑group for this unique species. In our larval

Hypostomal bristles 6 in number per side. Rectal papillaspecimens of S. (M. ) papuense it is also found that theabdominal segments 1‑5 each have a dorsolateral pair ofprotuberances, a character not mentioned in the originalcompound, each of 3 Iobes with 3‑5 finger‑like, secon‑
dary lobules. Posterior circlet with 82‑88 rows of up to
14 hooklets per row.

description of S. (M.) papuense (Wharton, 1948). Thisis the first record of this species from the Guadalcanal

TYPE SPECIMENS. Holotype pupa, slide‑mounted,
collected from Danf River, Guadalcanal Island, Solomon
Islands, 2. IX. 1992, by H. Suzuki. Paratypes: 2 pupae
and 11 mature larvae, same data as holotype.

DISTRIBUTION. Solomon Islands (GuadalcanalIsland.

SPECIMENS EXAMINED. I

reared from pupa, 1

pupa and 7 mature larvae, collected from a slow stream,
Babe, Guadalcanal Island, Solomon Islands, 6, 111. 1992,

by H. Suzuki.

Island) .

REMARKS. The species name kawagishii is given after

DISTRIBUTION. Papua New Guinea and Solomon
Islands (Guadalcanal Island) (new record) .

Mr. N. Kawagishi, ex charg6 d'affaires to Solomon, whokindly helped HS during his stays in the Solomon(O ungrouped species

Islands.

This new species also belongs to the clathrinum‑group by having the pit‑like organ (Fig. 42) , as well asfour gill filaments. The small pit‑like organ (plo ratio0.25) of this species separates it from the three preced‑

ing new species. Among five known species of thisspecies‑group, S. (M.) raunsimnae from New Britain

7. Simulium (Morops) sherwoodi Stone and Maffi,
1971

Simulium sherwoodi Stone and Maffi, 1971: 299‑300.

Simulium (Morops) sherwoodi: Crosskey, 1988: 458;
Crosskey, 1989: 225.

Island and S. (M.) g giduense from Morobe District ofPapua New Guinea (Smart and Clifford, 1965) have aThis species was originally described based on the266

197D. The adult female, male and pupa of S. (M.)

brown; trochanter yellowish white; femur dark yellow toyellowish brown with apical cap brown; tibia white on
basal 3/5, brown to brownish black on distal 2/5; poste‑

sherwoodi are here redescribed, and the mature larva isrior surface of tibia widely white shining when illuminat‑

described for the first time. It is clarified that the female

ed; tarsus brown with basal 3/5 of basitarsus and basal
1/2 of 2nd tarsomere white; basitarsus slender, parallel‑

adult female, male and pupal specimens cellected from
Guadalcanal Island, Solomon Islands, (Stone and Maffi,tarsal claws have a minute basal tooth (not simple asmentioned in the original description). Further, thepupae show a transverse row of spine‑combs dorsally on

sided, ca. 6.9 x as long as wide. Calcipala (Fig. 52) well

the segments 6 to 9 (not segments 5 to 8) and four hooks
ventrally on segments 5 to 7 (not egments 6 to 8) .

developed, ca. 1.2 x as long as wide. Pedisulcus (Fig.
52) well developed. Hind femur and tibia 2.2 x and 1.7
x as wide as hind basitarsus, respectively. All femora

REDESCRIPTION. Female. Body length ca. 2.5 mm.

and tibiae densely covered with scale‑like hairs on outersurface. Claws (Fig. 53) each with a minute basal tooth.

Head. Slightly narrower than thorax. Frons (Fig. 48)
dark brown, shiny, covered with several yellowish whitehairs near lower margin and several dark hairs along
both lateral margins, without scale‑1ike, recumbent
pubescence; frontal ratio 1.6 : 1.0 : 2.1. Frons‑head ratio

1.0:5.6. Fronto‑ocular area (Fig. 49) well developed.

Clypeus dark brown, densely covered with yellowish
white and dark hairs interspersed with whitish yellow,
scale‑like, recumbent pubescence on lower 1/2. Antenna

Wing. Length 1.5 mm. Costa with spinules as well as
hairs. Subcosta haired. Hair tuft on stem vein darkbrown. Basal portion of radius fully haired. Abdomen.
Basal scale brown with fringe of yellowish white hairs.
Dorsal surface of abdomen black, with dark hairs; ter‑gite of 2nd segment whitish pruinose, tergites of seg‑ments 5‑8 shiny. Genitalia. Nearly as in S. (M.) noroen‑se sp. nov. (Fig. 71). Sternal plate undeveloped on
abdominal segment 7. Sternite 8 bare medially, with 12‑composed of 2 + 9 segments, brownish black except

13 dark macrosetae on each side. Anterior gonapo‑

scape, pedicel and base of Ist flagellar segment yellow‑

ish. Maxillary palp with 5 segments; proportional

physis triangular, thin, membraneous, covered denselywith microsetae interspersed with a few setae; innerlengths of 3rd, 4th and 5th segments 1.0 : 1.1:2.4; 3rd

margin well sclerotized. Genital fork of usual inverted‑

segment somewhat enlarged, sensory vesicle (Fig. 50)
elongate, 2.1 x as long as wide, 0.4 x as long as 3rd

Y form, with arms rather broad, folded medially, eachwith a small projection directed forward. Paraproct

segment, with a medium opening slightly distally. Maxil‑
lary lacinia with 9 or 10 inner and 12 or 13 outer teeth.

(Fig. 54) of usual form. Cercus (Fig. 54) ca. 0.5 x as
10ng as wide, with posterior border gently rounded when

Mandible with ca. 25 small inner teeth and lacking outerones. Cibarium (Fig. 51) smooth. Thorax. Scutum black‑ish brown in ground color, whitish grey pruinose, dense‑viewed laterally. Spermatheca (Fig. 55) ovoid, wellsclerotized except tube and tubal base unsclerotized;
internal setae present.

ly covered with whitish yellow or yellow, scale‑like,

Male. Body length ca. 2.5 mm. Head. Wider thanrecumbent pubescence except median large area withthorax. Upper eye consisting of 9 or 10 vertical columns

brown pubescence. Scutellum blackish brown withand 11‑13 horizontal rows of large facets. Clypeus

dark, scale‑like, recumbent pubescence and long, erect,
dark hairs along posterior margin. Postscutellum black‑brownish black, whitish pruinose, covered densely withish brown, whitish grey pruinose (shining in light) , bare.

yellow pubescence, interspersed with dark hairs.
Antenna composed of 2 + 9 segments, yellow with 6

Pleural membrane with ca. 25 yellowish white pubescen‑ce. Katepisternum longer than deep, dark brown, whit‑

apical segments brown; Ist flagellomere somewhat elon‑
gate, ca. 1.6 x as long as 2nd one. Maxillary palp withish grey pruinose (shining in light) , with numerous dark

5 segments; proportional lengths of 3rd, 4th and 5thhairs and yellowish white pubescence; sulcus distinct.

segments 1.0: 1.2 : 2.9; sensory vesicle oblong, 0.33 x as

Legs. Foreleg: coxa and trochanter whitish yellow;
femur dark yellow to pale brown; tibia white with distal

10ng as 3rd segment, with opening near distal end.
Thorax. Nearly as inexcept pleural membrane with

1/4 brownish black, outer surface widely white shiningca. 15 hairs. Legs. Foreleg: coxa and trochanter yellow;

when illuminated; tarsus black; basitarsus somewhat

femur yellow with distal 1/3 brown; tarsus brownish

dilated, ca. 4.7 x as long as its greatest width. Midleg:coxa brown; trochanter whitish yellow; femur yellow;

black; basitarsus somewhat dilated ca. 5.0 X as long as
its greatest width. Midleg: coxa dark brown; trochanter

tibia whitish yellow on basal 2/5, brown on rest; poste‑rior surface of tibia widely white shining when illuminat‑
ed; tarsus brown. Hind leg: coxa yellowish brown or paleand femur yellow; tibia brown with basal 1/4 yellow;
tarsus brown. Hind leg: coxa dark yellow; trochanter
yellow; femur yellow, somewhat darkened toward distal

267

end, with distal cap brown; tibia yellow on basal 1/2,
brown to brownish black on distal 1/2; tarsus brown tobrownish black except basal 3/5 of basitarsus and basal
1/2 of 2nd tarsomere yellow; basitarsus slender, parallel‑

connected in middle. Genitalia. Coxite (Fig. 56) nearlyreqtangular in ventral view, much longer than wide.
Style (Fig. 56) shorter than coxite, tapered towardapical tip, gently curved inward, with apical spine.
Ventral plate (Figs. 57‑59) transverse, ca. 0.4 x as long

sided, ca. 5.6 x as long as wide. Hind femur and tibia
subequal in greatest width, ca. 1.8 x as wide as hindbasitarsus. Calcipala well developed, ca. 1.2 x as longproduced ventrally along posterior margin, with mi‑

as wide. Pedisulcus well developed. All femora andcrosetae on ventral, posterior and dorsal surface; basal

tibiae densely covered with scale‑like hairs on outersurface. Wing. Length 1.5 mm. Other features as inexcept subcosta bare. Abdomen. Basal scale brown, witheral small ones (nearly as in S. (M.) noroense sp. nov.,
Fig. 76). Median sclerite (Fig. 60) weakly sclerotized,

fringe of brown hairs. Dorsal surface of abdominal
segments brownish black, covered with dark hairs; seg‑ments 2, 5‑7 each with a dorsolateral pair of shiny,

whitish pruinose patches; those on segment 2 widelyl¥as wide, with both sides much angulated, widelyarms converging. Paramere with 3 Iong hooks and sev‑

plate‑like, slightly widened toward tip, with round tip.
Pupa. Body length (excluding gill filaments) ca. 2.5 mm.

Head. Integument yellowish brown, moderately covered
with round tubercles; a pair of facial trichomes and 2

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Figs. 48‑60. Adult characters of Simulium (Morops) sherwoodi. 48, frons; 49, fronto‑ocular area; 50, 3rd

maxillary palpal segment; 51, cibarium; 52, calcipala and pedisulcus; 53, claw with a minute tooth;54, paraproct and cercus (side view) ; 55, spermatheca; 56, coxite and style (ventral view); 57‑59,
ventralilate (57, ventral view; 58, end view; 59, side view); 60, median sclerite. 48‑55,; 56‑60, c l.

268

minute tubercles. Abdomen. Terga I and 2 pale yellow‑
ish brown, without tubercles; tergum I with I Iong setaon each side, tergum 2 with 5 simple spinous setae and
1 much longer seta on each side. Terga 3 and 4 eachwith 4 hooked spines directed forward on each side.
Terga 6‑9 each with a transverse row of spine‑combs
and comb‑1ike groups of minute spines directed caudad
on each side. Tergum 9 with a pair of spatulate, termi‑
nal hooks (Fig. 62). Sternum 4 with I bifid hook and 1

pairs of frontal trichomes all long and bifid. Antennal

sheath with smooth surface. Thorax. Integument yel‑lowish brown, bare except anterior surface moderately
covered with round tubercles and posterodorsal surface
moderately covered with small, cone‑shaped tubercles; 5
anterodorsal pairs of trichomes all long and bifid, 1

posterolateral pair long and simple. Pit‑1ike organ
absent. Gill (Fig. 6D composed of paired ventral fila‑
ments and 4 much slender dorsal filaments arising frombasal stalk of paired filaments; paired filaments sub‑simple hook (or spinous seta in sorne pupae) and 2

equal in length (2.5‑3.0 mm) and thickness, Iying horizon‑

slender simple setae on each side; sternum 5 with a pairof bifid or trifid hooks on each side; sterna 6 and 7 eachtally, parallel‑sided or somewhat diverging, directing

forwards and tapered toward apical tip; surface of
paired filaments with many transverse ridges (except
apical portion) forming reticulate patterns, covered
with somewhat larger tubercles on ridges and withsmaller ones on interspaces; 4 slender filaments ar‑with a pair of inner bifid or trifid and outer simple orbifid hooks on each side; Iast segment without grapnel‑
like hooklets ventrolaterally on each side. Cocoon. Wall‑pocket‑shaped, moderately woven, widely extending

ranged in pairs with variable lengths of stalks (usuallyventrolaterally; anterior margin thickly woven; individ‑
ual threads visible; interspaces thinly walled, without

sessile or very short‑stalked, but in one pupa secondary

perf orations.

stalks very long, ca. I mm long) , subequal in length (1.5‑1.8

Mature larva. Body length 3.6‑4.2 mm. Body usually
greyish, coloring on thorax and abdominal segment 3
weak and much paler than others. Cephalic apotomemm) and thickness, with weak annular ridges and fur‑rows on their surface of basal 1/3, and covered with

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66

Figs. 61‑66. Pupal and larval characters of Simulium (Morops) sheruroodi and S. (M.) herei. 61 & 62, pupa; 63‑
66, Iarva. 61, basal portion of gill filaments (side view) ; 62, terminal hooks (end view) ; 63, apex ofmandible; 64 & 66, head capsule (ventral view) showing hypostomium and postgenal cleft; 65,
hypostomium. 61‑64, S. (M.) sherwoodi; 65 & 66, S. (M.) kerei.

269

pale yellow with faint, positive head spots. Antenna
with 3 segments and apical sensillum, Ionger than stemof labral fan; proportional lengths of 3 segments frombase to tip 1.0 : 0.7 : 1.0. Labral fan with ca. 36 main rays.

Mandible (Fig. 63) with comb‑teeth decreasing in size
from Ist to 3rd; mandibular serration composed of 2teeth (1 Iarge and I smalD , without supernumerary
serrations. Hypostomium (Fig. 64) with a row of 9
apical teeth, of which median tooth subequal to orslightly longer than each corner tooth; 6 intermediate
teeth subequal in size to each other, and much smaller
than corner teeth; Iateral margin not serrate; hypos‑
tomal bristles 6 in number per side lying slightly diver‑gent posteriorly from lateral margin. Postgenal cleft(Fig. 64) nearly triangular, gradually narrowed ante‑riorly, ca. 5.0 X as long as postgenal bridge (in somelarvae anterior tip with a narrow extension approaching
to posterior margin of hypostomium) . Thoracic cuticle
bare. Abdominal cuticle bare except each side of analsclerite of last segrnent moderately covered with numer‑
ous colorless setae. Rectal papilla compound, each of 3

REMARKS. This species is characterized in the male by
the presence of the parameral hooks and in the pupa by
the simple, wall‑pocket‑shaped cocoon and the six slen‑
der gill filaments, of which dorsal four filaments aremuch slender and shorter than the ventral paired fila‑
ments, as described by Stone and Maffi (1971). It isnoticeable that the female genitalia are very similar tothat of S. ( G.) hiroshii, described from the Solomon
Islands (Takaoka, 1994), especially in the genital fork
with a projection directed anteriorly on each arm, andthe spermatheca with internal setae. Takaoka (1983)
noted that eight Morops species found in the Philippines
have more similarities to S. (M.) sherwoodi than all theother Morops species reported from Papua New Guinea
and Australia in having the parameral hooks in the male
genitalia, and 6‑filamented gill in the pupa. It is now

clear that S. (M.) sherwoodi and all the Philippine
Morops species are very similar in the male and maturelarva, but the former species is readily separated fromthe latters in the female by the almost bare frons, tarsal

claws with a minute todth, and the spermatheca withlobes with I or 2 finger‑like, secondary lobules oninternal setae, and in the pupa by the frontal integument

posterior surface. Anal sclerite of usual X‑form, withanterior arms ca. 0.8 x as long as posterior ones,

with two pairs of trichomes, the absence of grapnel‑likehooklets and plate‑like, terminal hooks.

broadly sclerotized at base. Ventral papillae present.
Posterior circlet with 82 rows of up to 14 hooklets per

8. Simulram (Morops) noroense sp nov

row. ̲

*, 11 c71, all reared fromDESCRIPTION. Female. Body length ca. 2.2 mm. Head.
Slightly narrower than thorax. Frons (Fig. 67) dark

pupa, 52 pupae, 4 pupal exuviae and 29 mature larvae,

brown, whitish grey pruinose, shiny in certain angles of

all in alcohol, collected from a stream, Selwyn College,

light, moderately covered with yellowish white, scale‑

Guadalcanal Island, 12. VIII. 1993, by H. Suzuki; 4pupae, 3 pupal exuviae and 5 mature larvae, all indark hairs along both lateral margins; frontal ratio 1.7 :SPECIMENS EXAMINED. 9

alcohol, collected from Poha River, Guadalcanal Island,
15. VIII. 1993, by H. Suzuki; 2 cf', both reared from pupa,

12 pupae, I pupal exuvia and 21 mature larvae, all in

alcohol, collected from Tenaru River, GuadalcanalIsland, 10. VIII. 1992, by H. Suzuki; 2, 5 c71, all reared

from pupa, 20 pupae, 2 pupal exuviae and 21 maturelarvae, all in alcohol, collected from Tenaru River,
Guadalcanal Island, 16. VIII. 1992, by H. Suzuki; I pupa
and 18 mature larvae, all in alcohol, collected from Gold

Ridge River, Guadalcanal Island, 30. VIII. 1992, by H.
Suzuki; 5, 4 c?1 , all reared from pupa, 41 pupae, 6 pupal

exuviae and 8 mature larvae, all in alcohol, collectedfrom Tanatita River, Guadalcanal Island, 16. VIII. 1992,
by H. Suzuki.

like, recumbent pubescence, interspersed with a few
1.0:2.6. Frons‑head ratio 1.0 : 5.3. Fronto‑ocular area
(Fig. 68) well developed. Clypeus dark brown, densely

covered with yellowish white, scale‑like, recumbent
pubescence, interspersed with several dark hairs.
Antenna composed of 2 + 9 segments, brownish black
except scape, pedicel and base of Ist flagellar segment
yellowish. Maxillary palp with 5 segments; proportional
lengths of 3rd, 4th and 5th segments 1.0:1.3:2.7; 3rdsegment somewhat enlarged, sensory vesicle (Fig. 69)
elongate, 1.9 x as long as wide, 0.4 x as long as 3rd
segment, with a large opening near distal end. Maxilla‑ry lacinia with 9 inner and 11 or 12 outer teeth.
Mandible with ca. 21 small inner teeth and lacking outerones. Cibarium smooth. Thorax. Scutum blackish
brown in ground color, whitish grey pruinose, densely

DISTRIBUTION. Solomon Islands (Guadalcanalcovered with whitish yellow, recumbent pubescence

Island) .

except median large area with dark brown pubescence;
3 Iongitudinal vittae of whitish yellow, recumbent pubes‑

270cence (1 median vitta very narrow and 2 submedian ones
rather wide) , which are connected anteriorly and poste‑
riorly to areas of the concolored pubescence. Scutellum

brown; tibia white on basal 1/3, brownish black on rest,

blackish brown with brown pubescence and long, erect,
dark hairs along posterior margin. Postscutellum black‑black. Hind leg: coxa yellowish brown; trochanter yel‑
10wish white; femur brown with apical cap dark brown;
tibia white on basal 2/3, brown to brownish black on
rest, though its border not well defined; posterior sur‑face of tibia widely white shining when illuminated;tarsus brown with basal 3/5 of basitarsus and basal 1/2

though its border not well defined; posterior surface oftibia widely white shining when illuminated; tarsus

ish brown, whitish grey pruinose (shining in light) , bare.

Pleural membrane with ca. 25 yellowish white pubescen‑ce. Katepisternum longer than deep, dark brown, whit‑
ish grey pruinose (shining in light) , with numerous darkof 2nd tarsomere white; basitarsus slender, parallel‑
sided, ca. 6.6 x as long as wide. Calcipala well devel‑
oped, ca. 1.3 x as long as wide. Pedisulcus well devel‑
oped. Hind femur and tibia 2.4 x and 1.9 x as wide as

hairs; sulcus distinct. Legs. Foreleg: coxa and trochanter

yellow; femur brown; tibia white with distal 1/4 brown‑

ish black, outer surface widely white shining when il‑luminated; tarsus black; basitarsus dilated, ca. 4.6 >< aslong as its greatest width. Midleg: coxa dark brown;
trochanter yellow; femur brown with apical cap darkhind basitarsus, respectively. All femora and tibiae
densely covered with scale‑like hairs on outer surface.

C! O: OOO
l C!
; " . OCiO) :O) . i ; ;O":i : 0C

/

o)o

! o.

r

Q

O:

o

68

: ir; ';i' fi!

. .fl' ii ""! ;?:i::: :;'?!; i:"'l j' !j,!:{7:"::';},.S{$‑; :; "'a'i? ' ::' ' ":t*^"' l:L:i, "'h'a:1'i:;'l " "'t '" t i"' ''i '*"St'}' t' : ¥x : '"' s:"':'1"'" *!" ;: i

69

:*'*
"¥ **

71

67
70
, ;7 /IP ' l‑V:1̲! )J

.

75

t1 t

.1" I

S
' s

s ;'i

i":' :'.

' ,.:'f.' .t, ,'.ii: '

L

:.
I

l

72

74

. 76

73

Figs. 67‑76. Adult characters of Simulium (Morops) noroense sp. nov. 67, frons; 68, fronto‑ocular area; 69, 3rd

maxillary palpal segment; 70, claw with a minute tooth; 71, genitalia in situ (ventral view) ; 72,
paraproct and cercus (side view) ; 73, calcipala and pedisulcus; 74, coxite and style (ventral view) ;75, ventral plate (ventral view) ; 76, paramere (ventral view) . 67‑72, ; 73‑76, c .

271

Claws (Fig. 70) each with a rudimentary, minute basal

tooth. Wing. Length 1.5 mm. Costa with spinules aswell as hairs. Subcosta haired. Hair tuft on stem veindark brown. Basal portion of radius fully haired.
Abdomen. Basal scale brown with fringe of yellowish
white hairs. Dorsal surface of abdomen black, withdark hairs; tergite of 2nd segment whitish pruinose,
tergites of segments 5‑8 shiny. Genitalia (Figs. 71 &72) . Sternal plate undeveloped on abdominal segment 7.

Sternite 8 bare medially, with 15‑18 dark macrosetae oneach side. Anterior gonapophysis triangular, thin, mem‑braneous, covered densely with microsetae interspersedwith a few setae; inner margin well sclerotized. Genital

fork of usual inverted‑Y form, with arms rather broad,folded medially, each with a distinct projection directed

Abdomen. Basal scale yellow, with fringe of yellow
hairs. Dorsal surface of abdominal segments brownish
black, covered with dark hairs; segments 2, 5‑7 eachwith a dorsolateral pair of shiny, whitish pruinose
patches; those on segment 2 widely connected in middle.
Genitalia. Nearly as in S. (M.) sherwoodi except ventral
plate. Ventral plate (Fig. 75) transverse, 0.45 x as longas wide, not markedly angulate on both lateral margins.

Pupa. As in S. (M.) sherwoodi except following fea‑tures: Head. Integument yellow. Thorax. Integument
yellow, moderately covered with round tubercles except
posterodorsal surface with srnall, cone‑shaped ones.

Abdomen. Terga I and 2 almost transparent. Tergum 6
with no or a few spine‑combs.
Mature larva. As in S. (M.) sherwoodi except following

forward. Paraproct of usual form. Cercus 0.56 x as long

features: Body length 3.8‑4.0 mm. Body uniformly

as wide, with posterior border gently rounded whengreyish. Labral fan with ca. 34 main rays. Rectal papillaviewed laterally. Spermatheca ellipsoidal, well sclero‑
tized except tube and tubal base unsclerotized; internal

compound, each of 3 Iobes with 2 or 3 finger‑like,secondary lobules on posterior surface.

setae present.

Male. Body length ca. 2.5 mm. Head. Wider than thorax.
Upper eye consisting of 9 or 10 vertical columns and 11

horizontal rows of large facets. Clypeus brownish
black, whitish pruinose, covered densely with yellow

pubescence, interspersed with several dark hairs.

TYPE SPECIMENS. Holotype, reared from pupa,

slide‑mounted together with pupal exuvia and cocoon,collected from a stream, Noro, New Georgia Island,Western Province, Solomon Islands, 5. IX. 1992, by H.
Suzuki. Allotype c71 , reared from pupa, slide‑mounted,, 3 c71 , all reared

Antenna composed of 2 + 9 segments, yellow with 4 or 5
apical segments brown; Ist flagellomere somewhat elon‑
gate, ca. 1.5 x as long as 2nd one. Maxillary palp with
5 segments; proportional lengths of 3rd, 4th and 5thsame data as holotype. Paratypes: I

segments 1.0: 1.2 : 3.0; sensory vesicle oblong, 0.27 x as

alcohol, same data as holotype except date: 21. II. 1993.

from pupa, 2 pupae and 3 mature larvae, all in alcohol,same data as holotype; 2, 2 c l , all reared from pupa,

15 pupae, I pupal exuvia and 25 mature larvae, inlong as 3rd segment, with opening somewhat distally (ornear distal end) . Thorax. As in

except pleural

DISTRIBUTION. Solomon Islands (New Georgia

membrane with ca. 20 yellow pubescences. Legs.

Island) .

Foreleg: coxa and trochanter whitish; femur brown;
tibia yellow (though appearing brown due to dense
covering of dark pubescence) with distal 1/3 dark

REMARKS. This species is very similar to S. (M.)

brown, silvery shining in light largely on outer surface;tarsus brownish black; basitarsus somewhat dilated, ca.
5.5 x as long as its greatest width. Midleg: dark brown
to brownish black except trochanter and base of tibia
yellow. Hind leg: coxa brown; trochanter yellow; femur
dark brown; tibia white on basal 1/2, brownish black on
distal 1/2; tarsus brownish black except basal 3/5 ofbasitarsus and basal 1/2 of 2nd tarsomere white; basitar‑sus slender, almost parallel‑sided, ca. 5.3 x as long as
wide. Hind femur and tibia ca. 1.8 x and 1.6 x width ofhind basitarsus, respectively. Calcipala (Fig. 73) welldeveloped, 1.45 x as long as wide. Pedisulcus (Fig. 73)

well developed. All femora and tibiae densely covered
with scale‑like hairs on outer surface. Wing. Length 1.5

mm. Other features as inexcept subcosta bare.

sherwoodi in many features including the genitalia ofboth sexes and the pupal gill filaments. However, S.
(M.) noroense differs from the latter species by the darkcoloration of the femora of both adult sexes, fernale
frons furnished with scale‑like pubescence (Fig. 67),
male ventral plate not markedly angulate laterally (Fig.

75), and pupal thorax moderately covered with tuber‑cles. The tarsal claw of the female S. (M.) noroense hasa minute basal tooth (Fig. 70), which is much smaller
than that of S. (M.) sherwoodi (Fig. 53).

9. Simulium (Morops)kerei Takaoka and Suzuki, 1994

Simulium (Morops) kerez Takaoka and Suzuki 1994
207‑210.

272

This species was recently described based on adult

History Museum, London, U. K. for loaning the typemale and pupal specimens collected from New Georgia
Island, Solomon Islands (Takaoka and Suzuki, 1994).

specimens of several Papua New Guinean and Aus‑

The mature larva is here described.

tralian species of the clathrinum ‑group. HS thanks Dr.

N. Kere, Ministry of Health and Medical Services,
Solomon Islands, for his assistance during the collection

DESCRIPTION. Mature larva. Body length ca. 5.0 mm.
Body greyish black. Cephalic apotome pale yellow with
distinct, positive head spots. Antenna with 3 segments
and apical sensillum, Ionger than stem of labral fan;
proportional lengths of 3 segments from base to tip 1.00 :trips. This work was in part supported by the Grant‑
In‑Aid from the Ministry of Education, Science andCulture of Japan for International Scientific Research

programme (No. 03041080; representative, Prof. M.
Sasa) .

0.96 : 0.81. Labral fan with ca. 45 main rays. Mandible asin S. (M.) sherwoodi. Hypostomium (Fig. 65) with a rowof 9 apical teeth, of which median tooth is subequal in

REFERENCES

size to each corner tooth; 6 intermediate teeth different1 ) Colbo, M. H. (1976): Four new species of Simuliumin length and much shorter than corner teeth; Iateral

Latreille (Diptera: Simuliidae) from Australia, J. Aust.

margin not serrate; hypostomal bristles 7 or 8 in number

Ent. Soc., 15, 253‑269.

per side lying subparallel to lateral margin. Postgenal
cleft (Fig. 66) nearly triangular, gradually narrowed
anteriorly, ca. 4.0 X as long as postgenal bridge. Thora‑
cic cuticle bare. Abdominal cuticle bare except each sideof anal sclerite of last segment moderately covered withnumerous colorless setae. Rectal papilla compound, eachof 3 Iobes with 2 or 3 finger‑like, secondary lobules onposterior surface. Anal sclerite of usual X‑form, withanterior arms ca. 0.9 x as long as posterior ones,
broadly sclerotized at base. Ventral papillae present.
Posterior circlet with ca. 100 rows of up to 18 hooklets

per row.

2 ) Crosskey, R. W. (1967): The classification of Simulium

Latreille (Diptera: Simuliidae) from Australia, New
Guinea and the Western Pacific, J. Nat. Hist. 1, 23‑51.

3 ) Mackerras. M. J. and Mackerras, I. M. (1948):
Simuliidae (Diptera) from Queensland, Aust. J. Scient.
Res.. Ser. B, 1, 231‑270.
4 ) Srnart, J. and Clifford, E. A. (1965): Simuliidae (Dipter‑a) of the territory of Papua and New Guinea, PacificInsects, 7, 505‑619.

5 ) Stone, A. and Maffi, M. (1971): A new: species ofSimulium from Guadalcanal, Solomon Islands (Diptera:
Simuliidae) , J. Med. Entomol., 3, 299‑300.

6 ) Takaoka, H. (1983): The blackflies (Diptera:
Simuliidae) of the Philippines, pp. 212. The Japan Soci‑ety for the Promotion of Science, Tokyo

SPECIMENS EXAMINED. 5 pupal exuviae and 2

7 ) Takaoka, H. (1994): A new blackfly species ofmature larvae, Noro, New Georgia Island, Western

Simulium (Gomphostilbia) from Solomon Islands, South

Province, Solomon Islands, 21. II. 1993, by H. Suzuki.

Pacific (Diptera: Simuliidae) , Jpn. J. Trop. Med. Hyg.,22, 103‑108.

DISTRIBUTION. Solomon Islands (New Georgra
Island) .

ACKNOWLEDGEMENTS
We are grateful to Dr. A. Shelley, Ms C. Lowry andMiss T. Howard, Department of Entomology, Natural

8 ) Takaoka. H. and Davies, D. M. (1995): The black flies(Diptera: Simuliidae) of Java. Indonesia, Bishop
Museum Entomology Monograph, in press
9 ) Takaoka, H. and Suzuki, H. (1994): A new blackfly
species of Simulium (Morops) from Solomon Islands,
South Pacific (Diptera: Simuliidae) , Jpn. J. Trop. Med.
Hyg., 22, 207‑210.

VOL．23NO．4

回eoember1995

CONTENTS
Original

articleNakashima，K．and

Iwanaga，M．

ImmuneCholeraResponse

Componentsofof7伽歪oPatients

ohol6π〜召

againstthe一227−231………………・響Uni，S．，Hayashi，S．，Fukunaga，A．，Takahashi，K，AbdElmaksoud，N．M
Kimata，1．and
Neutralization

Iseki，M．
ofSporozoites

C乃4）孟o司ウoガ4づz6ηzbyRabbitzz67乞s・・233−238

Anti−C郷％廊Serum…

Takaoka，H．
The

Simuliidae（Diptera）from

Papua

New

Island，
一239−252

Guinea……………・

Takaoka，H．andThe

BougainvilleSuzuki，H．

Simuliidae（Diptera）from

the

SolomonIslands，South

Pacific

・・253−272